Species-specific avoidance responses by blue crabs and fish to chronic and episodic hypoxia

Although both chronic and episodic hypoxia (O₂<2 mg l^–1) alter the distribution and abundance patterns of mobile animals within estuaries, recent evidence suggests that some animals may be more likely to remain within hypoxic or anoxic water than others, due to differences in physiological tolerance and movement responses to the dynamics of hypoxia. Determining avoidance responses to hypoxia is important for identifying the species most susceptible to the direct and indirect impacts of these events. A trawl survey was used to examine the avoidance responses of blue crabs (Callinectes sapidus) and several fish [pinfish (Lagodon rhomboides), spot (Leiostomus xanthurus), Atlantic croaker (Micropogonias undulatus), bay anchovy (Anchoa mitchilli), and paralichthid flounders (Paralichthys dentatus and Paralichthys lethostigma)] to chronic hypoxia and episodic hypoxic upwelling events in the Neuse River Estuary, North Carolina, USA. Trawl collections were made in three depth strata (3.0–4.6 m, 1.7–3.0 m, and 0.9–1.7 m depth) to quantify changes in the depth-specific distribution and abundance patterns of the six most common estuarine taxa during three dissolved oxygen conditions: normoxia, chronic hypoxia, and episodic hypoxic upwelling events. Pinfish, anchovies, blue crabs, and paralichthid flounder abundance increased with increasing dissolved oxygen concentrations. The two taxa most closely associated with the bottom (blue crabs and flounder) showed the strongest avoidance response to hypoxia. All taxa showed a stronger avoidance response to chronic hypoxia as compared to episodic hypoxic upwelling events. This difference is attributed to a reduced ability to avoid the rapid intrusions of hypoxic water during episodic events, or to increased risks of injury and predation in shallow refuge habitats, which may force some individuals back into hypoxic water.Communicated by J.P. Grassle, New Brunswick     

Behavioural responses of the Dungeness crab, <Emphasis Type="Italic">Cancer magister</Emphasis>, during feeding and digestion in hypoxic conditions

The Dungeness crab, Cancer magister, inhabits areas that are frequently subject to periods of hypoxia. This species can employ physiological mechanisms that allow it to cope with acute hypoxic episodes. When crabs feed there is a general increase in physiological variables; these may pose an additional physiological burden on crabs already attempting to maintain adequate oxygen uptake in hypoxia. In Barkley Sound, British Columbia, the inshore habitats of C. magister ranged in dissolved oxygen from 28 kPa at the water surface to less than 1.0 kPa just above the sediment–water interface. During short-term hypoxic events, crabs reduced both the amount of food eaten and the amount of time spent feeding. Crabs tended to cease feeding below 3.2 kPa oxygen, but resumed feeding when the dissolved oxygen tensions were rapidly raised to 6 kPa. In a high (10.5–21 kPa) oxygen gradient, both unfed and fed crabs showed no preference for any area of the gradient. In a low (2.5–10.5 kPa) dissolved oxygen gradient, both unfed and fed crabs preferred the highest oxygen regime. In the laboratory, crabs were less likely to enter hypoxic waters (below 3.2 kPa oxygen) to obtain and consume food; those that did moved the food to a higher oxygen regime prior to feeding and settled in higher oxygen regimes for digestion. Crab behaviour was also monitored in the field. Fed and unfed crabs were fitted with ultrasonic telemetry tags and tracked during a tidal cycle. Unfed crabs remained mobile, travelling up to 1,370 m within 6 h, while postprandial crabs settled in areas of high oxygen and moved very little during the first 48 h after release. The present study suggests that C. magister exhibits behavioural responses in order to minimise the use of physiological mechanisms, and maximise foraging and digestive processes. Thus the nutritional state of the individual may be important in regulating both its behaviour and distribution in its natural environment.     

