Spatial perseveration error by alpacas (<Emphasis Type="Italic">Vicugna pacos</Emphasis>) in an A-not-B detour task

Spatial perseveration has been documented for domestic animals such as mules, donkeys, horses and dogs. However, evidence for this spatial cognition behavior among other domestic species is scarce. Alpacas have been domesticated for at least 7000 years yet their cognitive ability has not been officially reported. The present article used an A-not-B detour task to study the spatial problem-solving abilities of alpacas (Vicugna pacos) and to identify the perseveration errors, which refers to a tendency to maintain a learned route, despite having another available path. The study tested 51 alpacas, which had to pass through a gap at one end of a barrier in order to reach a reward. After one, two, three or four repeats (A trials), the gap was moved to the opposite end of the barrier (B trials). In contrast to what has been found in other domestic animals tested with the same task, the present study did not find clear evidence of spatial perseveration. Individuals’ performance in the subsequent B trials, following the change of gap location, suggests no error persistence in alpacas. Results suggest that alpacas are more flexible than other domestic animals tested with this same task, which has important implications in planning proper training for experimental designs or productive purposes. These results could contribute toward enhancing alpacas’ welfare and our understanding of their cognitive abilities.     

Minding the gap: spatial perseveration error in dogs

We investigated a combination of perseveration and detour behaviour in 50 domestic dogs (Canis familiaris). They were required to make a detour through a gap at one end of a straight barrier in order to reach a target. After one, two, three or four repeats, the gap was moved to the opposite end of the barrier, and the detour behaviour of the dogs was recorded. Although the dogs could solve simple detour tasks (80% correct in the first trial), they committed a perseveration error of following the previously learned route despite the clearly visible change in the location of the gap. This ‘misbehaviour’ occurred in 29 of 30 dogs after only two learning trials. They never reached a 100% correct performance level again even after four runs through the second gap location. The results suggest that dogs are reluctant to unlearn acquired spatial motor responses and reinforced navigation, which has important implications for experimental design, everyday dog training and our understanding of their mental capacities.     

Mule cognition: a case of hybrid vigour?

This study compares the behaviour of the mule (Equus asinus × Equus caballus) with that of its parent species to assess the effects of hybridization on cognition. Six mules, six ponies (E. caballus) and six donkeys (E. asinus) were given a two choice visual discrimination learning task. Each session consisted of 12 trials and pass level was reached when subjects chose the correct stimulus for at least 9 out of the 12 trials in three consecutive sessions. A record was made of how many pairs each subject learnt over 25 sessions. The mules’ performance was significantly better than that of either of the parent species (Kruskal-Wallis: H_x = 8.11, P = 0.017). They were also the only group to learn enough pairs to be able to show a successive reduction in the number of sessions required to reach criterion level. This study provides the first empirical evidence that the improved characteristics of mules may be extended from physical attributes to cognitive function.     

Use of human-given cues by domestic dogs (Canis familiaris) and horses (Equus caballus)

Sixteen domestic dogs (Canis familiaris) and four horses (Equus caballus) were tested for their ability to use human-given manual and facial cues in an object-choice task. Two of the four horses used touch as a cue and one horse successfully used pointing. The performance of the dogs was considerably better, with 12 subjects able to use pointing as a cue, 4 able to use head orientation and 2 able to use eye gaze alone. Group analysis showed that the dogs performed significantly better in all experimental conditions than during control trials. Dogs were able to use pointing cues even when the cuer’s body was closer to the incorrect object. Working gundogs with specialised training used pointing more successfully than pet dogs and gundog breeds performed better than non-gundog breeds. The results of this experiment suggest that animals’ use of human given communicative signals depends on cognitive ability, the evolutionary consequences of domestication and enculturation by humans within the individual’s lifetime. Received: 15 July 1999 / Accepted after revision: 10 January 2000     

