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Ludwig Huber

University of Veterinary Medicine Vienna, Messerli Research Institute, Faculty Member

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Ludwig Huber is Professor of the Natural Science Foundations of Animal Ethics and Human-Animal Interactions at the Messerli Research Institute, an Interdisciplinary Institute of the University of Veterinary Medicine Vienna, the Medical University of Vienna, and the University of Vienna. There he is the head of the Division of Comparative Cognition, which includes the Clever Dog Lab Vienna, the Research Station on Cognition and Communication (Haidlhof) and is linked to the Wolf Science Center (Ernstbrunn). His research focuses on animal cognition in a broad, comparative manner, including such diverse species as humans, marmosets, dogs, kea, pigeons, red-footed tortoise, lizards, poison frogs and archer fish. He has written more than 100 research articles and book chapters about these topics, and is the co-editor of several books including The Evolution of Cognition (MIT Press 2000).

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Papers by Ludwig Huber

Canine perspective-taking

Animal Cognition

An important question in the study of canine cognition is how dogs understand humans, given that ... more An important question in the study of canine cognition is how dogs understand humans, given that they show impressive abilities for interacting and communicating with us. In this review, we describe and discuss studies that have investigated dogs’ perspective-taking abilities. There is solid evidence that dogs are not only sensitive to the gaze of others, but also their attention. We specifically address the question whether dogs have the ability to take the perspective of others and thus come to understand what others can or cannot perceive. From the latter, they may then infer what others know and use this representation to anticipate what others do next. Still, dogs might simply rely on directly observable cues and on what they themselves can perceive when they assess what others can perceive. And instead of making inferences from representations of others' mental states, they may have just learned that certain behaviours of ours lead to certain outcomes. However, recent rese...

Dogs follow human misleading suggestions more often when the informant has a false belief

Proceedings of the Royal Society B: Biological Sciences

We investigated whether dogs ( Canis familiaris ) distinguish between human true (TB) and false b... more We investigated whether dogs ( Canis familiaris ) distinguish between human true (TB) and false beliefs (FB). In three experiments with a pre-registered change of location task, dogs ( n = 260) could retrieve food from one of two opaque buckets after witnessing a misleading suggestion by a human informant (the ‘communicator’) who held either a TB or a FB about the location of food. Dogs in both the TB and FB group witnessed the initial hiding of food, its subsequent displacement by a second experimenter, and finally, the misleading suggestion to the empty bucket by the communicator. On average, dogs chose the suggested container significantly more often in the FB group than in the TB group and hence were sensitive to the experimental manipulation. Terriers were the only group of breeds that behaved like human infants and apes tested in previous studies with a similar paradigm, by following the communicator's suggestion more often in the TB than in the FB group. We discuss the re...

Vocal development in nestling kea parrots (Nestor notabilis)

Bioacoustics, 2020

This study investigates vocal development of nestling kea parrots (Nestor notabilis). First, we e... more This study investigates vocal development of nestling kea parrots (Nestor notabilis). First, we examine how many structurally distinct call types were present during the nestling period, and the age in which call types occurred. Based on studies with other avian species, we predicted that kea nestlings would have multiple call types, with some present at hatching and others emerging later in the nestling period. Results showed that nestlings have four distinct call types, two present at hatching, and two emerging when the nestlings undergo large physical changes in the second week of life. While two of the call types developed gradually towards more adult-like structures, the other two did not and were apparently only used for communication in the nest. Second, we tested whether nestlings could be discriminated individually based on their calls. All four call types were individually discriminable from hatching until the end of the study, thus providing evidence for vocal signatures. While the function of vocal signatures cannot be ascertained from this study, kea breeding biology rules out explanations based on preventing misdirected parental care. We suggest that vocal signatures may be relevant to the division of parental resources in the nest.

CASE STUDY 14A Movement Imitation in Monkeys

Intelligence, Culture, and Individualized Societies

Empathy in dogs: With a little help from a friend – a mixed blessing

Animal Sentience, 2017

Kujala (2017) presents an extensive overview of existing research on canine emotions in compariso... more Kujala (2017) presents an extensive overview of existing research on canine emotions in comparison to those of other non-human animals and humans. This commentary provides some additional research results on the intensively debated field of empathy in dogs. We focus on recent advances in the understanding of a fundamental building block of empathy-emotional contagion-and on dogs' remarkable sensitivity for human emotions, including the skills of assistance dogs.