Responses of estuarine crab megalopae to pressure,salinity and light: Implications for flood-tide transport

The megalopal larval stage of many estuarine brachyuran crabs appears to return to adult habitats by undergoing rhythmic vertical migrations which result in saltatory up-estuary transport on flood tides. Larval ascent into the water column during rising tides may be cued by changing hydrologic variables. To test this hypothesis, we investigated the responses of field-caught megalopae of the blue crab Callinectes sapidus and the fiddler crab Uca spp. to constant rates of pressure and salinity change under laboratory conditions. For both genera, pressure changes resulted in increased movement (barokinesis) and upward migration in the test chamber, with C. sapidus megalopae having a lower response threshold (2.8×10^-2 mbar s^-1) than Uca spp. larvae (5×10^-2 mbar s^-1). Similarly, larvae ascended in response to increasing salinity, with C. sapidus larvae being more sensitive. Larvae were negatively phototactic and failed to respond to pressure increases at light levels above 1.0×10¹⁵ and 1.0×10¹³ photons m^-2 s^-1 for C. sapidus and Uca spp. megalopae, respectively. Such responses are thought to explain the low abundances of larvae in the water column during daytime flood tides. Nevertheless, threshold sensitivities to increasing pressure for both genera were above levels experienced during floodtide conditions in the field. Similarly, it is unlikely that increasing salinity is sufficient to induce ascent in Uca spp. postlarvae. However, rates of salinity increase during midflood tide typically reach levels necessary to induce an ascent in C. sapidus megalopae. These results are consistent with the hypothesis that fiddler crab megalopae utilize an endogenous activity rhythm for flood-tide transport, while blue crab megalopae rely upon external cues, especially salinity changes, to time their sojourns in the water column.     

Comparison of the osmoregulatory capabilities of two portunid crabs,Callinectes sapidus and C. similis

An investigation of the osmoregulatory capabilities of two portunid crabs, Callinectes sapidus and C. similis, was conducted to determine if their differences in distributional patterns were reflected in their capacity to adjust physiologically to changes in salinity. After acclimation to 5, 20 and 35 S, measurements of hemolymph and muscle concentrations of Na⁺, Cl^- and K⁺ and muscle-free amino acids indicated that C. sapidus is a better osmoregulator at low salinity than C. similis, while both species osmoregulate equally well at high salinity. This difference in osmoregulatory capacity corresponds well with their distribution in coastal-plain estuaries.This research was supported under agreement (49-7)-5 between the National Marine Fisheries Service and the Energy Research and Development Administration.Communicated by M.R. Tripp, Newark     

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

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

Effects of periodic hypoxia on distribution of demersal fish and crustaceans

Effects of periodic hypoxia (O₂ < 2 mg l^–1) on distribution of three demersal fish species, spot (Leiostomus xanthurus), hogchoker (Trinectes maculatus) and croaker (Micropogonias undulatus), and of two crustacean species, mantis shrimp (Squilla empusa) and blue crab (Callinectes sapidus), were investigated in the lower York River, Chesapeake Bay, USA. Trawl collections were made in four depth strata (5 to 10, 10 to 14, 14 to 20 and > 20 m) during normoxia and hypoxia from 26 June to 20 October 1989. Three periods with hypoxia in the bottom water (below 10 m depth) occurred in mid-July, early August and early September, each with a duration of 6 to 14 d. The demersal fish and crustaceans studied were all affected by hypoxia, and a general migration from deeper to shallower water took place during July and August. However, when oxygen conditions improved after a hypoxic event all species, exceptS. empusa, returned to the deeper areas. The degree of vertical migration was related to levels of oxygen concentration and varied for the different species.M. undulatus was the most sensitive species to low oxygen, followed byL. xanthurus andC. sapidus. T. maculatus andS. empusa were more tolerant and survived in 14 to 25% oxygen saturation by increasing ventilation rate and, forS. empusa, by also increasing blood pigment (haemocyanin) concentration. Periodic hypoxia driven by the spring-neap tidal cycle may represent a natural phenomenon with which the fishes and crustaceans are in a delicate balance. Areas experiencing periodic short-lived hypoxia may be good nursery grounds for fisheries species, and there is no indication that the habitat value in the study area of lower York River is lessened. However, if eutrophication lengthens the time of hypoxia or brings the system closer to anoxia the system may change and become characteristically stressed. The migratory and physiological responses of these species to hypoxia are good indicators of the severity of oxygen stress and could be used as part of an early warning monitoring system for changes in environmental quality.Contribution of the Virginia Institute of Marine Science     

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.     