Social mimetic behaviour and social anticipation in dogs: preliminary results

Learning contributes to the development of mutual mimicry in group mates. The aim of our study was to investigate whether dogs would initiate walking a detour if they were repeatedly exposed to the detouring behaviour of their owner. Eight dog owners were asked to modify their usual way of approaching their home at the end of their daily walks, namely, to make a short detour before the entrance. Owners performed the detour at least 180 times, over a period of 3–6 months. During the first 30 detours (trials 1–30) all dogs followed the owner on the new route. Between trials 151 and 180, four dogs started to walk the detour before the owner displayed any intention to walk in that direction in 50–93% of the cases. Further observations that were carried out on one dog showed that the initialisation of the detours manifested sooner if a second familiar person started to walk the detours. Interestingly, the dog persisted in initialising detours long after the owners stopped detouring. We describe the observed phenomenon in the framework of social anticipation that manifests when an animal learns the proper sequence of an act performed by another animal, so that it can (1) predict the action in this sequence, and (2) as a result start either a similar or a complementary action as a response. These observations suggest that the dogs' social anticipation ability contributes to behavioural synchronisation and cooperative processes between dog and owner. Electronic Publication     

Domestic horses (<Emphasis Type="Italic">Equus ferus caballus</Emphasis>) fail to intuitively reason about object properties like solidity and weight

From early infancy, humans reason about the external world in terms of identifiable, solid, cohesive objects persisting in space and time. This is one of the most fundamental human skills, which may be part of our innate conception of object properties. Although object permanence has been extensively studied across a variety of taxa, little is known about how non-human animals reason about other object properties. In this study, we therefore tested how domestic horses (Equus ferus caballus) intuitively reason about object properties like solidity and height, to locate hidden food. Horses were allowed to look for a food reward behind two opaque screens, only one of which had either the proper height or inclination to hide food rewards. Our results suggest that horses could not intuitively reason about physical object properties, but rather learned to select the screen with the proper height or inclination from the second set of 5 trials.     

Personality correlates of Benton's visual retention test

Bivariate correlations revealed that only L was highly significantly (negatively) correlated with the ability to retain and reproduce geometrical figures (Benton Visual Retention Test). Personality groups were generated using extreme cut-offs for E and N, as well as P and L. Extraverts produced similar error profiles to introverts. High trait N Ss tended to emit more errors of omission compared to low trait N persons, otherwise the performance profiles were not statistically different. Several E × N interactions emerged however, with stable introverts yielding significantly more correct reproductions and a lower error frequency than the other groups, with dysthymics (introverted neurotics) performing worst, the latter group further characterised by a greater number of perseveration and mislocation errors. High and low P-scoring Ss did not differ significantly in their ability to reproduce figures, with the exception of perseveration errors which were more common amongst P + Ss. L + individuals displayed inferior performance compared to L− Ss. An analysis of error categories revealed that L + emitted more distortion, left- and right-handed errors. There was no indication of any P × L interaction effect.     

Spatial memory of domestic dogs (Canis familiaris) for hidden objects in a detour task

Spatial memory of domestic dogs (Canis familiaris) for hidden objects was investigated via a visible displacement problem of object permanence with a detour paradigm. Experiment 1 showed that dogs were able to spontaneously locate a disappearing object in a detour situation. In Experiments 2 and 3, dead reckoning and allocentric spatial information were put in conflict. Results revealed that dogs simultaneously encoded both sources of information when they had to bypass an obstacle to locate a hidden object. Experiment 3 also revealed that, over the course of testing, dogs gradually learned to rely predominantly on allocentric cues when the detour involved several reorientations. The current study reveals that spatial memory of dogs for hidden objects in a detour task was guided by flexibility in processing spatial information. Dogs could simultaneously encode dead reckoning and allocentric information to locate a disappearing object and used them hierarchically according to the complexity of the detour they encountered in the environment.     

The effect of training and breed group on problem-solving behaviours in dogs

Dogs have become the focus of cognitive studies looking at both their physical and social problem-solving abilities (Bensky et al. in Adv Stud Behav, 45:209–387, 2013), but very little is known about the environmental and inherited factors that may affect these abilities. In the current study, we presented a manipulation task (a puzzle box) and a spatial task (the detour) to 128 dogs belonging to four different breed groups: Herding, Mastiff-like, Working and Retrievers (von Holdt et al. in Nature 464:898–902, 2010). Within each group, we tested highly trained and non-trained dogs. Results showed that trained dogs were faster at obtaining the reward in the detour task. In the manipulation task, trained dogs approached the apparatus sooner in the first familiarization trial, but no effect of breed emerged on this variable. Furthermore, regardless of breed, dogs in the trained group spent proportionally more time interacting with the apparatus and were more likely to succeed in the test trial than dogs in the non-trained group, whereas regardless of training, dogs in the working breed group were more likely to succeed than dogs in the retriever and herding breed groups (but not the mastiff-like group). Finally, trained dogs were less likely to look at a person than non-trained dogs during testing, but dogs in the herding group more likely to do so than dogs in the retriever and working but not the mastiff-like breed groups. Overall, results reveal a strong influence of training experience but less consistent differences between breed groups on different components thought to affect problem solving.     

