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Rules and reciprocity in behavioural development: implications for rehabilitation.

This article considers the potential value of two bodies of thought arising from work on animals in connection with rehabilitating abnormal behaviour. One deals with the processes of catch-up and self-regulation, and the other with optimal periods of learning in development. The ideas about self-regulation are important because an individual with abnormal behaviour at one stage in development may cease to be abnormal of its own accord. The sensitive period concept in development has often been seen as being opposed to that of possible adult rehabilitation. Much of the animal evidence suggests that the opposition is false.

Animals↗

Action imitation in birds.

Action imitation, once thought to be a behavior almost exclusively limited to humans and the great apes, surprisingly also has been found in a number of bird species. Because imitation has been viewed by some psychologists as a form of intelligent behavior, there has been interest in how it is distributed among animal species. Although the mechanisms responsible for action imitation are not clear, we are now at least beginning to understand the conditions under which it occurs. In this article, I try to identify and differentiate the various forms of socially influenced behavior (species-typical social reactions, social effects on motivation, social effects on perception, socially influenced learning, and action imitation) and explain why it is important to differentiate imitation from other forms of social influence. I also examine some of the variables that appear to be involved in the occurrence of imitation. Finally, I speculate about why a number of bird species, but few mammal species, appear to imitate.

Adaptation, Psychological↗

Arousal, but not nursing, is necessary to elicit a decreased fear reaction toward humans in rabbit (Oryctolagus cuniculus) pups.

Rabbits that are handled at the time of feeding during the first week postpartum show reduced fear of humans later in their lives as compared to unhandled controls. Effective handling has been shown to be confined to a sensitive period. Our study aimed to investigate if feeding itself (provided by a second doe, 6 hr after the standard nursing time) affects the levels of fear of humans later in life. Our results showed that (a) the prenursing state of excitement is only characteristic of the standard nursing and is not elicited by a second feeding 6 hr past the usual nursing time, repeated daily across the first week postpartum; and (b) handling linked to a second feeding 6 hr after the standard nursing does not reduce fear responses toward humans at weaning. We conclude that the aroused state, per se, is essential for the reduction of a rabbit's fear response toward humans provoked by early handling.

Animals↗

Distinguishing social and asocial learning using diffusion dynamics.

Theoretical models predict that the cumulative number of individuals displaying a socially learned novel behavior will follow an accelerating pattern over time, whereas asocial processes have been associated with linear or decelerating functions. This raises the possibility that the shape of the diffusion curve may reveal something about the learning processes involved. If true, this would be particularly useful for identifying social transmission in observational field studies. Published data are reviewed and are found to provide limited support for this view. The use of accelerating curves as a diagnostic is challenging because (1) alternative theoretical models make similar predictions, (2) clear supporting empirical data are lacking, and (3) practical considerations frequently make accurate construction of the diffusion curve difficult.

Adaptation, Psychological↗

Information and aggression in fishes.

Aggressive interactions between fishes commonly take place in a social environment in which uninvolved individuals (bystanders) have an opportunity to gather information about interactants. Signals frequently used during such interactions are designed to transmit information about resource-holding power and/or intention. They are generally related to the level of escalation reached and the eventual outcome of a fight. We consider here the information available in signaling and nonsignaling aspects of aggressive interactions. We focus, in particular, on information available to bystanders. We summarize evidence that bystanders alter their behavior toward interactants on the basis of information acquired while bystanding, and we discuss the sources of information that may result in this change of behavior. In particular, we distinguish eavesdropping (i.e., extracting information from signaling interactions) as a subset of bystanding (i.e., extracting information from all available sources). We conclude that considerations of aggressive strategies should include potential costs and benefits resulting from wider social contexts in which aggression occurs.

Adaptation, Psychological↗

How do apes ape?

In the wake of telling critiques of the foundations on which earlier conclusions were based, the last 15 years have witnessed a renaissance in the study of social learning in apes. As a result, we are able to review 31 experimental studies from this period in which social learning in chimpanzees, gorillas, and orangutans has been investigated. The principal question framed at the beginning of this era, Do apes ape? has been answered in the affirmative, at least in certain conditions. The more interesting question now is, thus, How do apes ape? Answering this question has engendered richer taxonomies of the range of social-learning processes at work and new methodologies to uncover them. Together, these studies suggest that apes ape by employing a portfolio of alternative social-learning processes in flexibly adaptive ways, in conjunction with nonsocial learning. We conclude by sketching the kind of decision tree that appears to underlie the deployment of these alternatives.

Adaptation, Psychological↗

Social transmission of courtship behavior and mating preferences in brown-headed cowbirds, Molothrus ater.

Studies of the social learning of courtship behaviors and mating preferences of brown-headed cowbirds (Molothrus ater) are reviewed. Earlier work has suggested that cowbirds from behaviorally distinct populations mate preferentially with others from the same population. Studies are described which indicate that patterns of courtship behavior that differ by population can be socially transmitted across generations are described. Social background affects male songs, female preferences for males as mates, and courtship interactions between females and males. Thus, social traditions influence mating decisions and may, ultimately, impact reproductive success in this species. Recent work on possible social mechanisms involved in the ontogeny of courtship behavior is described. The implications of these findings are discussed, and future research directions are suggested.