Circatidal swimming behavior of brachyuran crab zoea larvae: implications for ebb-tide transport

Larvae of the blue crab Callinectes sapidus and fiddler crab Uca pugilator are exported from estuaries and develop on the continental shelf. Previous studies have shown that the zoea-1 larvae of some crab species use selective tidal-stream transport (STST) to migrate from estuaries to coastal areas. The STST behavior of newly hatched larvae is characterized by upward vertical migration during ebb tide followed by a descent toward the bottom during flood. The objectives of the study were (1) to determine if newly hatched zoeae of U. pugilator and C. sapidus possess endogenous tidal rhythms in vertical migration that could underlie STST, (2) to determine if the rhythms persist in the absence of estuarine chemical cues, and (3) to characterize the photoresponses of zoeae to assess the impact of light on swimming behavior and vertical distribution. Ovigerous crabs with late-stage embryos were collected from June to August 2002 and maintained under constant laboratory conditions. Following hatching, swimming activity of zoeae was monitored in darkness for 72 h. U. pugilator zoeae displayed a circatidal rhythm in swimming with peaks in activity occurring near the expected times of ebb currents in the field. Conversely, C. sapidus zoeae exhibited no clear rhythmic migration patterns. When placed in a light field that simulated the underwater angular light distribution, C. sapidus larvae displayed a weak positive phototaxis at the highest light levels tested, while U. pugilator zoeae were unresponsive. Swimming behaviors and photoresponses of both species were not significantly influenced by the presence of chemical cues associated with offshore or estuarine water. These results are consistent with predictions based on species-specific differences in spawning and the proximity of hatching areas to the mouths of estuaries. U. pugilator larvae are released within estuaries near the adult habitat. Thus, ebb-phased STST behavior by zoeae is adaptive since it enhances export. Selective pressures for a tidal migration in C. sapidus larvae are likely weaker than for U. pugilator since ovigerous females migrate seaward prior to spawning and hatching occurs near inlets and in coastal waters.     

Endogenous swimming rhythms in estuarine crab megalopae: implications for flood-tide transport

Up-estuary migration of crab larvae to adult habitats is thought to be accomplished by selective tidal transport in which late-stage larvae enter the water column on flood tides and remain on or near the bottom on ebb tides. This study measured endogenous rhythms in swimming by the last larval stage (megalopa) of blue crabs Callinectes sapidus and fiddler crabs Uca spp. Previous field studies found that megalopae of both species were only abundant in the estuarine water column on nocturnal rising tides. Megalopae were collected from the Newport River Estuary, North Carolina (34°41N; 76°40W) during August–September 1992 and swimming activity was recorded for 4.5 to 7 d under constant conditions with a video system. Rhythms exhibited by both genera in the laboratory were not identical to those recorded in the field. Uca spp. displayed a circatidal rhythm, with maximum swimming occurring near the time of high tide in the field. Rhythm amplitude increased when crushed oyster shells were present, which suggested that megalopae bury or cling to the substrate during quiescent periods. In contrast, C. sapidus had a circadian rhythm in which maximum swimming coincided with the day phase in the field. In most trials, the activity of blue crab megalopae was unrelated to the expected tidal cycle. It was concluded that a tidal rhythm in swimming was the behavioral basis of flood-tide transport for fiddler crab larvae. The endogenous rhythm in blue crabs does not participate in transport, which probably results from behavioral responses to environmental cues associated with flood tide.     

Microbial communities of the carapace, gut, and hemolymph of the Atlantic blue crab, Callinectes sapidus

The Atlantic blue crab Callinectes sapidus is an important fisheries resource that is subject to mortality and morbidity from hemolymph infections. We used culture-independent methods based on the analysis of 16S rRNA genes to characterize and quantify the microflora from the carapace, gut, and hemolymph of C. sapidus with the goals of (1) characterizing the C. sapidus microbial assemblage and (2) identifying the reservoirs of potential pathogens associated with the crab. We found that the carapace, gut and hemolymph microflora have a core Proteobacteria community with contributions from other phyla including Bacteroidetes, Firmicutes, Spirochetes, and Tenericutes. Within this Proteobacteria core, γ-Proteobacteria, including the members of the Vibrionaceae that are closely related to potential pathogens, dominate. Bacteria closely related to hemolymph pathogens were found on the carapace, supporting the hypothesis that punctures, molting damage, or broken dactyls may be routes for hemolymph infections. These results provide some of the first data on the blue crab microbial assemblage obtained with culture-independent techniques and offer insights into the routes of infection and potential bacterial pathogens associated with blue crabs.     

