Interactive effects of food deprivation state and hypoxia on the respiratory responses of postprandial rock crabs, Cancer irroratus.

Under the background of climate change, increasing attention has focused on the effects of ocean deoxygenation on marine organisms. However, few studies address the effects of different food deprivation states on hypoxia tolerance. We therefore investigated the metabolic responses of the Atlantic rock crab, Cancer irroratus (starved 28-35 days, fasted 3-5 days and recently fed). Starved-crab exhibited the lowest critical oxygen saturation (Scrit), while fed-crab had the highest Scrit. The fed-crab maintained an elevated postprandial oxygen consumption (MO2) even below the Scrit of fasted-crab indicating reserved aerobic scopes for critical activities in severe hypoxia. Following feeding, hypoxia (50% and 20% oxygen saturation, SO2) retarded the specific dynamic action resulting in lower peak MO2 and longer duration. The starved-crab exhibited a lower peak MO2, prolonged duration and higher energy expenditure than fasted-crab after feeding. The decline in arterial PO2 was most pronounced below the Scrit for both fasted- and starved-crab. The higher hemocyanin concentration ([Hc]) of fasted-crab (than starved-crab) suggested they had improved oxygen transport capacity, but hypoxia did not increase [Hc] during the 72-h experiment. Following feeding, the fasted-crab significantly increased L-lactate concentration ([L-lactate]) in 20% SO2, which was not observed in starved-crab. These results suggest starvation may trigger a cross-tolerance to hypoxia. Because crabs can undergo long periods of food deprivation in their natural environment, future studies should consider how this may affect their ability to deal with environmental perturbations.

Effect of Animal Stocking Density and Habitat Enrichment on Survival and Vitality of Wild Green Shore Crabs, Carcinus maenas, Maintained in the Laboratory.

The wide geographic distribution, large size and ease of capture has led to decapod crustaceans being used extensively in laboratory experiments. Recently in the United Kingdom decapod crustaceans were listed as sentient beings, resulting in their inclusion in animal care protocols. Ironically, little is known about how captive conditions affect the survival and general condition of wild decapod crustaceans. We used the green shore crab, Carcinus maenas, to investigate the effects of stocking density and shelter on survival and vitality indices during a 6 month period in the laboratory. Neither stocking density nor the presence of shelter affected survival. Stocking density also had no effect on the vitality indices (limb loss, claw strength, BRIX, righting time, leg flare and retraction). The presence of shelter did affect the number of limbs lost and the leg retraction response, but had no effect on the other vitality indices. All vitality indices changed, and mortality increased over time, independent of treatment: this became most apparent after 8 to 11 weeks storage in the laboratory. This decline in condition may have been due to repeated handling of the crabs, rather than the stocking conditions. In support of this, untracked, non-handled (control) individuals sustained a 4% mortality rate compared with 67% mortality in experimental crabs during the 6 month period. Although simple experimental monitoring of crabs with biweekly vitality tests only produced transient short-term stress events, the repeated handling over time apparently led to a cumulative stress and a deterioration in animal health. Bringing wild crustaceans into the laboratory and holding them, even with modest experimental manipulation, may result in high mortality rates. Researchers and animal care committees need to be aware that wild captive invertebrates will respond very differently to laboratory-bred vertebrates, and plan experiments accordingly.

Patterns of heart rate and cardiac pausing in unrestrained resting decapod crustaceans.

