Do rats in a two-action test encode movement egocentrically or allocentrically?

Two-action tests of imitation compare groups that observe topographically different responses to a common manipulandum. The general aim of the two experiments reported here was to find a demonstrator-consistent responding effect in a procedure that could be elaborated to investigate aspects of what was learned about the demonstrated lever response. Experiment 1 was a pilot study with rats of a variant of the two-action method of investigating social learning about observed responses. Groups of observer rats ( Rattus norvegicus) saw a demonstrator push a lever up or down for a food reward. When these observers were subsequently given access to the lever and rewarded for responses in both directions, their directional preferences were compared with two 'screen control' groups that were unable to see their demonstrators' behaviour. Demonstrator-consistent responding was found to be restricted to observers that were able to see demonstrator performance, suggesting that scent cues alone were insufficient to cue a preference for the demonstrators' response direction and thereby that the rats learned by observation about body movements (imitation) or lever movement (emulation). Experiment 2 assessed responding on two levers, one that had been manipulated by the demonstrator, and a second, transposed lever positioned some distance away. Demonstrator-consistent responding was abolished when actions were observed and performed in different parts of the apparatus, suggesting that observed movement was encoded allocentrically with respect to the apparatus rather than egocentrically with respect to the actor's body. With particular reference to the influence of scent cues, the results are discussed in relation to the strengths and weaknesses of this and other varieties of the two-action procedure as tests of imitation in animals and human infants.

Motor learning by observation: evidence from a serial reaction time task.

This study sought evidence of observational motor learning, a type of learning in which observation of the skilled performance of another person not only facilitates motor skill acquisition but does so by contributing to the formation of effector-specific motor representations. Previous research has indicated that observation of skilled performance engages cognitive processes similar to those occurring during action execution or physical practice, but has not demonstrated that these include processes involved in effector-specific representation. In two experiments, observer subjects watched the experimenter performing a serial reaction time (SRT) task with a six-item unique sequence before sequence knowledge was assessed by response time and/or free generation measures. The results suggest that: (1) subjects can acquire sequence information by watching another person performing the task (Experiments 1-2); (2) observation results in as much sequence learning as task practice when learning is measured by reaction times (RTs) and more than task practice when sequence learning is measured by free generation performance (Experiment 2, Part 1); and (3) sequence knowledge acquired by model observation can be encoded motorically--that is, in an effector-specific fashion (Experiment 2, Part 2).

Theory of mind and other domain-specific hypotheses.

The commentators do not contest the target article's claim that there is no compelling evidence of theory of mind in primates, and recent empirical studies further support this view. If primates lack theory of mind, they may still have other behavior control mechanisms that are adaptive in complex social environments. The Somatic Marker Mechanism (SMM) is a candidate, but the SMM hypothesis postulates a much weaker effect of natural selection on social cognition than the theory of mind hypothesis (on inputs to cognitive mechanisms, not on the mechanisms themselves), and there is currently no evidence that it is specific to social stimuli or to primates. "Two Guesser" training would make the goggles test too chauvinistic, and in its current form the goggles problem could not be solved by physical matching because, while wearing goggles, an individual cannot see itself seeing.

Theory of mind in nonhuman primates.

Since the BBS article in which Premack and Woodruff (1978) asked "Does the chimpanzee have a theory of mind?," it has been repeatedly claimed that there is observational and experimental evidence that apes have mental state concepts, such as "want" and "know." Unlike research on the development of theory of mind in childhood, however, no substantial progress has been made through this work with nonhuman primates. A survey of empirical studies of imitation, self-recognition, social relationships, deception, role-taking, and perspective-taking suggests that in every case where nonhuman primate behavior has been interpreted as a sign of theory of mind, it could instead have occurred by chance or as a product of nonmentalistic processes such as associative learning or inferences based on nonmental categories. Arguments to the effect that, in spite of this, the theory of mind hypothesis should be accepted because it is more parsimonious than alternatives or because it is supported by convergent evidence are not compelling. Such arguments are based on unsupportable assumptions about the role of parsimony in science and either ignore the requirement that convergent evidence proceed from independent assumptions, or fail to show that it supports the theory of mind hypothesis over nonmentalist alternatives. Progress in research on theory of mind requires experimental procedures that can distinguish the theory of mind hypothesis from nonmentalist alternatives. A procedure that may have this potential is proposed. It uses conditional discrimination training and transfer tests to determine whether chimpanzees have the concept "see." Commentators are invited to identify flaws in the procedure and to suggest alternatives.

Social learning in animals: categories and mechanisms.