Maternal care in the bromeliad crab Metopaulias depressus (Decapoda) : maintaining oxygen,pH and calcium levels optimal for the larvae

Summary Harsh physico-chemical conditions during early development may select for parental care. However, no study on the evolution of parental care has focused on the physico-chemical conditions in the environment, the physiological needs of early life stages and the significance of parental care. Early development of the bromeliad crab is completed in small rainwater reservoirs in bromeliad leaf axils. Axils not cared for by mother crabs are acidic, hypoxic and contain very little Ca²⁺. Maternal care buffers axil water and increases oxygen and Ca²⁺ availability. Our results show that (a) bromeliad crab larvae die at pH levels usually found in unattended axils, but develop successfully into the first crab stage at the pH typical of cared-for axils; (b) oxygen concentration in unattended axils is below the critical concentration for larvae, but is high enough for normal respiration in cared-for axils; (c) the calcium demand of larvae for moulting and development cannot be satisfied in unattended axils, but is met by the higher calcium content of cared-for axils. Therefore, physicochemical conditions in the bromeliad axil habitat exert strong selection for the maintenance of parental care in the bromeliad crab. Correspondence to: R. Diesel     

Comparative vulnerability to predators,and induced defense responses,of eastern oysters Crassostrea virginica and non-native Crassostrea ariakensis oysters in Chesapeake Bay

Management agencies are considering introducing the Suminoe oyster Crassostrea ariakensis into Chesapeake Bay, USA. It is unknown if the growth of feral populations of this non-native oyster would be regulated by the same predators that once controlled the abundance of the native eastern oyster C. virginica. In laboratory studies, we compared the relative susceptibility of juvenile diploids (shell height < 25 mm) of both oyster species to invertebrate predators of eastern oyster juveniles. Predators included four species of mud crabs [Rhithropanopeus harrisii (carapace width 7–11 mm), Eurypanopeus depressus (6–21 mm), Dyspanopeus sayi (8–20 mm), and Panopeus herbstii (9–29 mm)], the blue crab Callinectes sapidus (35–65 mm), and two sizes of polyclad flatworms (Stylochus ellipticus and possibly Euplana gracilis; planar area ≲5 mm² and ∼14 to 88 mm²). All four species of mud crab and the blue crab preyed significantly (ANOVA, P ≤ 0.05) more on C. ariakensis than on C. virginica, but predation by flatworms of both sizes did not differ significantly between oyster species. The greater susceptibility of C. ariakensis to crab predation was likely due to its shell compression strength being 64% lower than that of C. virginica (P = 0.005). To test for predator-induced enhancement of shell strength, we held oysters of both species for 54 days in the presence of, but protected from, C. sapidus and R. harrisii. Crabs were fed congeneric oysters twice weekly within each aquarium. Compared to controls, shell strength of C. virginica exposed to R. harrisii increased significantly (P < 0.043), as did shell strength of both oyster species exposed to C. sapidus (P < 0.01). Despite the changes in shell strength by both oyster species in the presence of C. sapidus, the shell of C. ariakensis remained 57% weaker than C. virginica. We conclude that, because C. ariakensis exposed to predators continued to have a weaker shell relative to C. virginica, the natural suite of crab and flatworm predators in Chesapeake Bay will likely serve to control the abundance of feral C. ariakensis. We caution that the situation in the natural environment may be sufficiently different in some locations that C. ariakensis may be able to compensate for its greater vulnerability to crab predation and hence become a nuisance species.     