Context specificity of inhibitory control in dogs

Across three experiments, we explored whether a dog’s capacity for inhibitory control is stable or variable across decision-making contexts. In the social task, dogs were first exposed to the reputations of a stingy experimenter that never shared food and a generous experimenter who always shared food. In subsequent test trials, dogs were required to avoid approaching the stingy experimenter when this individual offered (but withheld) a higher-value reward than the generous experimenter did. In the A-not-B task, dogs were required to inhibit searching for food in a previously rewarded location after witnessing the food being moved from this location to a novel hiding place. In the cylinder task, dogs were required to resist approaching visible food directly (because it was behind a transparent barrier), in favor of a detour reaching response. Overall, dogs exhibited inhibitory control in all three tasks. However, individual scores were not correlated between tasks, suggesting that context has a large effect on dogs’ behavior. This result mirrors studies of humans, which have highlighted intra-individual variation in inhibitory control as a function of the decision-making context. Lastly, we observed a correlation between a subject’s age and performance on the cylinder task, corroborating previous observations of age-related decline in dogs’ executive function.     

The effects of age,rank and neophobia on social learning in horses

Social learning is said to meet the demands of complex environments in which individuals compete over resources and cooperate to share resources. Horses (Equus caballus) were thought to lack social learning skills because they feed on homogenously distributed resources with few reasons for conflict. However, the horse’s social environment is complex, which raises the possibility that its capacity for social transfer of feeding behaviour has been underestimated. We conducted a social learning experiment using 30 socially kept horses of different ages. Five horses, one from each group, were chosen as demonstrators, and the remaining 25 horses were designated observers. Observers from each group were allowed to watch their group demonstrator opening a feeding apparatus. We found that young, low-ranking and more exploratory horses learned by observing older members of their own group, and the older the horse, the more slowly it appeared to learn. Social learning may be an adaptive specialisation to the social environment. Older animals may avoid the potential costs of acquiring complex and potentially disadvantageous feeding behaviours from younger group members. We argue that horses show social learning in the context of their social ecology and that research procedures must take such contexts into account. Misconceptions about the horse’s sociality may have hampered earlier studies.     

Detour behaviour in three species of birds: quails (Coturnix sp.), herring gulls (Larus cachinnans) and canaries (Serinus canaria)

Detour behaviour is the ability of an animal to reach a goal stimulus by moving round any interposed obstacle. It has been widely studied and has been proposed as a test of insight learning in several species of mammals, but few data are available in birds. A comparative study in three species of birds, belonging to different eco-ethological niches, allows a better understanding of the cognitive mechanism of such detour behaviour. Young quails (Coturnix sp.), herring gulls (Larus cachinnans) and canaries (Serinus canaria), 1 month old, 10–25 days old and 4–6 months old, respectively, were tested in a detour situation requiring them to abandon a clear view of a biologically interesting object (their own reflection in a mirror) in order to approach that object. Birds were placed in a closed corridor, at one end of which was a barrier through which the object was visible. Four different types of barrier were used: vertical bar, horizontal bar, grid and transparent. Two symmetrical apertures placed midline in the corridor allowed the birds to adopt routes passing around the barrier. After entering the apertures, birds could turn either right or left to re-establish social contact with the object in the absence of any local sensory cues emanating from it. Quails appeared able to solve the task, though their performance depended on the type of barrier used, which appeared to modulate their relative interest in approaching the object or in exploring the surroundings. Young herring gulls also showed excellent abilities to locate spatially the out-of-view object, except when the transparent barrier was used. Canaries, on the other hand, appeared completely unable to solve the detour task, whatever barrier was in use. It is suggested that these species differences can be accounted for in terms of adaptation to a terrestrial or aerial environment.     

