No evidence of morphine analgesia to noxious shock in the shore crab, Carcinus maenas

A number of criteria have been suggested for testing if pain occurs in animals, and these include an analgesic effect of opiates (Bateson, 1991). Morphine reduces responses to noxious stimuli in crustaceans but also reduces responsiveness in a non-pain context. Here we use a paradigm in which shore crabs receive a shock in a preferred dark shelter but not if they remain in an unpreferred light area. Analgesia should thus enhance movement to the preferred dark area because they should not experience 'pain'. However, morphine inhibits rather than enhances this movement even when no shock is given. Morphine produces a general effect of non-responsiveness rather than a specific analgesic effect and this could also explain previous studies claiming analgesia. However, we question the utility of this criterion of pain and suggest instead that behavioural criteria be employed. (C) 2011 Published by Elsevier B.V.

Effects of the organophosphorous pesticide, dimethoate, on cardiac and acetylcholinesterase (AChE) activity in the shore crab Carcinus maenas

The potential use of biochemical and physiological responses as biomarkers of organophosphate exposure and/or effect were assessed in the shore crab (Carcinus maenas). Male crabs were assigned to one of four dimethoate treatments (0, 0.5, 1.0 or 2.0 mg 1(-1)). Cardiac activity was measured non-invasively before and during dimethoate exposure using automated interpulse duration assessment. Heart rates decreased significantly in a concentration-dependent manner (p

Shock avoidance by discrimination learning in the shore crab (Carcinus maenas) is consistent with a key criterion for pain

Nociception allows for immediate reflex withdrawal whereas pain allows for longer-term protection via rapid learning. We examine here whether shore crabs placed within a brightly lit chamber learn to avoid one of two dark shelters when that shelter consistently results in shock. Crabs were randomly selected to receive shock or not prior to making their first choice and were tested again over 10 trials. Those that received shock in trial 2, irrespective of shock in trial 1, were more likely to switch shelter choice in the next trial and thus showed rapid discrimination. During trial 1, many crabs emerged from the shock shelter and an increasing proportion emerged in later trials, thus avoiding shock by entering a normally avoided light area. In a final test we switched distinctive visual stimuli positioned above each shelter and/or changed the orientation of the crab when placed in the chamber for the test. The visual stimuli had no effect on choice, but crabs with altered orientation now selected the shock shelter, indicating that they had discriminated between the two shelters on the basis of movement direction. These data, and those of other recent experiments, are consistent with key criteria for pain experience and are broadly similar to those from vertebrate studies. © 2013. Published by The Company of Biologists Ltd.

Breaking and Entering: Examining the role of stress and aerial exposure in predator-prey relationships between the common shore crab (Carcinus maenas) and cultivated blue mussels (Mytilus edulis).

During benthic cultivation Mytilus edulis (blue mussels) are subject to predation pressure from a number of predators including Carcinus maenas (shore crabs). This predator can be responsible for substantial losses of mussels from the fishery and a full understanding of the predator–prey relationship between M. edulis and C. maenas is required to ensure attempts that reduce predatory pressure and subsequent commercial loss are successful. Whilst much work has examined the prey–predator size relationships between C. maenas and M. edulis, far less research has investigated how stress, such as periods of extended aerial exposure, may affect these relationships. We tested whether profit in terms of calories gained by crabs consuming mussels stressed by aerial exposure for 48 h differed from that of mussels at ambient conditions and whether being stressed affected the mussel's likelihood of predation. We also tested whether the size relationship between predators and their prey differed when mussels were stressed. We found that the profitability of prey (calories gained per second of handling time) did not vary between stressed and unstressed mussels. Handling times for stressed and unstressed mussels were similar, even when crabs were presented with mussels of the maximum size that they are able to consume. Small crabs were more likely to reject a mussel of preferred size if it was unstressed, suggesting that crabs may be able to assess that these mussels would require extra effort to break into and consume. Our findings suggest that the predator–prey relationship between mussels and crabs is not altered when mussels are stressed. C. maenas remains a voracious predator and regardless of the condition of mussels laid on commercial beds there is a need to control this predator in attempt to reduce losses in the benthic fishery.