Measurement of heart rate (HR) has been used as an important physiological indicator in a broad range of taxa. In the present study HR patterns were measured in five species of unrestrained, resting decapod crustaceans. In addition to variation in HR among individuals, it was also very variable within an individual animal. While some of this variation was related to activity, there was also a non-locomotory component. Unstressed, resting crabs exhibited intermittent heart activity, whereas HR in stressed crabs was more stable, suggesting differential control of HR in resting crabs. Once the animals settled in the experimental apparatus they exhibited regular and extended cardiac pauses (acardia) of 15-300-s duration. As with HR, there was a significant variation in the frequency and length of acardic events, which were only observed in inactive crabs. Regaining of HR, following a period of acardia, was characterized by small adjustments in position and movement of the mouthparts. This rhythmic pattern, and the fact that entry into and out of acardia was not instantaneous, suggested that these events were related to release of neurohormones and their subsequent degradation in the system, rather than direct neural control of the heart. Because HR was variable and interrupted by regular periods of acardia, caution is recommended when calculating baseline levels of HR, or using HR alone as an indicator of physiological stress. Incorporating a coefficient of variation for HR and/or measuring the periods of acardia may be a more reliable indicator of physiological stress in decapod crustaceans.

Acclimation to tidal conditions alters the physiological responses of the green shore crab, Carcinus maenas, to subsequent emersion.

Animals inhabiting the intertidal zone are exposed to abrupt changes in environmental conditions associated with the rise and fall of the tide. For convenience, the majority of laboratory studies on intertidal organisms have acclimated individuals to permanently submerged conditions in seawater tanks. In this study, green shore crabs, Carcinus maenas, were acclimated to either a simulated tidal regime of continuous emersion-immersion ('tidal') or to permanently submerged conditions ('non-tidal') to assess their physiological responses to subsequent emersion. Tidal crabs exhibited an endogenous rhythm of oxygen consumption during continuous submersion with lower oxygen consumption during periods of anticipated emersion, which was not detected in non-tidal crabs. During emersion, tidal crabs were able to buffer apparent changes in acid-base balance and exhibited no change in venous pH, whereas non-tidal crabs developed an acidosis associated with a rise in lactate levels. These results indicate that tidal crabs were better able to sustain aerobic metabolism and had lower metabolic costs during emersion than non-tidal crabs. It is likely that the elevated levels of haemocyanin exhibited by tidal crabs allowed them to maintain oxygen transport and buffer pH changes during emersion. This suggests that acclimation of C. maenas to submerged conditions results in a loss of important physiological mechanisms that enable it to tolerate emersion. The results of this study show that caution must be taken when acclimating intertidal organisms to submerged conditions in the laboratory, as it may abolish important physiological responses and adaptations that are critical to their performance when exposed to air.

Changes in precipitation may alter food preference in an ecosystem engineer, the black land crab, Gecarcinus ruricola.

Gecarcinid land crabs are ecosystem engineers playing an important role in nutrient recycling and seedling propagation in coastal forests. Given a predicted future decline in precipitation for the Caribbean, the effects of dehydration on feeding preferences of the black land crab Gecarcinus ruricola were investigated. G. ruricola were offered novel food items of lettuce, apple, or herring to test for food choice based on water and nutritional (energetic) content in single and multiple choice experimental designs. The effect of dehydration was incorporated by depriving crabs of water for 0, 4, or 8 days, leading to an average body water loss of 0%, 9%, and 17%, respectively, (crabs survived a body water loss of 23% + 2% and 14-16 days without access to water). The results were consistent between the single and multiple choice experiments: crabs consumed relatively more apple and fish and only small amounts of lettuce. Overall, no selective preferences were observed as a function of dehydration, but crabs did consume less dry food when deprived of water and an overall lower food intake with increasing dehydration levels occurred. The decrease in feeding was likely due to loss of water from the gut resulting in the inability to produce ample digestive juices. Future climatic predictions suggest a 25-50% decline in rainfall in the Caribbean, which may lead to a lower food intake by the crabs, resulting in compromised growth. The subsequent reduction in nutrient recycling highlights possible long-term effects on coastal ecosystems and highlights the importance of future work on climate relative behavioral interactions that influence ecosystem function.

Application of Miniature Heart Rate Data Loggers for Use in Large Free-Moving Decapod Crustaceans: Method Development and Validation.