Expectancy Violations about Physical Properties of Animated Objects in Dogs

Dogs are not particularly known for complex physical cognitive abilities. However, a number of re... more Dogs are not particularly known for complex physical cognitive abilities. However, a number of recent violation-of-expectation studies have challenged this view. In the current eye-tracking study, we further investigated dogs’ (N=15) reaction to physically implausible events, particularly in the context of support, occlusion, and launching events. In Experiment 1, the dogs watched a rolling ball moving over a gap in a surface either falling down or hovering over the gap. In Experiment 2, the dogs saw a ball rolling behind a narrow pole either disappearing behind it or re-appearing on the other side. In Experiment 3, the dogs observed launching events either with or without contact between the balls. The dogs’ pupil dilation response and looking times suggest that they form implicit expectations about occlusion and launching events but not about gravity-related events at least in the context of animated objects on a screen.

Overimitation in Dogs: Is There a Link to the Quality of the Relationship with the Caregiver?

Animals, 2022

Overimitation, the copying of causally irrelevant or non-functional actions, is well-known from h... more Overimitation, the copying of causally irrelevant or non-functional actions, is well-known from humans but completely absent in other primates. Recent studies from our lab have provided evidence for overimitation in canines. Previously, we found that half of tested pet dogs copied their human caregiver’s irrelevant action, while only few did so when the action was demonstrated by an unfamiliar experimenter. Therefore, we hypothesized that dogs show overimitation as a result of socio-motivational grounds. To test this more specifically, here we investigated how the relationship with the caregiver influenced the eagerness to overimitate. Given the high variability in the tendency to overimitate their caregiver, we hypothesized that not only familiarity but also relationship quality influences whether dogs faithfully copy their caregiver. For this purpose, on the one hand we measured the overimitation tendency (with the same test as in the two studies before) and on the other hand the ...

1 The Evolutionary Origins of Consciousness Comments to John F. Kihlstrom’s “Revising Scientific Models of Unconscious Mental Activity”

What is consciousness? The conference is about consciousness, one of the most fascinating but als... more What is consciousness? The conference is about consciousness, one of the most fascinating but also most problematic phenomena studied by contemporary empirical science. While be attractive enough for philosophers to produce an endless stream of hot debates, empirical scientists have long hesitated or refused to consider consciousness a valid or fruitful scientific problem. Even worse, many people would be happy to see the workings of the human mind not being explained away, because it touches so fundamentally the way we see ourselves in the universe, shaking the hope for our own personal survival, which many see very much in terms of the continuity of their own consciousness. All that may end up in curiously sardonic definitions of consciousness, like “Consciousness is a fascinating but elusive phenomenon; it is impossible to specify what it is, what it does, or why it evolved. Nothing worth reading has been written about it ” (Sutherland 1989), or, even more cynical, “Nobody has th...

Kea, Nestor notabilis, achieve cooperation in dyads, triads, and tetrads when dominants show restraint

Learning & Behavior, 2021

Animal cooperation in the wild often involves multiple individuals that must tolerate each other ... more Animal cooperation in the wild often involves multiple individuals that must tolerate each other in close proximity. However, most cooperation experiments in the lab are done with two animals, that are often also physically separated. Such experiments are useful for answering some pertinent questions, for example about the understanding of the role of the partner and strategies of partner control, but say little about factors determining successful cooperation with multiple partners in group settings. We explored the influence of dominance, rank distance, tolerance, affiliation, and coordination by testing kea parrots with a box requiring two, three, or four chains to be pulled simultaneously to access food rewards. The reward could be divided unevenly, but not monopolized completely. Eventually dyadic, triadic, and tetradic cooperation tasks were solved, showing that non-human animals are capable of tetradic cooperation in an experimental setup. Starting with two chains, we found t...

Do Nonhumans Seek Explanations?

Animal Behavior and Cognition, 2020

From an early age, children explore their environment in a way suggesting that they reason about ... more From an early age, children explore their environment in a way suggesting that they reason about causal variables and seek causal explanations. Indeed, following extensive studies of problem-solving abilities in chimpanzees, Povinelli (Folk Physics for Apes, Oxford University Press, 2000) proposed that this ability to reason about unobservable variables is unique to humans. Following on from this, Povinelli and Dunphy-Lelii (Canadian Journal of Experimental Psychology, 55(2), 187–195, 2001) addressed the question whether chimpanzees would explore objects with the aim of elucidating unobservable and surprising object properties. Chimpanzees, unlike preschool children, did not show increased object exploration following a change in the unobservable properties of an object. We critically discuss these findings and argue that more research using a greater variety of methods and with a larger number of species is required to support the hypothesis that only humans engage in explanation s...