Effects of baited crab pots on cultivated mussel (Mytilus edulis) survival rates

The shore crab, Carcinus maenas, is recognized as a voracious predator of blue mussels, Mytilus edulis, having the potential to greatly reduce stocks in the benthic cultivation industry. As a consequence, baited crab pots are often deployed on and around cultivated benthic mussel beds to trap and remove crabs, in an attempt to reduce predatory pressure. Little is known about how C. maenas behaves around crab pots, but for many other crustacean fisheries, the trapping efficiency of pots is often low. Crabs may be attracted towards but not enter pots, instead feeding on cultivated mussels outside pots on the surrounding substratum. We tested whether the rate of loss of mussels attached to plates differed in areas next to baited pots compared with unbaited pots and to areas without any pots, at two sea loughs (60 km apart) in Northern Ireland. In Strangford Lough, more mussels were lost from plates next to baited pots than the other treatments. In Carlingford Lough, however, we found no difference in the number of mussels lost from plates in any treatment. This difference could be attributed to the different assemblages of mobile benthic predators at the two loughs. The presence of the starfish Asterias rubens, which was absent from experimental sites in Carlingford Lough, was thought to be responsible for increased predation rates near baited pots in Strangford. It is, therefore, important to consider local predator communities when deploying crab pots as a predator mitigation technique to ensure predation rates are in fact reduced and not enhanced. This study is of relevance not only to attempts to limit predation on commercial stocks of benthic cultivated mussels but also in situations where baited traps are deployed close to species vulnerable to mobile benthic predators.

No discrimination shock avoidance with sequential presentation of stimuli but shore crabs still reduce shock exposure

Insights into the potential for pain may be obtained from examination of behavioural responses to noxious stimuli. In particular, prolonged responses coupled with long-term motivational change and avoidance learning cannot be explained by nociceptive reflex but are consistent with the idea of pain. Here, we placed shore crabs alternately in two halves of a test area divided by an opaque partition. Each area had a dark shelter and in one repeated small electric shocks were delivered in an experimental but not in a control group. Crabs showed no specific avoidance of the shock shelter either during these trials or in a subsequent test in which both were offered simultaneously; however they often emerged from the shock shelter during a trial and thus avoided further shock. More crabs emerged in later trials and took less time to emerge than in early trials. Thus, despite the lack of discrimination learning between the two shelters they used other tactics to markedly reduce the amount of shock received. We note that a previous experiment using simultaneous presentation of two shelters demonstrated rapid discrimination and avoidance learning but the paradigm of sequential presentation appears to prevent this. Nevertheless, the data show clearly that the shock is aversive and tactics, other than discrimination learning, are used to avoid it. Thus, the behaviour is only partially consistent with the idea of pain.

Size-balanced community reorganization in response to nutrients and warming

It is widely accepted that global warming will adversely affect ecological communities. As ecosystems are simultaneously exposed to other anthropogenic influences, it is important to address the effects of climate change in the context of many stressors. Nutrient enrichment might offset some of the energy demands that warming can exert on organisms by stimulating growth at the base of the food web. It is important to know whether indirect effects of warming will be as ecologically significant as direct physiological effects. Declining body size is increasingly viewed as a universal response to warming, with the potential to alter trophic interactions. To address these issues, we used an outdoor array of marine mesocosms to examine the impacts of warming, nutrient enrichment and altered top-predator body size on a community comprised of the predator (shore crab Carcinus maenas), various grazing detritivores (amphipods) and algal resources. Warming increased mortality rates of crabs, but had no effect on their moulting rates. Nutrient enrichment and warming had near diametrically opposed effects on the assemblage, confirming that the ecological effects of these two stressors can cancel each other out. This suggests that nutrient-enriched systems might act as an energy refuge to populations of species under metabolic constraints due to warming. While there was a strong difference in assemblages between mesocosms containing crabs compared to mesocosms without crabs, decreasing crab size had no detectable effect on the amphipod or algal assemblages. This suggests that in allometrically balanced communities, the expected long-term effect of warming (declining body size) is not of similar ecological consequence to the direct physiological effects of warming, at least not over the six week duration of the experiment described here. More research is needed to determine the long-term effects of declining body size on the bioenergetic balance of natural communities.

Physiological stress responses in the edible crab, Cancer pagurus, to the fishery practice of de-clawing

We examined physiological stress responses in the edible crab, Cancer pagurus, subjected to the commercial fishery practice of manual de-clawing. We measured haemolymph glucose and lactate, plus muscular glycogen and glycogen mobilisation, in three experiments where the crabs had one claw removed. In the first, crabs showed physiological stress responses when 'de-clawed' as compared to 'handled only over the short term of 1-10 min. In the second, de-clawing and the presence of a conspecific both increased the physiological stress responses over the longer term of 24 h. In the third, de-clawing was shown to be more stressful than 'induced autotomy' of claws. Further, the former practice caused larger wounds to the body and significantly higher mortality than the latter. Since the fishery practice is to remove both claws, the stress response observed and mortality data reported are conservative.

Fighting for shells: how private information about resource value changes hermit crab pre-fight displays and escalated fight behaviour

Pre-fight displays typically provide honest, but sometimes dishonest, information about resource holding potential and may be influenced by assessment of resource value and hence motivation to acquire the resource. These assessments of potential costs and benefits are also predicted to influence escalated fight behaviour. This is examined in shell exchange contests of hermit crabs in which we establish an information asymmetry about a particularly poor quality shell. The poor shell was created by gluing sand to the interior whereas control shells lacked sand and the low value of the poor shell could not be accurately assessed by the opponent. Crabs in the poor shell showed changes in the use of pre-fight displays, apparently to increase the chances of swapping shells. When the fights escalated, crabs in poor shells fought harder if they took the role of attacker but gave up quickly if in the defender role. These tactics appear to be adaptive but do not result in a major shift in the roles taken or outcome. We thus link resource assessment with pre-fight displays, the roles taken, tactics used during escalation and the outcome of these contests.