Cardiovascular responses of decapod crustaceans to environmental challenges have received extensive attention. However, nearly all of these studies have been restricted to lab-based experiments; here we describe a methodology that will enable measurement of heart rate (HR) in free-moving decapods in the field. Data storage tag heart rate and temperature loggers (DST micro-HRT; Star-Oddi) were used to record electrocardiograms (ECG) and HR in large decapod crustaceans. These loggers were originally designed for use in vertebrates and must be surgically implanted in the body cavity near the heart in order to function. We adapted these loggers for external use in large decapod crustaceans. The method involved abrading the carapace directly above the heart and placing the electrodes of the logger directly on top of the dermal tissue. The logger was then secured in place with periphery wax. This method negated some of the more intricate operations used for vertebrates. The rapid setup time of approximately 5 min suggested that animals could be easily instrumented in the field and without the use of anesthetic. The logger was calibrated by simultaneously measuring the HR changes of a West Indian spider crab Mithrax spinosissimus with a pulsed-Doppler flowmeter. The data gathered with the two methods showed a tight correlation during an increase in temperature. The loggers were also successfully implanted in a variety of other large species of aquatic and terrestrial decapods. The data obtained showed that the method works in a broad range of species, under different experimental conditions. In each case, the loggers comprised less than 1% of the body mass and would be suitable for use in animals >300 g. All animals survived the attachment procedures and were feeding and active after removal of the loggers. Nearly all previous cardiac measurements on decapods have been carried out in controlled laboratory settings. The use of these loggers will make significant advances in measuring HR in unrestrained, undisturbed animals in their natural environment during extended periods of time and has the potential to lead to novel findings.

Metabolic costs of the mechanical components of the apparent specific dynamic action in the Dungeness crab, Cancer magister.

The specific dynamic action (SDA) describes the postprandial increase in metabolism. It is a composite of mechanical and chemical digestion, nutrient transport and protein synthesis. How these individual events contribute to the overall SDA has not been worked out fully for any organism. The mechanical events associated with the SDA were investigated in Dungeness crabs, Cancer magister. Following consumption of a meal, oxygen consumption (MO2) remained elevated for several hours. When the crabs were presented with fish scent there was a 2 fold increase in MO2, which rapidly decreased once the stimulus was removed. Crabs were then offered fish in a perforated tube. There was a rapid increase in MO2 associated with handling which returned to pre-treatment levels within an hour of removal of the tube. Finally the crabs were fed a piece of foam that had been soaked in fish water to determine the costs of mechanical digestion. The mechanical breakdown of the meal accounted for 29.9±3.3% of the overall SDA. Since food handling produced a large increase in MO2, it was reinvestigated using crabs that had one or both claws removed. Although there were no statistically significant differences as a function of claw removal there was a consistent trend in the data. The maximum MO2, scope, duration and SDA increased from animals with 0 claws through 1 claw to 2 claws. The results showed that the mechanical portion of the SDA can account for a significant portion of the overall budget in decapod crustaceans.

Behavioral Thermoregulation and Trade-Offs in Juvenile Lobster Homarus americanus.

Water temperature influences the behavior and distribution patterns of both larval and adult American lobster Homarus americanus. However, very little is known about the responses of juvenile lobsters. The juvenile life stage is a critical period; high levels of mortality, combined with specific behavioral responses, can disconnect larval settlement from patterns of abundance of adults. We assessed behavioral thermoregulation in juvenile lobsters, and determined how thermal preferences can be altered by the presence of shelter and food. Juvenile lobsters avoided temperatures higher than 20 °C and lower than 8 °C, and had a mean temperature preference of 16.2 ± 1 °C. This preference was unaffected by prior acclimation, origin (laboratory-raised or wild), or size. When the animals were subjected to a temperature change (5-20 °C), activity rates peaked at 15 °C, and remained stable thereafter. Activity rates did not change when a shelter was added. The addition of food resulted in an increase in activity associated with food handling. When juvenile lobsters were offered a choice between temperature, shelter, and food, they always chose the environment with a shelter, even when it was in a thermally unfavorable temperature. Juveniles also spent more time in a thermally unfavorable environment when food was present; however, acquisition of a shelter was prioritized over food. Although juveniles had a similar thermal preference to adults, they are more vulnerable to predation; the innate shelter-seeking behavior of juveniles overrode their thermal preference. While temperature is an important environmental factor affecting the physiology, distribution, and growth of aquatic ectotherms, our findings suggest that trade-off behaviors occur in order to maintain optimal fitness and survival of the individual.