Extending the Reach of Tooling Theory: A Neurocognitive and Phylogenetic Perspective

Topics in Cognitive Science, 2021

Tool use research has suffered from a lack of consistent theoretical frameworks. There is a pleth... more Tool use research has suffered from a lack of consistent theoretical frameworks. There is a plethora of tool use definitions and the most widespread ones are so inclusive that the behaviors that fall under them arguably do not have much in common. The situation is aggravated by the prevalence of anecdotes, which have played an undue role in the literature. In order to provide a more rigorous foundation for research and to advance our understanding of the interrelation between tool use and cognition, we suggest the adoption of Fragaszy and Mangalam's (2018) tooling framework, which is characterized by the creation of a body-plus-object system that manages a mechanical interface between tool and surface. Tooling is limited to a narrower suite of behaviors than tool use, which might facilitate its neurocognitive investigation. Indeed, evidence in the literature indicates that tooling has distinct neurocognitive underpinnings not shared by other activities typically classified as tool use, at least in primates. In order to understand the extent of tooling incidences in previous research, we systematically surveyed the comprehensive tool use catalog by . We identified 201 tool use submodes, of which only 81 could be classified as tooling, and the majority of the tool use examples across species were poorly supported by evidence. Furthermore

Towards a Methodology for Data-Driven Automatic Analysis of Animal Behavioral Patterns

Proceedings of the Seventh International Conference on Animal-Computer Interaction, 2020

Measurement of behavior a major challenge in many animalrelated disciplines, including ACI. This ... more Measurement of behavior a major challenge in many animalrelated disciplines, including ACI. This usually requires choosing specific parameters for measuring, related to the investigated hypothesis. Therefore, a key challenge is determining a priori what parameters are informational for a given experiment. The scope of this challenge is raised even further by the emerging computational approaches for animal detection and tracking, as automatizing behavioral measurement makes the possibilities for measuring behavioral parameters practically endless. This paper approaches these challenges by proposing a framework for guiding the decision making of researchers in their future data analysis. The framework is data-driven in the sense that it applies data mining techniques for obtaining insights from experimental data for guiding the choice of certain behavioral parameters. Here, we demonstrate the approach using a concrete example of clustering-based analysis of trajectories which can identify 'prevalent areas of stay' of the animal subjects in the experimental setting. CCS CONCEPTS • Human-centered computing → HCI theory, concepts and models; • Information systems → Data mining.

Experimental Tests for Measuring Individual Attentional Characteristics in Songbirds

Animals, 2021

Attention is defined as the ability to process selectively one aspect of the environment over oth... more Attention is defined as the ability to process selectively one aspect of the environment over others and is at the core of all cognitive processes such as learning, memorization, and categorization. Thus, evaluating and comparing attentional characteristics between individuals and according to situations is an important aspect of cognitive studies. Recent studies showed the interest of analyzing spontaneous attention in standardized situations, but data are still scarce, especially for songbirds. The present study adapted three tests of attention (towards visual non-social, visual social, and auditory stimuli) as tools for future comparative research in the European starling (Sturnus vulgaris), a species that is well known to present individual variations in social learning or engagement. Our results reveal that attentional characteristics (glances versus gazes) vary according to the stimulus broadcasted: more gazes towards unusual visual stimuli and species-specific auditory stimul...

Are free-ranging Kune Kune pigs (Sus scrofa domesticus) able to solve a cooperative task?

Applied Animal Behaviour Science, 2021

Cooperative behaviour has been observed in many species and in several domains like cooperative h... more Cooperative behaviour has been observed in many species and in several domains like cooperative hunting, cooperative predator defence and cooperative breeding. From an evolutionary perspective, it is an efficient way to increase one's own success, which ultimately has positive fitness consequences. However, the underlying mechanism in terms of the motivational and cognitive processes is less clear. Pigs are cooperative breeders, highly sociable and possess advanced skills in various cognitive domains and sensory modalities. The aim of this study was to explore whether domestic pigs would cooperate to acquire resources. We tested this using the Joint-Log-Lift task, which requires that two pigs lift a wooden log simultaneously to obtain food. Ten same-sex dyads of free-ranging Kune Kune pigs were tested. They did not interact with the apparatus in at least one of a six-trial session and hence, did not solve the task spontaneously, but became successful after several training steps including both individual and partner training. Because the Joint-Log-Lift task requires behavioural coordination to be solved cooperatively and efficiently, we tested the pigs' understanding of the need for coordination in two classical tests. A 10 or 30 s delay condition tested whether the pig who was released earlier into the testing arena would wait for the partner before lifting the log. A visual barrier condition tested whether the inability to see the partner would reduce the synchronicity of their actions toward the log. The dyads did not respond differently in the two testing manipulations, therefore failing to show evidence for coordinated behaviour. We discussed these findings in terms of weak understanding of the cooperative requirements of the task, disadvantageous training and low impulse control. Further studies on the pigs' cognitive capabilities to adjust or coordinate their behaviour to conspecifics are important as they have potential implications for their welfare.