Paired-associate learning by normal children and retardates with relevant redundant compound stimuli

During daily sessions four first-grade boys each played a two-choice marble dropping game. When one colored light was presented a plain token was dispensed as soon as S dropped a marble, and when another light was on a token with a hole in its center was delivered. During choice trials that were interspersed among single-color trials S had to choose one of the two different colored holes. At the end of each session S traded tokens with holes for pennies and then turned in the remaining plain tokens which were not exchanged for money. After S had developed a preference for the response that earned the tokens with holes (tokens that were backed up with pennies) E instructed S to try to get as many plain tokens (unexchangeable tokens) as he could. All Ss immediately switched to the response that earned unexchangeable tokens but as soon as E left they returned to choosing the backed up response. When the adult reentered, Ss again chose to work for unexchangeable tokens. Two different Es (male and female) served and the instructions of each were effective in controlling responding as long as the adult was present, except for one S who did not respond to one E, but did respond as described for the other.     

Horses (Equus caballus) discriminate body odour cues from conspecifics

Knowledge about social recognition and memory in animals can help us to determine appropriate management and husbandry techniques. In this study, we used a habituation–discrimination procedure to investigate the ability of horses (Equus caballus) to distinguish between the body odour samples of unfamiliar conspecifics. To pick up body odour, we rubbed material on the coat of horses and presented these unknown body odours to 16 different conspecifics of the same sex and similar age. The test consisted of two successive two-min presentations of a sample from one individual (e.g. individual ‘A’) and a simultaneous presentation of samples from individual ‘A’ and a novel individual (e.g. individual ‘B’) during a final third presentation. The results showed that horses, regardless of sex, decreased the time they spent investigating conspecific body odour across the initial two presentations—demonstrating habituation. In the final presentation, the results demonstrated successful discrimination of the previously experienced odour because horses investigated the novel olfactory sample (‘B’) significantly more than the pre-exposed sample (‘A’). Taken together, these findings suggest, for the first time, that horses are able to discriminate two stimuli derived from body odours of unfamiliar conspecifics over short period of time.     

Social information in equine movement gestalts

One model of signal evolution is based on the notion that behaviours become increasingly detached from their original biological functions to obtain a communicative value. Selection may not always favour the evolution of such transitions, for instance, if signalling is costly due to predators usurping signal production. Here, we collected inertial movement sensing data recorded from multiple locations in free-ranging horses (Equus caballus), which we subjected to a machine learning algorithm to extract kinematic gestalt profiles. This yielded surprisingly rich and multi-layered sets of information. In particular, we were able to discriminate identity, breed, sex and some personality traits from the overall movement patterns of freely moving subjects. Our study suggests that, by attending to movement gestalts, domestic horses, and probably many other group-living animals, have access to rich social information passively but reliably made available by conspecifics, a finding that we discuss in relation with current signal evolution theories.     

Flexible route selection by pigeons (Columba livia) on a computerized multi-goal navigation task with and without an "obstacle"

The ability to select efficient routes while reaching several locations, in situations addressed as the "traveling salesperson problem (TSP)", seems to play an important role in the lives of nonhuman species as well as humans. One of our previous studies with pigeons (Columba livia) used different variations of computerized navigation tasks to move a target to one or more goals, and showed that the birds consistently exhibited tendencies to visit the nearest goal first (Miyata & Fujita, 2010). Is this inflexible tendency consistent across all situations, or is it abandoned if it leads to a nonefficient strategy? The present study compared pigeons' route selection strategies in two-goal navigation tasks with and without having an L-shaped line as an obstacle barrier between the starting location of the target and the nearer goal. The pigeons frequently started by visiting the nearer goal in trials with no obstacles, whereas in trials having obstacles the birds often visited the farther goal first. For the detour trials with obstacles, the birds started from visiting the farther goal significantly more often than chance, even though disparity between the two traveling sequences was relatively small. The data demonstrated a case in which a tendency to visit the nearest goal was flexibly abandoned when the route required a detour behavior. Detailed analyses of the movement paths further suggest that the pigeons made decisions to choose a farther goal during the initial few steps, although a past history of reinforcement to avoid the barrier might have guided the birds' behavior.     