Physiological responses to digestion in low salinity in the crabs Carcinus maenas and Cancer irroratus.

Osmoregulation and digestion are energetically demanding, and crabs that move into low salinity environments to feed must be able to balance the demands of both processes. Achieving this balance may pose greater challenges for weak than for efficient osmoregulators. This study examined the rate of oxygen consumption (MO2) of Carcinus maenas (efficient osmoregulator) and Cancer irroratus (weak osmoregulator) as a function of feeding and hyposaline stress. The MO2 increased 2-fold in both species following feeding. The MO2 increased and remained elevated in fasted crabs during acute hyposaline exposure. When hyposaline stress occurred after feeding, C. maenas responded with an immediate summation of the MO2 associated with feeding and hyposaline stress, whereas C. irroratus reacted with a partial summation of responses in a salinity of 24‰, but were unable to sum responses in 16‰. C. irroratus exhibited longer gut transit times. This may be due to an inability to regulate osmotic water onload as efficiently as C. maenas. Mechanical digestion in crabs can account for a significant portion of SDA, and a short term interruption led to the delay in summation of metabolic demands. Although protein synthesis is reported to account for the majority of SDA, this did not appear to be the case here. Protein synthesis rates were higher in C. irroratus but neither feeding or salinity affected protein synthesis rates of either species which suggests that protein synthesis can continue in low salinity as long as substrates are available.

Effect of meal type on specific dynamic action in the green shore crab, Carcinus maenas.

The effect of meal type on specific dynamic action was investigated in the green shore crab, Carcinus maenas. When the crabs were offered a meal of fish, shrimp, or mussel of 3 % of their body mass the duration of the SDA response and thus the resultant SDA was lower for the mussel, compared with the shrimp or fish meals. In feeding behaviour experiments the crabs consumed almost twice as much mussel compared with fish or shrimp. When the animals were allowed to feed on each meal until satiated, the differences in the SDA response were abolished. The mussel was much softer (compression test) than the fish or shrimp meal, and meal texture is known to affect the SDA response in amphibians and reptiles. When the crabs were offered a meal of homogenized fish muscle or whole fish muscle, the SDA for the homogenized meal was approximately 35 % lower. This suggested that a significant portion of the SDA budget in decapod crustaceans may be related to mechanical digestion. This is not unexpected since the foregut is supplied by over forty muscles which control the cutting and grinding movements of the gastric mill apparatus. There were slight, but significant differences in protein, lipid, moisture and total energy content of each meal type. Three prepared meals that were high in either protein, lipid or carbohydrate were offered to the crabs to determine if the nutrient content was also a contributing factor to the observed differences in the SDA. The crabs did not eat the prepared meals as readily as the natural food items and as they are messy feeders there was a large variation in the amount of food eaten. The lack of significant differences in the SDA response as a function of nutrient content was likely due to differences in amount of food eaten, which is a major factor determining the SDA response. The differences in SDA when consuming natural food items were likely due to a combination of the costs of mechanical digestion, variation in nutrient content and food preference: determining how each of these factors contributes to the overall SDA budget remains a pressing question for comparative physiologists.

Extracellular digestion during hyposaline exposure in the Dungeness crab, Cancer magister, and the blue crab, Callinectes sapidus.