Exploring the dog–human relationship by combining fMRI, eye-tracking and behavioural measures

Scientific Reports, 2020

Behavioural studies revealed that the dog–human relationship resembles the human mother–child bon... more Behavioural studies revealed that the dog–human relationship resembles the human mother–child bond, but the underlying mechanisms remain unclear. Here, we report the results of a multi-method approach combining fMRI (N = 17), eye-tracking (N = 15), and behavioural preference tests (N = 24) to explore the engagement of an attachment-like system in dogs seeing human faces. We presented morph videos of the caregiver, a familiar person, and a stranger showing either happy or angry facial expressions. Regardless of emotion, viewing the caregiver activated brain regions associated with emotion and attachment processing in humans. In contrast, the stranger elicited activation mainly in brain regions related to visual and motor processing, and the familiar person relatively weak activations overall. While the majority of happy stimuli led to increased activation of the caudate nucleus associated with reward processing, angry stimuli led to activations in limbic regions. Both the eye-trackin...

How Dogs Perceive Humans and How Humans Should Treat Their Pet Dogs: Linking Cognition With Ethics

Frontiers in Psychology, 2020

Humans interact with animals in numerous ways and on numerous levels. We are indeed living in an ... more Humans interact with animals in numerous ways and on numerous levels. We are indeed living in an “animal”s world,’ in the sense that our lives are very much intertwined with the lives of animals. This also means that animals, like those dogs we commonly refer to as our pets, are living in a “human’s world” in the sense that it is us, not them, who, to a large degree, define and manage the interactions we have with them. In this sense, the human-animal relationship is nothing we should romanticize: it comes with clear power relations and thus with a set of responsibilities on the side of those who exercise this power. This holds, despite the fact that we like to think about our dogs as human’s best friend. Dogs have been part of human societies for longer than any other domestic species. Like no other species they exemplify the role of companion animals. Relationships with pet dogs are both very widespread and very intense, often leading to strong attachments between owners or caregi...

Partial rewarding during clicker training does not improve naïve dogs’ learning speed and induces a pessimistic-like affective state

Animal Cognition, 2020

Clicker training is considered a welfare-friendly way of teaching novel behaviors to animals beca... more Clicker training is considered a welfare-friendly way of teaching novel behaviors to animals because it is mostly based on the positive reinforcement. However, trainers largely vary in their way of applying this training technique. According to the most, a reward (e.g., food) should follow every click, while others claim that dogs learn faster when the reward is sometimes omitted. One argument against the use of partial rewarding is that it induces frustration in the animal, raising concerns over its welfare consequences. Here, we investigated the effect of partial rewarding not only on training efficacy (learning speed), but also on dogs’ affective state. We clicker-trained two groups of dogs: one group received food after every click while the other group received food only 60% of the time. Considering previous evidence of the influencing role of personality on reactions to frustrated expectations, we included measurements of dogs’ emotional reactivity. We compared the number of t...

Do puffins use tools?

Proceedings of the National Academy of Sciences, 2020

Tailored haemodynamic response function increases detection power of fMRI in awake dogs (Canis familiaris)

Functional magnetic resonance imaging (fMRI) of awake and unrestrained dogs (Canis familiaris) ha... more Functional magnetic resonance imaging (fMRI) of awake and unrestrained dogs (Canis familiaris) has been established as a novel opportunity for comparative neuroimaging, promising important insights into the evolutionary roots of human brain function and cognition. However, data processing and analysis pipelines are often derivatives of methodological standards developed for human neuroimaging, which may be problematic due to profound neurophysiological and anatomical differences between humans and dogs. Here, we explore whether dog fMRI studies would benefit from a tailored dog haemodynamic response function (HRF). In two independent experiments, dogs were presented with different visual stimuli. BOLD signal changes in the visual cortex during these experiments were used for (a) the identification and estimation of a tailored dog HRF, and (b) the independent validation of the resulting dog HRF estimate. Time course analyses revealed that the BOLD signal in the primary visual cortex ...

Selective overimitation in dogs

Learning & Behavior, 2020

Dogs have not only shown different kinds of social learning, from either conspecifics or humans, ... more Dogs have not only shown different kinds of social learning, from either conspecifics or humans, including do-as-I-do imitation, deferred imitation, and selective imitation, but in two previous studies they have also shown an eagerness to copy causally irrelevant actions. This so-called overimitation is prevalent in humans but is totally absent in great apes. Whereas in one of two previous studies dogs copied actions from an experimenter (Johnston, Holden, & Santos in Developmental Science, 20, e12460, 2017), in the other a reasonable number of the dogs copied the irrelevant actions from their human caregiver (Huber, Popovová, Riener, Salobir, & Cimarelli in Learning & Behavior, 46, 387-397, 2018). Dogs have not only been domesticated to live and work with us, but many companion dogs develop strong affiliative relationships with their caregiver, which are akin to the attachment bonds between human children and their mother. We therefore assumed that overimitation in dogs might be strongly motivated by social factors, such as affiliation or conformity. To test this hypothesis, we confronted dogs with the same demonstration of causally relevant and irrelevant actions as in the previous study (Huber et al. in Learning & Behavior, 46, 387-397, 2018), but this time with an unfamiliar experimenter instead of the caregiver as the demonstrator. The results strongly supported our hypothesis: Whereas half of the subjects in the previous study replicated the causally irrelevant action demonstrated by their caregiver, only very few did so when the actions were demonstrated by the experimenter. We conclude that the eagerness of dogs to learn from humans and to copy even unnecessary actions is strongly facilitated by their relationship with the particular human.

