Ludwig Huber

True imitation in marmosets

Marmosets, Callithrix jacchus, observed a demonstrator removing the lids from a series of plastic canisters to obtain a mealworm. When subsequently allowed access to the canisters, marmosets that observed a demonstrator using its hands to remove the lids used only their hands. In contrast, marmosets that observed a demonstrator using its mouth also used their mouth to remove the lids. Since hand and mouth demonstrators brought about identical changes in the canisters, the differential test behaviour of the observer groups suggests that they learned about the demonstrators behaviour. Furthermore, marmosets that had not been given the opportunity to observe a demonstrator prior to testing had a low probability of mouth opening, even if the canisters were previously opened by a mouth-opening demonstrator in an olfactory control experiment. Corroborating Bugnyar & Hubers (1997, Animal Behaviour, 54, 817-831) earlier findings, our results provide further evidence that marmosets can imitate. Copyright 2000 The Association for the Study of Animal Behaviour.

Push or pull: an experimental study on imitation in marmosets

A laboratory experiment was conducted in order to explore the possibility of imitation, that is, response learning by observation, in marmosets, Callithrix jacchusInexperienced individuals were allowed to observe a skilful model that demonstrated one of two possible techniques (pushing or pulling a pendulum-door) to get food from inside a wooden box. Their initial manipulative actions, performed when exposed to the box in a subsequent test, were compared with those of naive control subjects (non-observers). The observers showed less exploratory behaviour than the non-observers and, more importantly, some showed a strong tendency to use the demonstrated opening technique in the initial test phase. This initial preference disappeared in the course of five test sessions and the observers converged towards the simpler, alternative solution that was generally preferred by the non-observers. Despite fundamental individual differences in the observer group and the failure to find a significant group effect, the results indicate that marmosets are capable of learning simple motor skills through conspecific observation.Copyright 1997 The Association for the Study of Animal Behaviour1997The Association for the Study of Animal Behaviour

Obey or not obey? Dogs (Canis familiaris) behave differently in response to attentional states of their owners.

Sixteen domestic dogs (Canis familiaris) were tested in a familiar context in a series of 1-min trials on how well they obeyed after being told by their owner to lie down. Food was used in 1/3 of all trials, and during the trial the owner engaged in 1 of 5 activities. The dogs behaved differently depending on the owners attention to them. When being watched by the owner, the dogs stayed lying down most often and/or for the longest time compared with when the owner read a book, watched TV, turned his or her back on them, or left the room. These results indicate that the dogs sensed the attentional state of their owners by judging observable behavioral cues such as eye contact and eye, head, and body orientation.

Technical intelligence in animals: The kea model.

The ability to act on information flexibly is one of the cornerstones of intelligent behavior. As particularly informative example, tool-oriented behavior has been investigated to determine to which extent nonhuman animals understand means–end relations, object affordances, and have specific motor skills. Even planning with foresight, goal-directed problem solving and immediate causal inference have been a focus of research. However, these cognitive abilities may not be restricted to tool-using animals but may be found also in animals that show high levels of curiosity, object exploration and manipulation, and extractive foraging behavior. The kea, a New Zealand parrot, is a particularly good example. We here review findings from laboratory experiments and field observations of keas revealing surprising cognitive capacities in the physical domain. In an experiment with captive keas, the success rate of individuals that were allowed to observe a trained conspecific was significantly higher than that of naive control subjects due to their acquisition of some functional understanding of the task through observation. In a further experiment using the string-pulling task, a well-probed test for means–end comprehension, we found the keas finding an immediate solution that could not be improved upon in nine further trials. We interpreted their performance as insightful in the sense of being sensitive of the relevant functional properties of the task and thereby producing a new adaptive response without trial-and-error learning. Together, these findings contribute to the ongoing debate on the distribution of higher cognitive skills in the animal kingdom by showing high levels of sensorimotor intelligence in animals that do not use tools. In conclusion, we suggest that the ‘Technical intelligence hypothesis’ (Byrne, Machiavellian intelligence II: extensions and evaluations, pp 289–211, 1997), which has been proposed to explain the origin of the ape/monkey grade-shift in intelligence by a selection pressure upon an increased efficiency in foraging behavior, should be extended, that is, applied to some birds as well.

Social learning in a non-social reptile (Geochelone carbonaria).

The ability to learn from the actions of another is adaptive, as it is a shortcut for acquiring new information. However, the evolutionary origins of this trait are still unclear. There is evidence that group-living mammals, birds, fishes and insects can learn through observation, but this has never been investigated in reptiles. Here, we show that the non-social red-footed tortoise (Geochelone carbonaria) can learn from the actions of a conspecific in a detour task; non-observer animals (without a conspecific demonstrator) failed. This result provides the first evidence that a non-social species can use social cues to solve a task that it cannot solve through individual learning, challenging the idea that social learning is an adaptation for social living.