Extracellular digestive processes were examined in the Dungeness crab, Cancer magister and the blue crab, Callinectes sapidus, during hyposaline exposure. Both species are found in estuaries as adults, but vary in their ability to balance the cardiovascular and respiratory demands of concurrent osmoregulation and digestion. The weak osmoregulator, C. magister, is unable to balance the demands of osmoregulation and digestion. Concordant with observed decreases in oxygen consumption and mechanical digestion, proteolytic digestion within the foregut and hepatopancreas was delayed, resulting in a relative reduction of circulating amino acids post-feeding in low salinity. In contrast, the efficient osmoregulator, C. sapidus, balances the demands of osmoregulation and digestion, and mechanical digestion continues unabated in low salinity. Protease activity in the gut fluid and hepatopancreas showed either no change or a reduction over time. The transport of amino acids into the cells post-feeding is opposed by an efflux of amino acids at the cellular level, and resulted in a build up of amino acids in the hemolymph. Despite differences in the extracellular responses to low salinity exposure following feeding, both species were able to maintain high digestive efficiencies.

Effect of meal size and body size on specific dynamic action and gastric processing in decapod crustaceans.

Meal size and animal size are important factors affecting the characteristics of the specific dynamic action (SDA) response across a variety of taxa. The effects of these two variables on the SDA of decapod crustaceans are based on just a couple of articles, and are not wholly consistent with the responses reported for other aquatic ectotherms. Therefore, the effects of meal size and animal size on the characteristics of SDA response were investigated in a variety of decapod crustaceans from different families. A 6 fold increase in meal size (0.5%-3% body mass) resulted a pronounced increase in the duration of increased oxygen consumption, resulting in an increase in the SDA of Callinectes sapidus, Cancer gracilis, Hemigrapsus nudus, Homarus americanus, Pugettia producta and Procambarus clarkii. Unlike many other aquatic ectotherms a substantial increase between meal sizes was required, with meal size close to their upper feeding limit (3% body mass), before changes were evident. In many organisms increases in both duration and scope contribute to the overall SDA, here changes in scope as a function of meal size were weak, suggesting that a similar amount of energy is required to upregulate gastric processes, regardless of meal size. The SDA characteristics were less likely to be influenced by the size of the animal, and there was no difference in the SDA (kJ) as a function of size in H. americanus or Cancer irroratus when analysed as mass specific values. In several fish species characteristics of the SDA response are more closely related to the transit times of food, rather than the size of a meal. To determine if a similar trend occurred in crustaceans, the transit rates of different sized meals were followed through the digestive system using a fluoroscope. Although there was a trend towards larger meals taking longer to pass through the gut, this was only statistically significant for P. clarkii. There were some changes in transit times as a function of animal size. The foregut clearance times for Cancer magister increased with increasing body size, while smaller Carcinus maenas cleared the hindgut region at a faster rate than larger individuals. Unlike fish there was no clear relationship between transit rates and any of the SDA characteristics. While the fluoroscopy method is useful for assessing foregut activity and food passage, it is limited when inferring connections between nutrient assimilation and post-absorptive processes in crustaceans. Therefore, at least with respect to meal size, transit rates do not make a good proxy for determining the SDA characteristics in crustaceans.

A review of gastric processing in decapod crustaceans.