Canine perspective-taking

Animal Cognition

An important question in the study of canine cognition is how dogs understand humans, given that ... more An important question in the study of canine cognition is how dogs understand humans, given that they show impressive abilities for interacting and communicating with us. In this review, we describe and discuss studies that have investigated dogs’ perspective-taking abilities. There is solid evidence that dogs are not only sensitive to the gaze of others, but also their attention. We specifically address the question whether dogs have the ability to take the perspective of others and thus come to understand what others can or cannot perceive. From the latter, they may then infer what others know and use this representation to anticipate what others do next. Still, dogs might simply rely on directly observable cues and on what they themselves can perceive when they assess what others can perceive. And instead of making inferences from representations of others' mental states, they may have just learned that certain behaviours of ours lead to certain outcomes. However, recent rese...

Dogs follow human misleading suggestions more often when the informant has a false belief

Proceedings of the Royal Society B: Biological Sciences

We investigated whether dogs ( Canis familiaris ) distinguish between human true (TB) and false b... more We investigated whether dogs ( Canis familiaris ) distinguish between human true (TB) and false beliefs (FB). In three experiments with a pre-registered change of location task, dogs ( n = 260) could retrieve food from one of two opaque buckets after witnessing a misleading suggestion by a human informant (the ‘communicator’) who held either a TB or a FB about the location of food. Dogs in both the TB and FB group witnessed the initial hiding of food, its subsequent displacement by a second experimenter, and finally, the misleading suggestion to the empty bucket by the communicator. On average, dogs chose the suggested container significantly more often in the FB group than in the TB group and hence were sensitive to the experimental manipulation. Terriers were the only group of breeds that behaved like human infants and apes tested in previous studies with a similar paradigm, by following the communicator's suggestion more often in the TB than in the FB group. We discuss the re...

Vocal development in nestling kea parrots (Nestor notabilis)

Bioacoustics, 2020

This study investigates vocal development of nestling kea parrots (Nestor notabilis). First, we e... more This study investigates vocal development of nestling kea parrots (Nestor notabilis). First, we examine how many structurally distinct call types were present during the nestling period, and the age in which call types occurred. Based on studies with other avian species, we predicted that kea nestlings would have multiple call types, with some present at hatching and others emerging later in the nestling period. Results showed that nestlings have four distinct call types, two present at hatching, and two emerging when the nestlings undergo large physical changes in the second week of life. While two of the call types developed gradually towards more adult-like structures, the other two did not and were apparently only used for communication in the nest. Second, we tested whether nestlings could be discriminated individually based on their calls. All four call types were individually discriminable from hatching until the end of the study, thus providing evidence for vocal signatures. While the function of vocal signatures cannot be ascertained from this study, kea breeding biology rules out explanations based on preventing misdirected parental care. We suggest that vocal signatures may be relevant to the division of parental resources in the nest.

CASE STUDY 14A Movement Imitation in Monkeys

Intelligence, Culture, and Individualized Societies

Empathy in dogs: With a little help from a friend – a mixed blessing

Animal Sentience, 2017

Kujala (2017) presents an extensive overview of existing research on canine emotions in compariso... more Kujala (2017) presents an extensive overview of existing research on canine emotions in comparison to those of other non-human animals and humans. This commentary provides some additional research results on the intensively debated field of empathy in dogs. We focus on recent advances in the understanding of a fundamental building block of empathy-emotional contagion-and on dogs' remarkable sensitivity for human emotions, including the skills of assistance dogs.

Expectancy Violations about Physical Properties of Animated Objects in Dogs

Dogs are not particularly known for complex physical cognitive abilities. However, a number of re... more Dogs are not particularly known for complex physical cognitive abilities. However, a number of recent violation-of-expectation studies have challenged this view. In the current eye-tracking study, we further investigated dogs’ (N=15) reaction to physically implausible events, particularly in the context of support, occlusion, and launching events. In Experiment 1, the dogs watched a rolling ball moving over a gap in a surface either falling down or hovering over the gap. In Experiment 2, the dogs saw a ball rolling behind a narrow pole either disappearing behind it or re-appearing on the other side. In Experiment 3, the dogs observed launching events either with or without contact between the balls. The dogs’ pupil dilation response and looking times suggest that they form implicit expectations about occlusion and launching events but not about gravity-related events at least in the context of animated objects on a screen.