There has been relatively little research on the psychological mechanisms of social learning. This may be due, in part, to the practice of distinguishing categories of social learning in relation to ill-defined mechanisms (Davis, 1973; Galef, 1988). This practice both makes it difficult to identify empirically examples of different types of social learning, and gives the false impression that the mechanisms responsible for social learning are clearly understood. It has been proposed that social learning phenomena be subsumed within the categorization scheme currently used by investigators of asocial learning. This scheme distinguishes categories of learning according to observable conditions, namely, the type of experience that gives rise to a change in an animal (single stimulus vs. stimulus-stimulus relationship vs. response-reinforcer relationship), and the type of behaviour in which this change is detected (response evocation vs. learnability) (Rescorla, 1988). Specifically, three alignments have been proposed: (i) stimulus enhancement with single stimulus learning, (ii) observational conditioning with stimulus-stimulus learning, or Pavlovian conditioning, and (iii) observational learning with response-reinforcer learning, or instrumental conditioning. If, as the proposed alignments suggest, the conditions of social and asocial learning are the same, there is some reason to believe that the mechanisms underlying the two sets of phenomena are also the same. This is so if one makes the relatively uncontroversial assumption that phenomena which occur under similar conditions tend to be controlled by similar mechanisms. However, the proposed alignments are intended to be a set of hypotheses, rather than conclusions, about the mechanisms of social learning; as a basis for further research in which animal learning theory is applied to social learning. A concerted attempt to apply animal learning theory to social learning, to find out whether the same mechanisms are responsible for social and asocial learning, could lead both to refinements of the general theory, and to a better understanding of the mechanisms of social learning. There are precedents for these positive developments in research applying animal learning theory to food aversion learning (e.g. Domjan, 1983; Rozin & Schull, 1988) and imprinting (e.g. Bolhuis, de Vox & Kruit, 1990; Hollis, ten Cate & Bateson, 1991). Like social learning, these phenomena almost certainly play distinctive roles in the antogeny of adaptive behaviour, and they are customarily regarded as 'special kinds' of learning (Shettleworth, 1993).(ABSTRACT TRUNCATED AT 400 WORDS)

Imitation in rats (Rattus norvegicus): The role of demonstrator action.

In a bidirectional control procedure, rats had their first opportunity to push a joystick immediately after observing, from an adjacent compartment, the joystick moving 50 times either to the right or to the left, with each movement signalling the delivery of inaccessible food. Half of these animals observed the joystick moving automatically, and half observed a conspecific demonstrator pushing the joystick. When they were given direct access to the joystick, the observers were rewarded for both left and right pushes. Rats that had observed the joystick moving through the action of a conspecific demonstrator showed a response bias in favour of the observed direction of joystick movement (Experiment 1), while rats that had observed the joystick moving automatically, either in the presence or absence of a passive conspecific, did not show observation-consistent responding (Experiments 1 and 2). These results apparently confirm that rats are capable of imitation or observational learning.

Pharmacological mechanisms and animal models of cognition.

Requirements for an effective animal model of cognition are discussed with special reference to the cholinergic hypothesis of Alzheimer's disease. It is argued, with reference to research on vasopressin and ACE inhibitors, that many putative animal models of cognition lack predictive clinical validity because they either confound the effects of cognitive and arousal processes, or fail to model a specific component of cognitive functioning. A survey of recent research on the cholinergic hypothesis illustrates how these weaknesses can be overcome. Studies involving scopolamine and basal forebrain excitatory amino acid lesion models of the cholinergic deficit in Alzheimer's disease have employed a delayed-matching-to-position test in rodents which, unlike passive avoidance, allows the effects of memory and attentional variables to be distinguished. In combination with recent human studies, these experiments suggest that the cholinergic system has a major role in executive control of attentional resources, and lead to the recommendation of a 'top down' strategy in the investigation of neurochemical processes and pharmacological mechanisms underlying cognition.

Social blockade of taste-aversion learning in Norway rats (Rattus norvegicus): is it a social phenomenon?

In Experiment 1, hooded rats (Rattus norvegicus) were exposed to a novel diet in a food dish or on a conspecific; they were allowed to consume the same diet and then were injected with a toxin LiCl. Later both groups ate more of the novel diet than animals that had not been exposed, and the conspecific-exposed group ate more than the dish-exposed group. Reducing aversion learning by exposure on a conspecific is known as social blockade. We examined if this effect is because a conspecific intensifies dietary cues and thereby increases latent inhibition. Experiment 2 failed to show that diet on a conspecific is a more effective conditioned stimulus for taste-aversion learning than diet in a dish, and Experiment 3 showed that diet in a dish is an effective overshadowing stimulus in aversion learning but diet on a conspecific is not. These results suggest that social blockade cannot readily be assimilated to a latent-inhibition model and may be a distinctly social form of learning.

A demonstration of observational learning in rats using a bidirectional control.

Hungry rats observed a conspecific demonstrator pushing a single manipulandum, a joystick, to the right or to the left for food reward and were then allowed access to the joystick from a different orientation. The effects of right-pushing vs left-pushing observation experience on (1) response acquisition, (2) reversal of a left-right discrimination, and (3) responding in extinction, were examined. Rats that had observed left-pushing made more left responses during acquisition than rats that had observed right-pushing, and rats that had observed demonstrators pushing in the direction that had previously been reinforced took longer to reach criterion reversal and made more responses in extinction than rats that had observed demonstrators pushing in the opposite direction to that previously reinforced. These results provide evidence that rats are capable of learning a response, or a response-reinforcer contingency, through conspecific observation.