Adaptation, Psychological↗

Dissociation between extension of the sensitive period for avian vocal learning and dendritic spine loss in the song nucleus lMAN.

Several instances of early learning coincide with significant rearrangements of neural connections in regions contributing to these behaviors. In fact developmentally restricted learning may be constrained temporally by the opportunity for experience to selectively maintain appropriate synapses amidst the elimination of exuberant connections. Consistent with this notion, during the normal sensitive period for vocal learning in zebra finches (Taenopygia guttata), there is a decline in the density of dendritic spines within a region essential for song development, the lateral magnocellular nucleus of the anterior nidopallium (lMAN). Moreover, in birds isolated from conspecific song shortly after hatching, both the closure of the sensitive period for vocal learning and the pruning of spines from lMAN neurons is delayed. Here, we employed a more subtle form of deprivation to delay the close of the sensitive period for song learning, and found that late song learning occurred without obvious alterations in the pruning of dendritic spines on lMAN neurons. At posthatch day (PHD) 65 (beyond the end of the normal sensitive period for song memorization in zebra finches), birds isolated from song beginning on PHD30 did not differ from normally reared birds in measures of dendritic spine density on Golgi-Cox stained lMAN neurons. Moreover, tutor exposure from PHD65 to 90 did not increase spine elimination in these isolates (who memorized new song material) relative to controls (who did not). Thus, we conclude that the extent of normally occurring lMAN spine loss is not sufficient to account for the timing of the sensitive period for zebra finch song learning.

Animals↗

Approaches to the study of traditional behaviors of free-living animals.

I review literature on four different approaches to the study of traditions in animals: observation of free-living animals, laboratory experiment, armchair analysis, and field experiment. Because, by definition, a tradition entails social learning of some kind, it is difficult, perhaps impossible, to establish that a behavior is in fact traditional without knowledge of how it develops. Observations of free-living animals often provide strong circumstantial evidence of a tradition. However, even in the view of several researchers who have studied possibly traditional behaviors in natural populations, observation alone has not proven sufficient to show that social learning contributes to development of behaviors of interest. The relevance of laboratory experiments to the understanding of the development of behaviors in free-living animals is always open to challenge. Armchair analyses of field data can produce interesting hypotheses but cannot test them. Field experiments to determine how behaviors of interest develop in population members provide a promising way forward.

Adaptation, Psychological↗

Social learning about predators: a review and prospectus.

In comparison with social learning about food, social learning about predators has received little attention. Yet such research is of potential interest to students of animal cognition and conservation biologists. I summarize evidence for social learning about predators by fish, birds, eutherian mammals, and marsupials. I consider the proposal that this phenomenon is a case of S-S classical conditioning and suggest that evolution may have modified some of the properties of learning to accommodate for the requirements of learning socially about danger. I discuss some between-species differences in the properties of socially acquired predator avoidance and suggest that learning may be faster and more robust in species in which alarm behavior reliably predicts high predatory threat. Finally, I highlight how studies of socially acquired predator avoidance can inform the design of prerelease antipredator training programs for endangered species.

Adaptation, Psychological↗

Testing social learning in a wild mountain parrot, the kea (Nestor notabilis).

Huber, Taborsky, and Rechberger (2001) reported an experiment in which the efficiency with which captive keas opened a complex food container was increased by observation of a skilled conspecific. However, only testing social learning in free-ranging animals can demonstrate social learning in natural conditions. For that purpose, a tube-lifting paradigm was developed and tested on keas both in captivity and in Mount Cook National Park, New Zealand. The task was to remove a tube from an upright pole in order to gain access to a reward inside the tube. The top of the pole was higher than a standing kea, so that, to remove the tube, an individual had to simultaneously climb onto the pole and manipulate the tube up the pole with its bill. Because only 1 naive bird managed to remove a tube twice in 25 half-hour sessions and disappeared after success, another bird was trained to solve the task and to provide demonstrations for others. Even under such conditions, only 2 of at least 15 birds learned to remove the tube in 28 sessions. There was no indication that observer birds' use of bill and feet when exploring the tube changed as the number of observations of tube removal increased in a way that would, in principle, increase the likelihood of tube removal. The results suggest a dissociation of social learning potential as assessed in laboratory animals, and social transmission of foraging techniques in natural populations.

Adaptation, Psychological↗

Social learning strategies.

In most studies of social learning in animals, no attempt has been made to examine the nature of the strategy adopted by animals when they copy others. Researchers have expended considerable effort in exploring the psychological processes that underlie social learning and amassed extensive data banks recording purported social learning in the field, but the contexts under which animals copy others remain unexplored. Yet, theoretical models used to investigate the adaptive advantages of social learning lead to the conclusion that social learning cannot be indiscriminate and that individuals should adopt strategies that dictate the circumstances under which they copy others and from whom they learn. In this article, I discuss a number of possible strategies that are predicted by theoretical analyses, including copy when uncertain, copy the majority, and copy if better, and consider the empirical evidence in support of each, drawing from both the animal and human social learning literature. Reliance on social learning strategies may be organized hierarchically, their being employed by animals when unlearned and asocially learned strategies prove ineffective but before animals take recourse in innovation.

Adaptation, Psychological↗