Overimitation in Dogs: Is There a Link to the Quality of the Relationship with the Caregiver?

Animals, 2022

Overimitation, the copying of causally irrelevant or non-functional actions, is well-known from h... more Overimitation, the copying of causally irrelevant or non-functional actions, is well-known from humans but completely absent in other primates. Recent studies from our lab have provided evidence for overimitation in canines. Previously, we found that half of tested pet dogs copied their human caregiver’s irrelevant action, while only few did so when the action was demonstrated by an unfamiliar experimenter. Therefore, we hypothesized that dogs show overimitation as a result of socio-motivational grounds. To test this more specifically, here we investigated how the relationship with the caregiver influenced the eagerness to overimitate. Given the high variability in the tendency to overimitate their caregiver, we hypothesized that not only familiarity but also relationship quality influences whether dogs faithfully copy their caregiver. For this purpose, on the one hand we measured the overimitation tendency (with the same test as in the two studies before) and on the other hand the ...

1 The Evolutionary Origins of Consciousness Comments to John F. Kihlstrom’s “Revising Scientific Models of Unconscious Mental Activity”

What is consciousness? The conference is about consciousness, one of the most fascinating but als... more What is consciousness? The conference is about consciousness, one of the most fascinating but also most problematic phenomena studied by contemporary empirical science. While be attractive enough for philosophers to produce an endless stream of hot debates, empirical scientists have long hesitated or refused to consider consciousness a valid or fruitful scientific problem. Even worse, many people would be happy to see the workings of the human mind not being explained away, because it touches so fundamentally the way we see ourselves in the universe, shaking the hope for our own personal survival, which many see very much in terms of the continuity of their own consciousness. All that may end up in curiously sardonic definitions of consciousness, like “Consciousness is a fascinating but elusive phenomenon; it is impossible to specify what it is, what it does, or why it evolved. Nothing worth reading has been written about it ” (Sutherland 1989), or, even more cynical, “Nobody has th...

Kea, Nestor notabilis, achieve cooperation in dyads, triads, and tetrads when dominants show restraint

Learning & Behavior, 2021

Animal cooperation in the wild often involves multiple individuals that must tolerate each other ... more Animal cooperation in the wild often involves multiple individuals that must tolerate each other in close proximity. However, most cooperation experiments in the lab are done with two animals, that are often also physically separated. Such experiments are useful for answering some pertinent questions, for example about the understanding of the role of the partner and strategies of partner control, but say little about factors determining successful cooperation with multiple partners in group settings. We explored the influence of dominance, rank distance, tolerance, affiliation, and coordination by testing kea parrots with a box requiring two, three, or four chains to be pulled simultaneously to access food rewards. The reward could be divided unevenly, but not monopolized completely. Eventually dyadic, triadic, and tetradic cooperation tasks were solved, showing that non-human animals are capable of tetradic cooperation in an experimental setup. Starting with two chains, we found t...

Do Nonhumans Seek Explanations?

Animal Behavior and Cognition, 2020

From an early age, children explore their environment in a way suggesting that they reason about ... more From an early age, children explore their environment in a way suggesting that they reason about causal variables and seek causal explanations. Indeed, following extensive studies of problem-solving abilities in chimpanzees, Povinelli (Folk Physics for Apes, Oxford University Press, 2000) proposed that this ability to reason about unobservable variables is unique to humans. Following on from this, Povinelli and Dunphy-Lelii (Canadian Journal of Experimental Psychology, 55(2), 187–195, 2001) addressed the question whether chimpanzees would explore objects with the aim of elucidating unobservable and surprising object properties. Chimpanzees, unlike preschool children, did not show increased object exploration following a change in the unobservable properties of an object. We critically discuss these findings and argue that more research using a greater variety of methods and with a larger number of species is required to support the hypothesis that only humans engage in explanation s...

Extending the Reach of Tooling Theory: A Neurocognitive and Phylogenetic Perspective

Topics in Cognitive Science, 2021

Tool use research has suffered from a lack of consistent theoretical frameworks. There is a pleth... more Tool use research has suffered from a lack of consistent theoretical frameworks. There is a plethora of tool use definitions and the most widespread ones are so inclusive that the behaviors that fall under them arguably do not have much in common. The situation is aggravated by the prevalence of anecdotes, which have played an undue role in the literature. In order to provide a more rigorous foundation for research and to advance our understanding of the interrelation between tool use and cognition, we suggest the adoption of Fragaszy and Mangalam's (2018) tooling framework, which is characterized by the creation of a body-plus-object system that manages a mechanical interface between tool and surface. Tooling is limited to a narrower suite of behaviors than tool use, which might facilitate its neurocognitive investigation. Indeed, evidence in the literature indicates that tooling has distinct neurocognitive underpinnings not shared by other activities typically classified as tool use, at least in primates. In order to understand the extent of tooling incidences in previous research, we systematically surveyed the comprehensive tool use catalog by . We identified 201 tool use submodes, of which only 81 could be classified as tooling, and the majority of the tool use examples across species were poorly supported by evidence. Furthermore