This article reviews the mechanical processes associated with digestion in decapod crustaceans. The decapod crustacean gut is essentially an internal tube that is divided into three functional areas, the foregut, midgut, and hindgut. The foregut houses the gastric mill apparatus which functions in mastication (cutting and grinding) of the ingested food. The processed food passes into the pyloric region of the foregut which controls movement of digesta into the midgut region and hepatopancreas where intracellular digestion takes place. The movements of the foregut muscles and gastric mill are controlled via nerves from the stomatogastric ganglion. Contraction rates of the gastric mill and foregut muscles can be influenced by environmental factors such as salinity, temperature, and oxygen levels. Gut contraction rates depend on the magnitude of the environmental perturbation and the physiological ability of each species. The subsequent transit of the digesta from the foregut into the midgut and through the hindgut has been followed in a wide variety of crustaceans. Transit rates are commonly used as a measure of food processing rates and are keys in understanding strategies of adaptation to trophic conditions. Transit times vary from as little as 30 min in small copepods to over 150 h in larger lobsters. Transit times can be influenced by the size and the type of the meal, the size and activity level of an animal and changes in environmental temperature, salinity and oxygen tension. Ultimately, changes in transit times influence digestive efficiency (the amount of nutrients absorbed across the gut wall). Digestive efficiencies tend to be high for carnivorous crustaceans, but somewhat lower for those that consume plant material. A slowing of the transit rate allows more time for nutrient absorption but this may be confounded by changes in the environment, which may reduce the energy available for active transport processes. Given the large number of articles already published on the stomatogastric ganglion and its control mechanisms, this area will continue to be of interest to scientists. There is also a push towards studying animals in a more natural environment or even in the field and investigation of the energetic costs of the components of digestion under varying biotic and environmental conditions will undoubtedly be an area that expands in the future.

Specific dynamic action in the sunflower star, Pycnopodia helianthoides.

The effects of meal size and meal type on specific dynamic action (SDA) were investigated in a large, active asteroid, the sunflower star, Pycnopodia helianthoides. When the sunflower stars were fed clam flesh totalling 5%, 10%, or 20% of their body weight there was a step-wise increase in the scope, time to peak oxygen consumption, duration of the response and total SDA. The change in the rate of oxygen consumption was slower than other organisms, and oxygen uptake remained elevated for over 12d following consumption of the largest meal. There were also differences in the characteristics of the SDA if sunflower stars consumed a whole clam versus the shucked flesh of a clam. The time to reach peak oxygen consumption was greater for sunflower stars consuming a whole clam. This occurred because the clam had to be opened before they could digest the flesh; a smaller initial peak comprising 3.5% of the total SDA represented the energy require to open the clam valves. When the sunflower stars were fed different prey items (e.g. butter clam, purple urchin and herring) of similar wet organic mass, there was no difference in the time to peak, peak oxygen uptake or total SDA despite the fact that the prey items differed in protein, lipid and caloric content. There was an increased duration for which oxygen uptake remained elevated for sea stars that consumed the urchin meal. Five of the seven sunflower stars that consumed urchins exhibited a smaller second peak in oxygen uptake, totalling approximately 8.5% of the SDA energy budget. This likely represented the energy required to eject the urchin test from the stomach. Although the sunflower star is much larger and more active than other sea stars, it displayed similar SDA responses to other members of the Asteroidea, indicative of the low metabolic rate of this class.

Assimilation of water and dietary ions by the gastrointestinal tract during digestion in seawater-acclimated rainbow trout.

Recent studies focusing on the consequences of feeding for ion and water balance in freshwater fish have revealed the need for similar comparative studies in seawater fish. A detailed time course sampling of gastrointestinal (GI) tract contents following the ingestion of a single meal of a commercial diet revealed the assimilation of both water and dietary ions (Na(+), Cl(-), K(+), Ca(2+), Mg(2+)) along the GI tract of seawater-acclimated rainbow trout (Oncorhynchus mykiss) which had been fasted for 1 week. Consumption of the meal did not change the drinking rate. There was a large secretion of fluid into the anterior intestine and caecae (presumably bile and/or pancreatic secretions). As a result, net assimilation (63%) of the ingested water along the GI tract was lower than generally reported for fasted trout. Mg(2+) was neither secreted into nor absorbed from the GI tract on a net basis. Only K(+) (93% assimilated) and Ca(2+) (43% assimilated) were absorbed in amounts in excess of those provided by ingested seawater, suggesting that dietary sources of K(+) and Ca(2+) may be important to seawater teleosts. The oesophagus-stomach served as a major site of absorption for Na(+), Cl(-), K(+), Ca(2+), and Mg(2+), and the anterior intestine and caecae as a major site of net secretion for all of these ions, except Cl(-). Despite large absorptive fluxes of these ions, the ionic composition of the plasma was maintained during the digestion of the meal. The results of the present study were compared with previous work on freshwater-acclimated rainbow trout, highlighting some important differences, but also several similarities on the assimilation of water and ions along the gastrointestinal tract during digestion. This study highlights the complicated array of ion and water transport that occurs in the intestine during digestion while revealing the importance of dietary K(+) and Ca(2+) to seawater-acclimated rainbow trout. Additionally, this study reveals that digestion in seawater-acclimated rainbow trout appears to compromise intestinal water absorption.