Social learning affects object exploration and manipulation in keas, Nestor notabilis

Both the pattern of exploration and the manipulation efficiency of young keas attempting to open a complex food container proved to be influenced by the observation of an experienced group member. Five individuals were allowed to observe a trained conspecific that iteratively demonstrated several techniques to open a large steel box. The lid of the box could be opened only after several locking devices had been dismantled: a bolt had to be poked out, a split pin had to be pulled, and a screw had to be twisted out. The observers’ initial manipulative actions were compared with those of five nao¨ve control subjects (nonobservers). Although the observers failed to open the box completely and thus to get the reward in their first attempts, they explored more, approached the locking devices sooner and were more successful at opening them. Although their initial attempts did not match the response topography or the sequence of the model’s actions (movement or sequence imitation), their improved efficiency at unlocking the devices seemed to reflect the acquisition of some functional understanding of the task through observation (emulation learning).  2001 The Association for the Study of Animal Behaviour

Imitation as faithful copying of a novel technique in marmoset monkeys.

Imitative learning has received great attention due to its supposed role in the development of culture and the cognitive demands it poses on the individual. Evidence for imitation in non-human primate species, therefore, could shed light on the early origins of proto-cultural traits in the primate order. Imitation has been defined as the learning of an act by seeing it done or, more specifically, as the copying of a novel or otherwise improbable act. But despite a century of research and the detection of mirror neurons the empirical basis for this most advanced form of observational learning is weak. Few, if any, studies have shown that the observer has learned the response topography, i.e., the specific action by which the response is made. In an experimental set-up we confronted marmoset monkeys (Callithrix jacchus) with a conspecific model that was previously trained to open a plastic box in a peculiar way. Employing detailed motion analyses we show that the observers precisely copied the movement patterns of the novel action demonstrated by the model. A discriminant analysis classified 13 out of 14 observer movements (92.86%) as model movements and only one as non-observer movement. This evidence of imitation in non-human primates questions the dominant opinion that imitation is a human-specific ability. Furthermore, the high matching degree suggests that marmosets possess the neuronal mechanism to code the actions of others and to map them onto their own motor repertoire, rather than priming existing motor-templates.

Dogs Can Discriminate Emotional Expressions of Human Faces

The question of whether animals have emotions and respond to the emotional expressions of others has become a focus of research in the last decade [1-9]. However, to date, no study has convincingly shown that animals discriminate between emotional expressions of heterospecifics, excluding the possibility that they respond to simple cues. Here, we show that dogs use the emotion of a heterospecific as a discriminative cue. After learning to discriminate between happy and angry human faces in 15 picture pairs, whereby for one group only the upper halves of the faces were shown and for the other group only the lower halves of the faces were shown, dogs were tested with four types of probe trials: (1) the same half of the faces as in the training but of novel faces, (2) the other half of the faces used in training, (3) the other half of novel faces, and (4) the left half of the faces used in training. We found that dogs for which the happy faces were rewarded learned the discrimination more quickly than dogs for which the angry faces were rewarded. This would be predicted if the dogs recognized an angry face as an aversive stimulus. Furthermore, the dogs performed significantly above chance level in all four probe conditions and thus transferred the training contingency to novel stimuli that shared with the training set only the emotional expression as a distinguishing feature. We conclude that the dogs used their memories of real emotional human faces to accomplish the discrimination task.

Flexibility in Problem Solving and Tool Use of Kea and New Caledonian Crows in a Multi Access Box Paradigm

Parrots and corvids show outstanding innovative and flexible behaviour. In particular, kea and New Caledonian crows are often singled out as being exceptionally sophisticated in physical cognition, so that comparing them in this respect is particularly interesting. However, comparing cognitive mechanisms among species requires consideration of non-cognitive behavioural propensities and morphological characteristics evolved from different ancestry and adapted to fit different ecological niches. We used a novel experimental approach based on a Multi-Access-Box (MAB). Food could be extracted by four different techniques, two of them involving tools. Initially all four options were available to the subjects. Once they reached criterion for mastering one option, this task was blocked, until the subjects became proficient in another solution. The exploratory behaviour differed considerably. Only one (of six) kea and one (of five) NCC mastered all four options, including a first report of innovative stick tool use in kea. The crows were more efficient in using the stick tool, the kea the ball tool. The kea were haptically more explorative than the NCC, discovered two or three solutions within the first ten trials (against a mean of 0.75 discoveries by the crows) and switched more quickly to new solutions when the previous one was blocked. Differences in exploration technique, neophobia and object manipulation are likely to explain differential performance across the set of tasks. Our study further underlines the need to use a diversity of tasks when comparing cognitive traits between members of different species. Extension of a similar method to other taxa could help developing a comparative cognition research program.

The evolution of imitation: what do the capacities of non-human animals tell us about the mechanisms of imitation?

In this paper, we review reports and present new empirical data from studies with marmosets and dogs that address the correspondence problem of imitation research. We focus on the question of how it is possible to transform visual information into matching motor acts. Here, the important issue is not the learning of a complex skill, but determining the copying fidelity of animals at different levels of behavioural organization. As a theoretical framework, we suggest a classification in terms of movement, action and result, which shows a positive relationship between the organizational level of imitation and matching degree. While the monkey studies have provided evidence of very precise copying of movements and, to a lesser degree, of behaviours, the dog studies have provided evidence of action copying and the reproduction of results. In a Do-as-I-do study, a dog attempted to reproduce the results of demonstrated object manipulations at the expense of movement details. Transitive actions were more easily replicated than intransitive ones, and familiarity of actions had a major influence. The discussion of these findings addresses the question of the neuronal mechanisms underlying imitation and whether a single mechanism is sufficient to explain the different levels of copying fidelity.