Towards a Methodology for Data-Driven Automatic Analysis of Animal Behavioral Patterns

Proceedings of the Seventh International Conference on Animal-Computer Interaction, 2020

Measurement of behavior a major challenge in many animalrelated disciplines, including ACI. This ... more Measurement of behavior a major challenge in many animalrelated disciplines, including ACI. This usually requires choosing specific parameters for measuring, related to the investigated hypothesis. Therefore, a key challenge is determining a priori what parameters are informational for a given experiment. The scope of this challenge is raised even further by the emerging computational approaches for animal detection and tracking, as automatizing behavioral measurement makes the possibilities for measuring behavioral parameters practically endless. This paper approaches these challenges by proposing a framework for guiding the decision making of researchers in their future data analysis. The framework is data-driven in the sense that it applies data mining techniques for obtaining insights from experimental data for guiding the choice of certain behavioral parameters. Here, we demonstrate the approach using a concrete example of clustering-based analysis of trajectories which can identify 'prevalent areas of stay' of the animal subjects in the experimental setting. CCS CONCEPTS • Human-centered computing → HCI theory, concepts and models; • Information systems → Data mining.

Experimental Tests for Measuring Individual Attentional Characteristics in Songbirds

Animals, 2021

Attention is defined as the ability to process selectively one aspect of the environment over oth... more Attention is defined as the ability to process selectively one aspect of the environment over others and is at the core of all cognitive processes such as learning, memorization, and categorization. Thus, evaluating and comparing attentional characteristics between individuals and according to situations is an important aspect of cognitive studies. Recent studies showed the interest of analyzing spontaneous attention in standardized situations, but data are still scarce, especially for songbirds. The present study adapted three tests of attention (towards visual non-social, visual social, and auditory stimuli) as tools for future comparative research in the European starling (Sturnus vulgaris), a species that is well known to present individual variations in social learning or engagement. Our results reveal that attentional characteristics (glances versus gazes) vary according to the stimulus broadcasted: more gazes towards unusual visual stimuli and species-specific auditory stimul...

Are free-ranging Kune Kune pigs (Sus scrofa domesticus) able to solve a cooperative task?

Applied Animal Behaviour Science, 2021

Cooperative behaviour has been observed in many species and in several domains like cooperative h... more Cooperative behaviour has been observed in many species and in several domains like cooperative hunting, cooperative predator defence and cooperative breeding. From an evolutionary perspective, it is an efficient way to increase one's own success, which ultimately has positive fitness consequences. However, the underlying mechanism in terms of the motivational and cognitive processes is less clear. Pigs are cooperative breeders, highly sociable and possess advanced skills in various cognitive domains and sensory modalities. The aim of this study was to explore whether domestic pigs would cooperate to acquire resources. We tested this using the Joint-Log-Lift task, which requires that two pigs lift a wooden log simultaneously to obtain food. Ten same-sex dyads of free-ranging Kune Kune pigs were tested. They did not interact with the apparatus in at least one of a six-trial session and hence, did not solve the task spontaneously, but became successful after several training steps including both individual and partner training. Because the Joint-Log-Lift task requires behavioural coordination to be solved cooperatively and efficiently, we tested the pigs' understanding of the need for coordination in two classical tests. A 10 or 30 s delay condition tested whether the pig who was released earlier into the testing arena would wait for the partner before lifting the log. A visual barrier condition tested whether the inability to see the partner would reduce the synchronicity of their actions toward the log. The dyads did not respond differently in the two testing manipulations, therefore failing to show evidence for coordinated behaviour. We discussed these findings in terms of weak understanding of the cooperative requirements of the task, disadvantageous training and low impulse control. Further studies on the pigs' cognitive capabilities to adjust or coordinate their behaviour to conspecifics are important as they have potential implications for their welfare.

Exploring the dog–human relationship by combining fMRI, eye-tracking and behavioural measures

Scientific Reports, 2020

Behavioural studies revealed that the dog–human relationship resembles the human mother–child bon... more Behavioural studies revealed that the dog–human relationship resembles the human mother–child bond, but the underlying mechanisms remain unclear. Here, we report the results of a multi-method approach combining fMRI (N = 17), eye-tracking (N = 15), and behavioural preference tests (N = 24) to explore the engagement of an attachment-like system in dogs seeing human faces. We presented morph videos of the caregiver, a familiar person, and a stranger showing either happy or angry facial expressions. Regardless of emotion, viewing the caregiver activated brain regions associated with emotion and attachment processing in humans. In contrast, the stranger elicited activation mainly in brain regions related to visual and motor processing, and the familiar person relatively weak activations overall. While the majority of happy stimuli led to increased activation of the caudate nucleus associated with reward processing, angry stimuli led to activations in limbic regions. Both the eye-trackin...

