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Fischhoff, I. R., Sundaresan, S. R., Cordingley, J., Larkin, H. M., Sellier, M. - J., & Rubenstein, D. I. (2007). Social relationships and reproductive state influence leadership roles in movements of plains zebra, Equus burchellii. Anim. Behav., 73(5), 825–831.
Abstract: In animal groups, collective movements emerge from individual interactions. Biologists seek to identify how characteristics of actors in these groups, and their relationships, influence the decision-making process. We distinguished two basic factors determining leadership in group choices: identity and state. We hypothesized that identity is more important to leadership in groups with stable relationships, which permit the development of habitual roles. In groups with fluid membership, particular individuals or subgroups are less likely to emerge as consistent leaders. Instead, we predicted that movement initiation in unstable groups depends on individual state at the time of the decision. We characterized how identity and reproductive state influenced leadership patterns in the movements of plains zebra. As in many other mammals, lactation in this species significantly alters water and energy needs. We investigated leadership in tightly knit harems and loosely bonded herds of multiple harems. Harem females tended to have habitual roles in the initiation of harem movement. In herds, however, we found no consistent leaders among harems. At both levels of social organization, lactation was a key determinant of leadership. In harems, lactating females were more likely to initiate movement than nonlactating females. In turn, harems containing lactating females were more likely to lead herd movements. Thus, we conclude that social relationships and reproductive state together shape the interactions that produce group behaviours. One benefit to lactating females of leading herd movements is preferential access to scarce water. Thus, leadership roles in group decisions may have fitness consequences.
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Lingle, S., Rendall, D., Wilson, W. F., DeYoung, R. W., & Pellis, S. M. (2007). Altruism and recognition in the antipredator defence of deer: 2. Why mule deer help nonoffspring fawns. Anim. Behav., 73(5), 907–916.
Abstract: Both white-tailed deer, Odocoileus virginianus, and mule deer, O. hemionus, females defend fawns against coyotes, Canis latrans, but only mule deer defend nonoffspring conspecific and heterospecific fawns. During a predator attack, females may have to decide whether to defend a fawn while having imperfect information on its identity obtained from hearing a few distress calls. Although imperfect recognition can influence altruistic behaviour, few empirical studies have considered this point when testing functional explanations for altruism. We designed a series of playback experiments with fawn distress calls to test alternative hypotheses (by-product of parental care, kin selection, reciprocal altruism) for the mule deer's defence of nonoffspring, specifically allowing for the possibility that females mistake these fawns for their own. White-tailed deer females approached the speaker only when distress calls of white-tailed deer fawns were played and when their own fawn was hidden, suggesting that fawn defence was strictly a matter of parental care in this species. In contrast, mule deer females responded similarly and strongly, regardless of the caller's identity, the female's reproductive state (mother or nonmother) or the presence of their own offspring. The failure of mule deer females to adjust their responses to these conditions suggests that they do not defend nonoffspring because they mistake them for their own fawns. The lack of behavioural discrimination also suggests that kin selection, reciprocal altruism and defence of the offspring's area are unlikely to explain the mule deer's defence of nonoffspring. We identify causal and functional questions that still need to be addressed to understand why mule deer defend fawns so indiscriminately.
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Zentall, T. R. (2006). Imitation: definitions, evidence, and mechanisms. Anim. Cogn., 9(4), 335–353.
Abstract: Imitation can be defined as the copying of behavior. To a biologist, interest in imitation is focused on its adaptive value for the survival of the organism, but to a psychologist, the mechanisms responsible for imitation are the most interesting. For psychologists, the most important cases of imitation are those that involve demonstrated behavior that the imitator cannot see when it performs the behavior (e.g., scratching one's head). Such examples of imitation are sometimes referred to as opaque imitation because they are difficult to account for without positing cognitive mechanisms, such as perspective taking, that most animals have not been acknowledged to have. The present review first identifies various forms of social influence and social learning that do not qualify as opaque imitation, including species-typical mechanisms (e.g., mimicry and contagion), motivational mechanisms (e.g., social facilitation, incentive motivation, transfer of fear), attentional mechanisms (e.g., local enhancement, stimulus enhancement), imprinting, following, observational conditioning, and learning how the environment works (affordance learning). It then presents evidence for different forms of opaque imitation in animals, and identifies characteristics of human imitation that have been proposed to distinguish it from animal imitation. Finally, it examines the role played in opaque imitation by demonstrator reinforcement and observer motivation. Although accounts of imitation have been proposed that vary in their level of analysis from neural to cognitive, at present no theory of imitation appears to be adequate to account for the varied results that have been found.