Control of populations of the coral-feeding nudibranch Phestilla sibogae by fish and crustacean predators

Juvenile lesser blue crabs, Callinectes similis Williams, were exposed to a range of salinities for measurement of survival and bioenergetics. Effects of salinity on survival were determined by exposing juvenile crabs to salinity treatments ranging from 0 to 74‰. All crabs survived 21 d of exposure to 5 and 45‰S. The 21 d LC₅₀ values for salinity tolerance (calculated from survival data) were 2.6 and 60.8‰S at low and high salinities, respectively. Energy-budget components and scope for growth were determined for crabs exposed to 2.5, 10, 25, 35 and 50‰S. Energetic absorption rates were highest at 2.5 and 35‰S. Energetic expenditure rates (energy lost to respiration and excretion) were greatest at 2.5‰S, and decreased as salinity increased. Respiration constituted the majority of energetic expenditure at all salinities (92.3% average). Scope for growth was significantly affected by salinity and was highest in crabs exposed to 35‰S. Increased respiration at low salinity may indicate that C. similis incurs greater costs due to osmoregulation. The results of this study indicate that C. similis is capable of surviving and growing in waters with salinities as low as 10‰. Received: 10 January 1997 / Accepted: 11 February 1997     

Some effects of temperature on the growth and metabolic rate of juvenile blue crabs,Callinectes sapidus,in the laboratory

Juvenile blue crabs, Callinectes sapidus Rathbun, were grown in the laboratory at different temperatures, and metabolic-rate determinations were made. Growth is shown to be dependent upon temperature. Crabs kept at high temperatures (34° and 27°C) grow faster than those kept at lower temperatures (13°, 15°, and 20°C). Increase in size per molt is less at higher temperatures than at lower ones. Mortality is directly proportional to temperature between 13° and 34°C and is very high during ecdysis at elevated temperatures. Metabolic rate increases with temperature, but various degrees of acclimation are seen after 4 weeks exposure. No acclimation of general activity to temperature was found. The findings are applied theoretically to crabs living in the region of heated discharge canals of electrical generators: the motile blue crab could extend its growing season without decreasing size at maturity by active selection of thermal surroundings.In part based on a thesis submitted in partial fulfillment of the requirements for the Degree of Master of Science at the University of Florida, USA.     

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.     

Morphological adaptations to chronic hypoxia in deep-sea decapod crustaceans from hydrothermal vents and cold seeps

Animals inhabiting hydrothermal vents and cold seeps face conditions that are challenging for survival. In particular, these two habitats are characterized by chronic hypoxia, sometimes reaching complete anoxia. The characteristics of the scaphognathite and gills were studied in four species of shrimp and three species of crabs from hydrothermal vents and cold seeps, in order to highlight potential adaptations that could enhance oxygen acquisition in comparison with shallow-water relatives. All the vent and seep species studied here exhibit significantly larger scaphognathites, likely allowing more water to flow over their gills per stroke of this appendage. This is probably more energetically efficient that prolonged hyperventilation. In contrast to annelids, vent and seep decapods usually do not possess enlarged gills, a phenomenon likely due to the physical limitations imposed by the size of the gill chamber. In the vent shrimp Rimicaris exoculata and the vent crab Bythograea thermydron, however, there is a significantly higher specific gill surface area linked to a higher number of lamellae per gram of gill. Again in contrast to annelids, the diffusion distance through the gills is not strikingly different between the vent shrimp Alvinocaris komaii and the shallow-water species Palaemon spp. This may indicate that the epithelium and cuticle of the decapod gills are already optimized for oxygen uptake and that reducing the thickness of these compartments is not physically possible without affecting the physical integrity of the gills.     

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.     

Accumulation and release of petroleum-derived aromatic hydrocarbons by four species of marine animals

Survival of Ctenodiscus crispatus during exposure to hypoxia (P _{O 2}<3 mm Hg) at 5°C is greater than that of any echinoderm reported in the literature, the LT₅₀ being 248 h; this is reduced to 236 h in the presence of hydrogen sulfide. Unlike Asterias vulgaris and A. forbesi, both of which lose the tube foot response to tactile stimulation long before death from hypoxia occurs, C. crispatus remains responsive until death. The extension of the highly protrusible epiproctal cone, which occurs in 75% of the mud stars simultaneously exposed to hypoxia and H₂S, serves to maintain burrow contact with the overlying water. The rate of oxygen consumption remains constant down to an ambient oxygen partial pressure of 10 to 25 mm Hg, becoming more oxygen-dependent after prior exposure of the asteroids to hypoxia. C. crispatus exhibits a clear oxygen-debt phenomenon as well as a compensatory reduction in the residual P _{O 2} (oxygen partial pressure at which oxygen consumption ceases) from 2.4 to 0.2 mm Hg after hypoxic exposure.     