Cardiovascular system of the Majidae (Crustacea: Decapoda).

The cardiovascular system of majid crabs was mapped using corrosion casting techniques. The general form of the circulatory system was comparable to that of other malacostracan crustaceans, but with distinct differences between several arterial systems. The anterior aorta exited from the anterior surface of the heart supplying hemolymph to the antennae, eyestalks, gastric muscles and brain. This artery was more complex compared with other decapods. The anterolateral arteries exited from the anterior dorsal surface of the heart and supplied hemolymph to the hypodermis, stomach, antennal gland and mandibular muscles. The hepatic arteries were larger and more complex compared with other decapod families, branching profusely within the hepatopancreas and gonads. The small posterior aorta exited from the posterior-ventral surface of the heart. Standard sex-specific differences in this artery were observed. Exiting from the ventral surface of the heart, the sternal artery supplied each pereiopod in a segmental arrangement. The sternal artery arrangement was different to other brachyuran crabs, possibly a symplesiomorphy with segmented ancestors. In accordance with anatomical descriptions of blue crabs and Cancer crabs it would also seem appropriate to classify the circulatory system of the Majidae as one that is "incompletely closed".

Respiratory and digestive responses of postprandial Dungeness crabs, Cancer magister, and blue crabs, Callinectes sapidus, during hyposaline exposure.

Respiratory responses and gastric processing were examined during hyposaline exposure in two crab species of differing osmoregulatory ability. The efficient osmoregulator, Callinectes sapidus, displayed an immediate increase in oxygen uptake when exposed to low salinity in isolation. In contrast, the weak osmoregulator, Cancer magister, showed no change in oxygen uptake upon acute exposure (<6 h), but slight increases in oxygen uptake tended to occur over longer time scales (12-24 h). These changes were likely attributable to an increase in avoidance activity after 6 h hyposaline exposure. Following feeding in 100% SW, oxygen uptake doubled for both species and remained elevated for 15 h. When postprandial crabs were exposed to low salinities, C. sapidus were able to sum the demands of osmoregulation and digestion. Thus, gastric processes continued unabated in low salinity. Conversely, postprandial C. magister prioritized responses to low salinity over those of digestion, resulting in a decrease in oxygen uptake when exposed to low salinity. This decrease in oxygen uptake corresponded to a reduction in the rate of contraction of the pyloric stomach and a subsequent doubling of gastric evacuation time. The current study is one of the few to illustrate how summation or prioritization of competing physiological systems is manifested in digestive processes.

Cardiovascular system of anomuran crabs, genus Lopholithodes.