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Weatherly, J. N., Arthur, E. I. L., & Tischart, L. M. (2003). Altering “motivational” variables alters induction produced by upcoming food-pellet reinforcement. Anim. Cogn., 6(1), 17–26.
Abstract: Previous research has demonstrated that rats will increase their rates of lever pressing for sucrose rewards in the first half of an experimental session when food pellets, rather than the same sucrose, continually serve as the reward in the second half of the session. This effect has been coined induction, and the present study investigated whether it could be altered by altering “motivational” variables. Experiment 1 manipulated subjects' motivation by altering, across conditions, their level of food deprivation. Predictably, the size of induction varied directly with level of deprivation. Experiments 2 and 3 manipulated subjects' motivation by feeding them food pellets and sucrose, respectively, prior to their responding in the experimental session. These pre-session feedings decreased the size of the observed induction in both experiments. The results from the present study indicate that the size of induction is correlated with subjects' motivation to respond for the available reinforcers. They are also consistent with the idea that operant processes underlie the effect. The notion that induction might encompass the concept of “anticipation” is also discussed.
Keywords: Animals; *Conditioning, Operant; Food Deprivation; Male; *Motivation; Rats; Rats, Sprague-Dawley
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Waite, T. A. (2002). Interruptions improve choice performance in gray jays: prolonged information processing versus minimization of costly errors. Anim. Cogn., 5(4), 209–214.
Abstract: Under the assumption that selection favors minimization of costly errors, erroneous choice may be common when its fitness cost is low. According to an adaptive-choice model, this cost depends on the rate at which an animal encounters the choice: the higher this rate, the smaller the cost of choosing a less valuable option. Errors should thus be more common when interruptions to foraging are shorter. A previous experiment supported this prediction: gray jays, Perisoreus canadensis, were more error prone when subjected to shorter delays to access to food rewards. This pattern, though, is also predicted by an attentional-constraints model. Because the subjects were able to inspect the rewards during delays, their improved performance when subjected to longer delays could have been a byproduct of the experimentally prolonged opportunity for information processing. To evaluate this possibility, a follow-up experiment manipulated both delay to access and whether rewards could be inspected during delays. Depriving jays of the opportunity to inspect rewards (using opaque lids) induced only a small, nonsignificant increase in error rate. This effect was independent of length of delay and so the jays' improved performance when subjected to longer delays was not simply a byproduct of prolonged information processing. More definitively, even when the jays were prevented from inspecting rewards during delays, their performance improved when subjected to longer delays. The findings are thus consistent with the adaptive-choice model.
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Tebbich, S., Bshary, R., & Grutter, A. S. (2002). Cleaner fish Labroides dimidiatus recognise familiar clients. Anim. Cogn., 5(3), 139–145.
Abstract: Individual recognition has been attributed a crucial role in the evolution of complex social systems such as helping behaviour and cooperation. A classical example for interspecific cooperation is the mutualism between the cleaner fish Labroides dimidiatus and its client reef fish species. For stable cooperation to evolve, it is generally assumed that partners interact repeatedly and remember each other's past behaviour. Repeated interactions may be achieved by site fidelity or individual recognition. However, as some cleaner fish have more than 2,300 interactions per day with various individuals per species and various species of clients, basic assumptions of cooperation theory might be violated in this mutualism. We tested the cleaner L. dimidiatus and its herbivorous client, the surgeon fish Ctenochaetus striatus, for their ability to distinguish between a familiar and an unfamiliar partner in a choice experiment. Under natural conditions, cleaners and clients have to build up their relationship, which is probably costly for both. We therefore predicted that both clients and cleaners should prefer the familiar partner in our choice experiment. We found that cleaners spent significantly more time near the familiar than the unfamiliar clients in the first 2 minutes of the experiment. This indicates the ability for individual recognition in cleaners. In contrast, the client C. striatus showed no significant preference. This could be due to a sampling artefact, possibly due to a lack of sufficient motivation. Alternatively, clients may not need to recognise their cleaners but instead remember the defined territories of L. dimidiatus to achieve repeated interactions with the same individual.