How Dogs Perceive Humans and How Humans Should Treat Their Pet Dogs: Linking Cognition With Ethics

Frontiers in Psychology, 2020

Humans interact with animals in numerous ways and on numerous levels. We are indeed living in an ... more Humans interact with animals in numerous ways and on numerous levels. We are indeed living in an “animal”s world,’ in the sense that our lives are very much intertwined with the lives of animals. This also means that animals, like those dogs we commonly refer to as our pets, are living in a “human’s world” in the sense that it is us, not them, who, to a large degree, define and manage the interactions we have with them. In this sense, the human-animal relationship is nothing we should romanticize: it comes with clear power relations and thus with a set of responsibilities on the side of those who exercise this power. This holds, despite the fact that we like to think about our dogs as human’s best friend. Dogs have been part of human societies for longer than any other domestic species. Like no other species they exemplify the role of companion animals. Relationships with pet dogs are both very widespread and very intense, often leading to strong attachments between owners or caregi...

Partial rewarding during clicker training does not improve naïve dogs’ learning speed and induces a pessimistic-like affective state

Animal Cognition, 2020

Clicker training is considered a welfare-friendly way of teaching novel behaviors to animals beca... more Clicker training is considered a welfare-friendly way of teaching novel behaviors to animals because it is mostly based on the positive reinforcement. However, trainers largely vary in their way of applying this training technique. According to the most, a reward (e.g., food) should follow every click, while others claim that dogs learn faster when the reward is sometimes omitted. One argument against the use of partial rewarding is that it induces frustration in the animal, raising concerns over its welfare consequences. Here, we investigated the effect of partial rewarding not only on training efficacy (learning speed), but also on dogs’ affective state. We clicker-trained two groups of dogs: one group received food after every click while the other group received food only 60% of the time. Considering previous evidence of the influencing role of personality on reactions to frustrated expectations, we included measurements of dogs’ emotional reactivity. We compared the number of t...

Do puffins use tools?

Proceedings of the National Academy of Sciences, 2020

Tailored haemodynamic response function increases detection power of fMRI in awake dogs (Canis familiaris)

Functional magnetic resonance imaging (fMRI) of awake and unrestrained dogs (Canis familiaris) ha... more Functional magnetic resonance imaging (fMRI) of awake and unrestrained dogs (Canis familiaris) has been established as a novel opportunity for comparative neuroimaging, promising important insights into the evolutionary roots of human brain function and cognition. However, data processing and analysis pipelines are often derivatives of methodological standards developed for human neuroimaging, which may be problematic due to profound neurophysiological and anatomical differences between humans and dogs. Here, we explore whether dog fMRI studies would benefit from a tailored dog haemodynamic response function (HRF). In two independent experiments, dogs were presented with different visual stimuli. BOLD signal changes in the visual cortex during these experiments were used for (a) the identification and estimation of a tailored dog HRF, and (b) the independent validation of the resulting dog HRF estimate. Time course analyses revealed that the BOLD signal in the primary visual cortex ...

Selective overimitation in dogs

Learning & Behavior, 2020

Dogs have not only shown different kinds of social learning, from either conspecifics or humans, ... more Dogs have not only shown different kinds of social learning, from either conspecifics or humans, including do-as-I-do imitation, deferred imitation, and selective imitation, but in two previous studies they have also shown an eagerness to copy causally irrelevant actions. This so-called overimitation is prevalent in humans but is totally absent in great apes. Whereas in one of two previous studies dogs copied actions from an experimenter (Johnston, Holden, & Santos in Developmental Science, 20, e12460, 2017), in the other a reasonable number of the dogs copied the irrelevant actions from their human caregiver (Huber, Popovová, Riener, Salobir, & Cimarelli in Learning & Behavior, 46, 387-397, 2018). Dogs have not only been domesticated to live and work with us, but many companion dogs develop strong affiliative relationships with their caregiver, which are akin to the attachment bonds between human children and their mother. We therefore assumed that overimitation in dogs might be strongly motivated by social factors, such as affiliation or conformity. To test this hypothesis, we confronted dogs with the same demonstration of causally relevant and irrelevant actions as in the previous study (Huber et al. in Learning & Behavior, 46, 387-397, 2018), but this time with an unfamiliar experimenter instead of the caregiver as the demonstrator. The results strongly supported our hypothesis: Whereas half of the subjects in the previous study replicated the causally irrelevant action demonstrated by their caregiver, only very few did so when the actions were demonstrated by the experimenter. We conclude that the eagerness of dogs to learn from humans and to copy even unnecessary actions is strongly facilitated by their relationship with the particular human.