Iain J. McGaw

Thermoregulatory behavior of the crayfish Procambarus clarki in a burrow environment

The behavioral thermoregulation of the red swamp crayfish, Procambarus clarki, was investigated in its burrow environment. In the field, air and water temperatures within crayfish burrows fluctuated less compared with surface temperatures in the Mojave Desert. However, crayfish could still experience sub-optimal temperature regimes inside burrows. In the laboratory, P. clarki heated and cooled more rapidly in water than in air. In a thermal gradient, the crayfish selected a water temperature of 22 degrees C and avoided water temperatures above 31 degrees C and below 12 degrees C. Observations of behavior in an artificial burrow showed that P. clarki displayed three main shuttling behaviors between water and air in response to temperature. The number of bilateral emersions and emigrations, as well as the amount of time spent in air (in a 24 h period), were significantly greater at 34 degrees C than at 12, 16, 22 or 28 degrees C. This reflected an increased use of the behavioral thermoregulation at temperatures approaching the critical thermal maximum of this species. Upon migrating from 34 degrees C water into 38 degrees C air, crayfish body temperature decreased significantly. These periods of emersion were interspersed with frequent dipping in the water, allowing the crayfish to gain the benefits of evaporative cooling, without the physiological costs incurred by long-term exposure to air.

Circulatory modification in the blue crab Callinectes sapidus, during exposure and acclimation to low salinity

Abstract A pulsed-Doppler flowmeter was used to measure heart rate and haemolymph flow rates in each arterial system of the blue crab, Callinectes sapidus, enabling calculation of stroke volume and cardiac output. During exposure to a 6–6–12 h salinity cycle of 100–25–100% seawater, there was an immediate increase in heart rate upon dilution of the medium. After this initial increase it decreased steadily, but remained elevated above levels in 100% seawater. A smaller increase in heart rate occurred when the salinity was raised, declining thereafter and reaching pre-treatment levels after 6 h in 100% SW. There was a slight decrease in stroke volume of the heart, but overall this resulted in an increase in cardiac output when the salinity was lowered. Differential haemolymph flow through each major arterial system also occurred. There was an increase in flow rates through the anterior aorta, anterolateral arteries and sternal artery during the first 2 h of low salinity exposure and smaller increases occurred again when the salinity was raised to 100% seawater. No significant changes in flow were observed in the hepatic arteries or posterior aorta. During a 72 h acclimation period in low salinity, similar increases in cardiac parameters and flow rates were observed in the first 6 h. These values declined to levels comparable to those in 100% seawater, after 40–50 h acclimation in low salinity. The changes in cardiovascular parameters are not directly related to the osmoregulatory physiology of this species, but appear to be due to specific behaviours occurring in response to low salinity. The results obtained here for this efficient osmoregulator are compared and contrasted with similar studies on Cancer magister, which is classified as a weak hyperosmoregulator.

The Interactive Effects of Exercise and Feeding on Oxygen Uptake, Activity Levels, and Gastric Processing in the Graceful Crab Cancer gracilis

Exercise and digestive processes are known to elevate the metabolic rate of organisms independently. In this study, the effects of simultaneous exercise and digestion were examined in the graceful crab Cancer gracilis. This species exhibited resting oxygen uptake levels between 29 and 42 mg O2 kg−1 h−1. In postprandial crabs, oxygen uptake was approximately double that of unfed crabs. During exercise, oxygen uptake increased three‐ to fourfold, reaching maximal levels of more than 130 mg O2 kg−1 h−1. However, there was no difference in oxygen uptake during activity between unfed and postprandial animals. There was also no difference in exercise endurance levels between unfed and postprandial animals; both sets of animals were unable to right themselves after being turned on their backs, reaching exhaustion after 13–15 attempts. To determine whether increased activity affected gastric processes, the passage of a meal through the digestive system was followed using a fluoroscope. Passage of digesta through the gut system was slower in active animals than in resting crabs. Resting crabs cleared the foregut after approximately 18 h, which was significantly faster than the 34.5 h for constantly active animals. Likewise, the midgut region of resting animals was cleared at a faster rate than that of active animals. Because of residual amounts of digesta remaining in the hindgut, no difference in clearance rates of this section of the gut was evident. The slower clearance times of the foregut were due to a significantly slower rate of mastication of food, as evidenced by a lower cardiac stomach contraction rate. Contraction of the pyloric region of the foregut functions to move the digesta along the midgut, and there was a direct correlation between slower contraction rates of this region and the increased time of passage for digesta through the midgut of active animals. Because increased activity levels affected gastric processing, the crabs exhibited a behavioral response. During a 24‐h period after feeding, there was a significant reduction in locomotor activity. The findings of this study suggest a prioritization of metabolic responses toward activity at the expense of digestion. This is discussed in relation to the ability of the crabs to balance the demands of competing physiological systems.

Prioritization or summation of events? Cardiovascular physiology of postprandial Dungeness crabs in low salinity.