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Shettleworth, S. J. (1985). Foraging, memory, and constraints on learning. Ann N Y Acad Sci, 443, 216–226.
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Seyfarth, R. M., & Cheney, D. L. (2003). Signalers and receivers in animal communication. Annu Rev Psychol, 54, 145–173.
Abstract: In animal communication natural selection favors callers who vocalize to affect the behavior of listeners and listeners who acquire information from vocalizations, using this information to represent their environment. The acquisition of information in the wild is similar to the learning that occurs in laboratory conditioning experiments. It also has some parallels with language. The dichotomous view that animal signals must be either referential or emotional is false, because they can easily be both: The mechanisms that cause a signaler to vocalize do not limit a listener's ability to extract information from the call. The inability of most animals to recognize the mental states of others distinguishes animal communication most clearly from human language. Whereas signalers may vocalize to change a listener's behavior, they do not call to inform others. Listeners acquire information from signalers who do not, in the human sense, intend to provide it.
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Meehan, C. L., & Mench, J. A. (2007). The challenge of challenge: Can problem solving opportunities enhance animal welfare? Appl. Anim. Behav. Sci., 102(3-4), 246–261.
Abstract: Cognitive mechanisms are an important part of the organization of the behavior systems of animals. In the wild, animals regularly face problems that they must overcome in order to survive and thrive. Solving such problems often requires animals to process, store, retrieve, and act upon information from the environment--in other words, to use their cognitive skills. For example, animals may have to use navigational, tool-making or cooperative social skills in order to procure their food. However, many enrichment programs for captive animals do not include the integration of these types of cognitive challenges. Thus, foraging enrichments typically are designed to facilitate the physical expression of feeding behaviors such as food-searching and food consumption, but not to facilitate complex problem solving behaviors related to food acquisition. Challenging animals by presenting them with problems is almost certainly a source of frustration and stress. However, we suggest here that this is an important, and even necessary, feature of an enrichment program, as long as animals also possess the skills and resources to effectively solve the problems with which they are presented. We discuss this with reference to theories about the emotional consequences of coping with challenge, the association between lack of challenge and the development of abnormal behavior, and the benefits of stress (arousal) in facilitating learning and memory of relevant skills. Much remains to be done to provide empirical support for these theories. However, they do point the way to a practical approach to improving animal welfare--to design enrichments to facilitate the cognitive mechanisms which underlie the performance of complex behaviors that cannot be performed due to the restrictions inherent to the captive environment.
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Søndergaard, E., Jensen, M. B., & Nicol, C. J. (). Motivation for social contact in horses measured by operant conditioning. Appl. Anim. Behav. Sci., In Press, Corrected Proof.
Abstract: Although horses are social animals they are often housed individually with limited social contact to other horses and this may compromise their welfare. The present study included eight young female horses and investigated the strength of motivation for access to full social contact, head contact and muzzle contact, respectively, to a familiar companion horse. Horses were housed individually next to their companion horse and separations between pens prevented physical contact. During daily test sessions horses were brought to a test area where they could access an arena allowing social contact. Arena access during 3 min was given after completion of a predetermined number of responses on a panel. Fixed ratios (FR) of 8, 16, 24, 32 and 40 responses per arena access were applied in a random order, one per daily test session, within each test week (Monday to Friday), and the number of rewards per daily test session was recorded. All horses could access all three types of social contact in a cross-over design, and an empty arena was used as control. Motivational strength was assessed using elasticity of demand functions, which were estimated based on the number of rewards earned and FR. Elasticities of demand for the three types of social contact were low (-0.20), and not significantly different, although increasing FR still resulted in a decrease in rewards obtained for all three types of social contact (P < 0.001). Across FR-levels horses earned more rewards for social contact than for an empty arena, as shown by much higher intercept values (2.51 vs. 0.99; P < 0.001). However, the elasticity of demand for infrequent access to an empty arena (-0.08) was lower than for social contact (P < 0.01) and not significantly different from zero (P = 0.07). Horses performed more social behaviour the lesser the restriction on social contact (full > head > muzzle). However, the finding that horses showed a similar and high motivation for all three types of social contact suggests that they are valued equally highly in a situation where the alternative is no social contact.
Keywords: Horse; Housing; Social behaviour; Operant conditioning; Motivation
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