Monoamines and decision making during contests in the hermit crab Pagurus bernhardus

During fights animals are expected to make a series of strategic decisions that involve interactions between information about the contest and the individual's nervous system that produce a change in behaviour. Biogenic monoamines such as serotonin ('5-HT') and dopamine are thought to prime decision-making centres for appropriate responses during aggressive interactions in crustaceans, and circulating levels vary both between individuals and during agonistic encounters. Aminergenic systems operate in diverse animal taxa and in this study we assayed circulating levels of S-HT and dopamine following shell fights in the common European hermit crab, Pagurus bernhardus. The two roles in these fights, attacker and defender, perform different activities but, in both, S-HT increased and dopamine declined in response to engaging in a fight. In defenders but not attackers, giving up was correlated with low 5-HT and dopamine. In attackers, motivation to initiate a fight was positively correlated with dopamine levels. Circulating monoamines are therefore involved in decision making during these aggressive encounters. (c) 2007 The Association for the Study of Animal Behaviour Published by Elsevier Ltd. All rights reserved.

Honest and dishonest displays, motivational state and subsequent decisions in hermit crab shell fights

Animal fights are typically preceded by displays and there is debate whether these are always honest. We investigated the prefight period in hermit crabs, Pagurus bernhardus, during which up to four types of display plus other activities that might provide information are performed. We determined how each display influences or predicts various fight decisions, and related these displays to the motivational state of the attacker, as determined by a startle response, and of the motivational state of the defender, as determined by the duration for which it resisted eviction from its shell. Two displays appeared to have consistent but different effects. Cheliped presentation, where the claws were held in a stationary position, often by both crabs but for longer by the larger, seemed to be honest, and allowed for mutual size assessment. This display enhanced the motivation and the success of the larger crab. In contrast, cheliped extension, involving the rapid thrust of the open chelae towards the opponent, did not seem to allow for mutual size assessment and may contain an element of bluff. It was performed more by the smaller crab and enhanced its success. The complexity of displays in this species appears to allow for both honesty and manipulation.

Motivational change during shell fights in the hermit crab Pagurus bernhardus

During agonistic interactions the motivation of each contestant is expected to vary because of increased information and changes in fighting ability. In shell fights between hermit crabs over gastropod shells, attackers rap their shell in a series of bouts against that of the defender whereas defenders remain withdrawn into their shells until the encounter is resolved; either the defender is evicted from its shell or the attacker 'gives up' and the defender retains its shell. We assessed the motivational state of attackers for performing rapping by measuring the duration of startle responses elicited by a novel stimulus. We staged fights between pairs of crabs in six different groups defined by the potential gain in shell quality available to attackers (high or low) and by the point at which the novel stimulus was applied (prior to rapping, after one bout or after four bouts). Startle response duration decreased during the first four bouts of fighting and showed a U-shaped relationship with the relative difference in size between the crabs. There was, no difference in startle response duration between high- and low-gain groups. Individuals showing short startle responses were likely to be victorious and we conclude that the relationship between the relative size difference of the opponents and. startle duration reflects that between size difference and the cost of gaining an eviction. (C) 2001 The Association for the Study of Animal Behaviour.

Decision rules, energy metabolism and vigour of hermit-crab fights

Aggressive interactions between animals are often settled by the use of repeated signals that reduce the risk of injury from combat but are expected to be costly. The accumulation of lactic acid and the depletion of energy stores may constrain activity rates during and after fights and thus represent significant costs of signalling. We tested this by analysing the concentrations of lactate and glucose in the haemolymph of hermit crabs following agonistic interactions over the ownership of the gastropod shells that they inhabit. Attackers and defenders play distinct roles of sender and receiver that are fixed for the course of the encounter. Attackers perform bouts of 'shell rapping', which vary in vigour between attackers and during the course of the encounter, and are a key predictor of victory. In contrast to the agonistic behaviour of other species, we can quantify the vigour of fighting. We demonstrate, to our knowledge for the first time, an association between the vigour of aggressive activity and a proximate cost of signalling. We show that the lactate concentration in attackers increases with the amount of shell rapping, and that this appears to constrain the vigour of subsequent rapping. Furthermore, attackers, but not defenders, give up when the concentration of lactate is high. Glucose levels in attackers also increase with the amount of rapping they perform, but do not appear to influence their decision to give up. Defenders are more likely to lose when they have particularly low levels of glucose. We conclude that the two roles use different decision rules during these encounters.