Decapod crustaceans commonly forage in estuarine environments. The osmoregulatory mechanisms that allow them to cope with periodic episodes of low salinity have been well documented. There is less information on how ventilatory and cardiovascular mechanisms aid survival in low salinity. Prior experiments have shown that most species exhibit a tachycardia coupled with an increase in ventilation rate and oxygen uptake. However, these previous experiments were conducted on animals that were starved before experimentation in order to avoid increases in metabolism associated with digestive processes. This study investigated how the Dungeness crab Cancer magister balances the demands of physiological systems during feeding and digestion in low salinity. Cardiac and ventilatory parameters increased during feeding. When the crabs were subjected to low salinity after feeding, heart rate increased in 25% seawater (SW) but decreased in 50% SW. Instead of an expected increase in ventilation rate during low‐salinity exposure, there was a decrease. Feeding was associated with an increase in sternal artery flow, with subsequent decreases in flows through the sternal and anterolateral arteries in low salinity. When low salinity was administered first, a tachycardia occurred, coupled with decreased stroke volume and cardiac output. There was also an increase in ventilation rate. When crabs were fed in low salinity, heart rate decreased in 50% SW but was maintained in 25% SW. Ventilation rate decreased when crabs fed in 50% and 25% SW. Flow through the sternal artery and anterolateral arteries decreased in low salinity, and except for transient increases while feeding, there were further decreases during digestion. Cardiac and ventilatory parameters were rapidly regained when control conditions were restored. The results suggest that events during low salinity are prioritized. Nevertheless, these alterations in physiological parameters may not be beneficial; although digestive processes did not affect osmoregulatory ability, postprandial crabs did not survive as long as starved crabs in 25% SW. The results show that the digestive state of an animal is important in modulating its physiological responses to environmental perturbations, underscoring the importance of an integrative approach to studying physiological responses at the organismal level.

A Review of the “Open” and “Closed” Circulatory Systems: New Terminology for Complex Invertebrate Circulatory Systems in Light of Current Findings

Invertebrate cardiovascular systems have historically been viewed as sluggish, poorly regulated, and “open”, where blood bathes the tissues directly as it moves through a system of ill-defined sinuses and/or lacunae without an endothelial boundary. When examining cardiovascular/circulatory morphology and physiology in a broader evolutionary context, one can question the very nature of the definition of a “closed” versus “open” circulatory system. Viewed in this context a number of invertebrates have evolved incomplete or even completely cell-lined vessels and or lacunae with a highly branched vasculature that allows for the production of significant driving pressures and flows to meet relatively high metabolic demands driven by active life styles. In light of our current understanding of invertebrate cardiovascular systems and their paralleled complexity to vertebrate systems, a number of long established paradigms must be questioned and new definitions presented to better align our understanding of the nature of “open” versus “closed” cardiovascular systems.

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 LT50 in 50% seawater (a practical salinity of 16, or 16‰) 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.

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. J. Morphol., 2008.

Cardiovascular and respiratory responses associated with limb autotomy in the blue crab, Callinectes sapidus

Cardiovascular and respiratory variables were recorded in the blue crab, Callinectes sapidus, during injury and subsequent autotomy of a chela. Cardiac function and haemolymph flow rates were measured using a pulsed-Doppler flowmeter. Oxygen uptake was recorded using an intermittent flow respirometry system. Crabs reacted to the loss of a chela with a rapid increase in heart rate, which was sustained for 2 h. Stroke volume of the heart also increased after the chela was autotomized. A combined increase in heart rate and stroke volume led to an increase in cardiac output, which was maintained for an hour after the loss of a chela. There was also differential haemolymph perfusion of various structures. There was no change in perfusion of the anterolateral arteries or posterior and anterior aortae, during injury of the chela or subsequent autotomy. Haemolymph flow rates did increase significantly through the sternal artery during injury and immediately following autotomy of the chela. This was at the expense of blood flow to the digestive gland: a sustained decrease in haemolymph flow through the hepatic arteries occurred for 3 h following autotomy. Fine-scale cardiac changes associated with the act of autotomy included a bradycardia and/or associated cardiac pausing before the chela was shed, followed by a subsequent increase in cardiac parameters. Changes in the cardiovascular physiology were paralleled by an increase in oxygen uptake, which was driven by an increased ventilation of the branchial chambers. Although limb loss is a major event, it appears that only acute changes in physiology occur. These may benefit the individual, allowing rapid escape following autotomy with a subsequent return to normal activity.

Gastric processing and evacuation during emersion in the red rock crab, Cancer productus

The passage of a radio-opaque meal was followed through the digestive system of the red rock crab, Cancer productus, using a fluoroscope. When the crabs were maintained in seawater, the food was apparent in the foregut as soon as the animals had fed. Release of food from the foregut was routinely slow and digesta appeared in the midgut only in small amounts at any one time. The foregut was emptied between 24 and 36 h, digesta was cleared from the midgut region at 36 h and by 48 h only a small amount of residual digesta was left in the posterior part of the hindgut. Contractions of the cardiac region of the foregut were somewhat sporadic and ranged between 6 and 11 min−1. Contractions of the pyloric region were more stable, varying between 45–65 min−1. In both cases, there was no change in rate during 18 h period in seawater. When crabs were subjected to both short- and long-term aerial exposure, release of food from the foregut was halted for the first 4–6 h of emersion. Although, small amounts of digesta appeared in the midgut and hindgut, there was no significant change in the amount in each region during emersion. There was a trend towards a depression of cardiac stomach contraction rates, but this was only significant in 3 h postprandial crabs during short-term emersion. A pronounced decrease in pyloric stomach contraction rate was maintained for the duration of the aerial exposure. When crabs were returned to seawater, contraction rates took 3–5 h to return to normal, but no significant change in gastric evacuation was observed during this period. During re-immersion, over 65% of the animals regurgitated the stomach contents. This regurgitation may act as a protective mechanism to avoid digestion and the subsequent specific dynamic action. The decrease in gastric processing in C. productus is probably part of an overall metabolic depression occurring during emersion.

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.