The cardiovascular systems of Puget Sound king crabs, Lopholithodes mandtii, and brown box crabs, Lopholithodes foraminatus, were mapped using corrosion casting techniques. Both species have a similar external morphology and a very similar cardiovascular system. Seven arteries (five arterial systems) arise from the heart. The small anterior aorta exits from the anterior surface of the heart and supplies hemolymph to the eyestalks and brain region. The pathway of the two sets of paired arteries, the anterolateral arteries and hepatic arteries, is close, and they intertwine with one another during their initial course. The anterolateral arteries exit from the anterior dorsal surface of the heart and supply hemolymph to the hypodermis, cardiac stomach, antennal gland, and mandibular muscles, whereas the hepatic arteries branch profusely within the hepatopancreas. The lithodids are believed to have evolved from hermit crab ancestors; indicative of these evolutionary origins the posterior aorta is well developed and supplies hemolymph to the large abdomen and the gonads. Exiting from the ventral surface of the heart, the sternal artery is the largest in the system. It branches to supply the mouthparts, chelae, and pereiopods. The differing arrangement of this vessel compared with that of the pagurid anomurans is due to the more carcinized (crab-like) morphological features of the lithodid anomurans. The arrangement of vessels supplying the gills is different compared with that of brachyuran crabs; the infrabranchial sinus joins to the afferent gill vessels at their midpoint, rather than along the ventral edge. In general, the circulatory system of the lithodid crabs is somewhat simpler than that of brachyuran crabs, with fewer branching capillary-like networks. Nevertheless, it is still very complex. In accordance with anatomical descriptions of blue crabs and cancrid crabs it would also seem appropriate to classify the lithodid circulatory system as one that is incompletely closed.

Gastric processing in the Dungeness crab, Cancer magister, during hypoxia.

The gastric physiology of the Dungeness crab, Cancer magister, was investigated over a range of oxygen tensions. Postprandial crabs reacted differently to hypoxia compared with unfed animals. The bradycardic response in postprandial animals was reduced, suggesting a summation of responses with feeding. A similar pattern was observed for ventilation rate. In unfed animals ventilation rate increased slightly as oxygen levels declined, but dropped significantly in oxygen tensions below 3.2 kPa, whereas in postprandial crabs it increased significantly in the lower oxygen regimes. Gastric processing of the meal was followed using a fluoroscope. Pyloric contraction rates were maintained during mild hypoxia, but decreased in 5.3 kPa oxygen tension and below. This led to an increase in clearance times of digesta from the foregut, midgut and hindgut regions. The slowing of gastric processing in the lower oxygen tensions suggested that the animals were unable to maintain their internal oxygen concentration. A significant reduction in efficiency of assimilation only occurred in the lowest oxygen regime tested (1.6 kPa). The range of hypoxia where gastric processing was affected corresponded closely to the levels of oxygen that modulate the foraging behaviour of C. magister. By using both physiological and behavioural mechanisms C. magister can maintain digestive processes, even in severely oxygen depleted environments.

Behavioral influences on the physiological responses of Cancer gracilis, the graceful crab, during hyposaline exposure.

The relationship between the behavioral and physiological responses to hyposaline exposure was investigated in Cancer gracilis, the graceful crab. The status of C. gracilis as an osmoconformer was confirmed. Survival decreased with salinity: the LT(50) in 50% seawater (a practical salinity of 16, or 16 per thousand) was 31.5 +/- 22.7 h and in 25% seawater (a salinity of 8) was 8.0 +/- 0.7 h. When exposed to a salinity gradient, most crabs moved towards the highest salinity. However, in the salinity range of 55% to 65% seawater, they became quiescent. This "closure response" was also evident at low salinities: the mouthparts were tightly closed and animals remained motionless for 2 to 2.5 h. During closure, crabs were able to maintain the salinity of water within the branchial chambers at a level that was about 30% higher than that of the surrounding medium. The closure response was closely linked to a short-term decrease in oxygen uptake. During closure, oxygen within the branchial chamber was rapidly depleted, with oxygen uptake returning to pretreatment levels upon the resumption of activity. In addition to the short-term decrease in oxygen uptake, there was a longer-term bradycardia, which may serve to further reduce diffusive ion loss across the gills. By exhibiting a closure response during acute hyposaline exposure and an avoidance reaction during prolonged or severe hyposaline exposure, C. gracilis is able to use behavior to exploit areas prone to frequent episodes of low salinity.