Analysis of multiple aspects of a repeated signal: power and rate of rapping during shell fights in hermit crabs

Repeated activities used by animals during contests are assumed to act as signals advertising the quality of the sender. However, their exact functions are not well understood and observations fit only a limited set of the predictions made by models of signaling systems. Experimental studies of contest behavior tend to focus on analysis of the rate of signaling, but individual performances may also vary in magnitude. Both of these features can vary between outcomes and within contests. We examined changes in the rate and power of shell rapping during shell fights in hermit crabs. We show that both rate and power decline during the course of the encounter and that the duration of pauses between bouts of shell rapping increases with an index of the total effort put into each bout. This supports the idea that the vigor of shell rapping is regulated by fatigue and could therefore act as a signal of stamina. By examining different interacting components of this complex activity, we gain greater insight into its function than would be achieved by investigating a single aspect in isolation.

Gender differences, responsiveness and memory of a potentially painful event in hermit crabs

Nonreflexive responses to a noxious event and prolonged memory are key criteria of a pain experience. In a previous study, hermit crabs, Pagurus bernhardus, that received a small electric shock within their shell often temporarily evacuated the shell and some groomed their abdomen and/or moved away from their vital resource. Most, however, returned to the shell. When offered a new shell 20 s later, shocked crabs were more likely than nonshocked crabs to approach and move into a new shell and did so more quickly (Elwood & Appel 2009, Animal Behaviour, 77, 1243-1246). Here we examined how increasing the time between the shock and the offering of a new shell influences the response. There was evidence of a memory of the aversive shock that lasted at least 1 day. Crabs tested after 30 min and 1 day were more likely to approach the shell and new shells were more likely to be taken 30 min after the shock. Shocked crabs approached the new shell more quickly and used fewer probes of the chelipeds prior to moving in and these results were stable over time and significant for specific times up to 1 day. Females were more likely than males to evacuate shells and did so after fewer shocks. These results extend previous work and demonstrate an extended memory of having been shocked. The findings are consistent with respect to criteria for pain that are accepted for vertebrates.

Motivational trade-offs and potential pain experience in hermit crabs

One criterion of pain experience is that the emotional response to pain may be traded-off against other motivational requirements. This was tested in hermit crabs, housed in either preferred or unpreferred species of shells, by subjecting their abdomens to electric shocks of gradually increasing intensity. The first observable response was not affected by shell species but those in preferred shells evacuated at a higher shock level than those in poor quality shells. Thus, they seem to trade-off the requirement to retain a high quality shell with that of avoidance of the noxious stimulus. Some crabs returned to their shells and those that got back into the preferred species did so with less probing of the aperture before getting in and subsequently thrust their abdomen in and out less often in further investigation, thus confirming their shell species preference. Not all crabs returned to the vicinity of the shell and some attempted to climb the wall of the experimental chamber. Others engaged in shell rapping as if in a fight and grooming of the abdomen was noted. These findings are consistent with the idea of a pain experience rather than a nociceptive reflex. (C) 2009 Elsevier B.V. All rights reserved.

Pain experience in hermit crabs?

Pain may be inferred when the responses to a noxious stimulus are not reflexive but are traded off against other motivational requirements, the experience is remembered and the situation is avoided in the future. To investigate whether decapods feel pain we gave hermit crabs, Pagurus bernhardus, small electric shocks within their shells. Only crabs given shocks evacuated their shells indicating the aversive nature of the stimulus, but fewer crabs evacuated from a preferred species of shell indicating a motivational trade-off. Some crabs that evacuated attacked the shell in the manner seen in a shell fight. Most crabs, however, did not evacuate at the stimulus level we used, but when these were subsequently offered a new shell, shocked crabs were more likely to approach and enter the new shell. Furthermore, they approached that shell more quickly, investigated it for a shorter time and used fewer cheliped probes within the aperture prior to moving in. Thus the experience of the shock altered future behaviour in a manner consistent with a marked shift in motivation to get a new shell to replace the one occupied. The results are consistent with the idea of pain in these animals. (C) 2009 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

The power of shell rapping influences rates of eviction in hermit crabs

Hermit crabs fight for ownership of shells, and shell exchange may occur after a period of shell rapping, involving the initiating or attacking crab bringing its shell rapidly and repeatedly into contact with the shell of the noninitiator or defender, in a series of bouts. The temporal pattern of rapping contains information about the motivation and/or relative resource holding potential (RHP) of the initiator and acts as a repeated signal of stamina. Here we investigated the role of the force with which the rapping is performed and how this is related to the temporal pattern of rapping by rubberizing the external surface of shells. Initiators that are prevented from rapping with their usual level of force persist with the activity for longer over the whole encounter but use fewer raps per bout and are less likely to effect an exchange than those supplied with control shells. The fact that the force of rapping affects the likelihood of a crab being victorious suggests that either the force of rapping contains information about motivation or RHP or that force directly affects noninitiators, reducing their ability to maintain an adequate grip on their shells. The data suggest that shell rapping is an agonistic signal rather than one that provides information useful to the noninitiator, as has been suggested by the negotiation model of shell exchange.

How resource quality differentially affects motivation and ability to fight in hermit crabs

Contesting animals typically gather information about the resource value and that information affects fight motivation. However, it is possible that particular resource characteristics alter the ability to fight independently of the motivation. Using hermit crabs, we investigate how the resource in terms of shell quality affects both motivation and ability to fight. These crabs fight for shells, but those shells have to be carried and may impose physiological costs that impede fight vigour. We find that the shell has different effects on motivation and ability. Potential attackers in very small shells were highly motivated to attack but, rather than having enhanced ability, unexpectedly quickly fatigued and subsequently were not more successful in the fights than were crabs in larger shells. We also examined whether defending crabs could gather information about the attacker's shell from the vigour of the attack. Defending crabs gave up quickly when a potential gain had been assessed, indicating that such information had been gathered. However, there was no indication that this could be owing to the activity of the attacker and the information is probably gathered via visual assessment of the shell.

Hermit crabs perceive the extent of their virtual bodies

A flexible body image is required by animals if they are to adapt to body changes and move effectively within a structurally complex environment. Here, we show that terrestrial hermit crabs, Coenobita rugosus, which frequently change shells, can modify walking behaviour, dependent on the shape of the shell. Hermit crabs walked along a corridor that had alternating left and right corners; if it was narrow at the corner, crabs rotated their bodies to avoid the wall, indicating an awareness of environmental obstacles. This rotation increased when a plastic plate was attached to the shell. We suggest that the shell, when extended by the plate, becomes assimilated to the hermit crab's own body. While there are cases of a tool being assimilated with the body, our result is the first example of the habitat where an animal lives and/or carries being part of a virtual body. This journal is © 2012 The Royal Society.