Synergistic effects of cadmium and salinity combined with constant and cycling temperatures on the larval development of two estuarine crab species

The developmental stages from megalopa to third crab of the blue crab Callinectes sapidus Rathbun were tested in 12 combinations of cadmium (0, 50, and 150 ppb) and salinity (10, 20, 30, and 40) at 25°C. A reduction in survival and a significant delay in development from megalopa to third crab occurred within each salinity regime in 50 ppb compared with the control. Comparison of the delay in development within each salinity regime revealed that the sublethal effect of cadmium was most pronounced in the salinities normally preferred by C. sapidus. A similar comparison within each cadmium concentration, however, showed that the developmental time from megalopa to third crab was approximately the same irrespective of salinity. The developmental stages from hatch to first crab of the mud-crab Rhithropanopeus harrisii (Gould) were examined in 63 combinations of cadmium (0, 50, and 150 ppb), salinity (10, 20, and 30), constant temperature (20°, 25°, 30°, and 35°C) and cycling temperature (20° to 25°C, 25° to 30°C, and 30° to 35°C). The results indicated that cycling temperatures may have a stimulating effect on survival of the larvae compared to constant temperatures, both in the presence and in the absence of cadmium. Effects of cadmium and salinity and their interaction on the survival of the larvae from zoeae to megalopa were documented at most of the temperatures by analyses of variance. The zoeal larvae were more susceptible to cadmium than the megalopa. Effects of different combinations of cadmium and salinity on the duration of larval development were assessed by a t-test.     

Effects of sulfide exposure history and hemolymph thiosulfate on oxygen-consumption rates and regulation in the hydrothermal vent crab Bythograea thermydron

The hydrothermal vent crab Bythograea thermydron is exposed to high environmental concentrations of sulfide and low levels of oxygen for extended periods of time. It has previously been shown that hydrogen sulfide is oxidized to the relatively non-toxic thiosulfate (S₂O ₃ ^2– ), which accumulates in the hemolymph. Hemolymph thiosulfate levels in freshly captured crabs vary significantly among crabs from different hydrothermal vent sites as well as between crabs from different microhabitats within the same site. Hemolymph thiosulfate concentrations were not significantly different between crabs captured at the same site 6 mo apart. Hemolymph thiosulfate concentrations ranged from 66 mol 1^–1 in a crab captured at a site with relatively low sulfide venting, to 3206 mol 1^–1 in an individual that was netted from an active smoker vent with much higher sulfide exposure. The differences in hemolymph thiosulfate between sites and the stability of hemolymph thiosulfate in crabs captured at the same site at different times suggest that sulfide exposure is significantly different between sites and that this exposure may not vary significantly over the course of a few months. B. thermydron experimentally exposed to sulfide had high levels of thiosulfate in their hemolymph and increased abilities to regulate oxygen consumption in conditions of low oxygen. This enhancement of regulatory abilities suggests that the previously demonstrated increased hemocyaninoxygen (Hc–O₂) affinity due to elevated thiosulfate may be adaptive in vivo. Average oxygen-consumption rates were much higher in crabs experimentally exposed to sulfide than in unexposed crabs. Crabs injected with isosmotic thiosulfate did not have increased oxygen-consumption rates as did the sulfide-exposed individuals, but did show a similar reduction in P _c (the critical partial pressure of oxygen at which crabs can no longer regulate oxygen consumption). This suggests that it is the sulfide exposure and/or detoxification rather than the elimination of thiosulfate that causes the increase in metabolic rate. Thiosulfate diffuses into dead crabs and into live crabs exposed to 15 mmol S₂O ₃ ^2- l^–1, indicating substantial permeability, and yet live crabs are able to eliminate thiosulfate when incubated in sea water containing 1.5 mmol S₂O ₃ ^2- l^–1, suggesting a process that has an active component.