Spatial learning in pigs: effects of environmental enrichment and individual characteristics on behaviour and performance

This study investigated the effects of both environmental enrichment and individual behavioural characteristics on spatial cognitive capabilities of pigs, using a novel latent spatial learning paradigm based on Tolman’s detour experiments (1948). Pigs were housed either in ‘barren’ pens or in pens enriched with straw bedding from birth. Pigs were restrained in a Backtest at 10 and 17 days postpartum. Based on their escape behaviour in this test, which has been shown to reflect their behavioural style, six ‘high-resisting’(HR) and six ‘low-resisting’ (LR) pigs were selected from each housing environment (n = 24 in total). At 12 weeks of age, pairs of pen mates (LR and HR) were exposed to a maze three times (exploration trials). Pigs were then placed individually in the maze, and social reinstatement proved to be a strong incentive to find the exit leading to the home pen. We subsequently blocked the direct route to the exit, forcing animals to find a detour (memory test 1, MT1). This test was repeated once to investigate the relative improvement, i.e. detour learning (memory test 2, MT2). Housing condition and Backtest response strongly affected exploration patterns. In spite of this, no effects on performance during the subsequent memory tests were found. Performance was substantially improved in MT2, indicating that once a goal is apparent, pigs are able to solve a complex spatial memory task easily. In conclusion, social reinstatement provided a good incentive to complete a spatial task, and the substantial improvement in performance between MT1 and MT2 stresses the need for task complexity when testing spatial memory in pigs. Housing conditions or individual behavioural style did not affect spatial memory during MT1 or MT2. However, housing environment and behavioural style strongly affected explorative behaviour of pigs in an unfamiliar maze during both exploration trials and memory tests. This implicates that apparent effects of environmental enrichment on spatial learning and memory in pigs might reflect differences in explorative patterns rather than in cognitive processes.     

Horses (<Emphasis Type="Italic">Equus caballus</Emphasis>) use human local enhancement cues and adjust to human attention

This study evaluates the horse (Equus caballus) use of human local enhancement cues and reaction to human attention when making feeding decisions. The superior performance of dogs in observing human states of attention suggests this ability evolved with domestication. However, some species show an improved ability to read human cues through socialization and training. We observed 60 horses approach a bucket with feed in a three-way object-choice task when confronted with (a) an unfamiliar or (b) a familiar person in 4 different situations: (1) squatting behind the bucket, facing the horse (2) standing behind the bucket, facing the horse (3) standing behind the bucket in a back-turned position, gazing away from the horse and (4) standing a few meters from the bucket in a distant, back-turned position, again gazing away from the horse. Additionally, postures 1 and 2 were tested both with the person looking permanently at the horse and with the person alternating their gaze between the horse and the bucket. When the person remained behind the correct bucket, it was chosen significantly above chance. However, when the test person was turned and distant from the buckets, the horses’ performance deteriorated. In the turned person situations, the horses approached a familiar person and walked towards their focus of attention significantly more often than with an unfamiliar person. Additionally, in the squatting and standing person situations, some horses approached the person before approaching the correct bucket. This happened more with a familiar person. We therefore conclude that horses can use humans as a local enhancement cue independently of their body posture or gaze consistency when the persons remain close to the food source and that horses seem to orientate on the attention of familiar more than of unfamiliar persons. We suggest that socialization and training improve the ability of horses to read human cues.     

Selective information use and perseveration in the search behavior of infants and young children

Two experiments examined the early development of selective information use in search. The first experiment tested 9- and 16-month-olds on a modification of Piaget's Stage IV object permanence task. It examined infants' use of information from previous experiences with an object (prior information) and from the most recent hiding (current information) to locate a hidden object. In the second experiment, 2-, $\text{2}\text{1}\text{2}$-, and 4-year-old children received these same sources of information along with new forms of prior and current information: information about the typical locations of objects (location specificity) and verbal information. No systematic perseveration was observed at 9 months, although previous findings related to perseveration were replicated. Perseveration was found at 16 months, but there was also evidence of selectivity at that age. When errors occurred, they tended to be to the prior location, but they were infrequent in comparison to correct searches at the current location. The preschoolers, while continuing to show perseveration, were more consistently selective than the infants. They also showed considerable generality in extending their selectivity to new sources of information.     

Dogs learn to solve the support problem based on perceptual cues

Numerous recent studies have investigated how animals solve means-end tasks and unraveled considerable variation in strategies used by different species. Domestic dogs (Canis familiaris) have typically performed comparably poorly in physical cognition tasks, but a recent study showed that they can solve the on–off condition of the support problem, where they are confronted with two boards, one with a reward placed on it and the other with a reward placed next to it. To explore which strategies dogs use to solve this task, we first tested 37 dogs with the on–off condition tested previously and then tested subjects that passed this condition with three transfer tasks. For the contact condition, the inaccessible reward was touching the second board. For the perceptual containment condition, the inaccessible reward was surrounded on three sides by the second board, but not supported by it, whereas for the gap condition, discontinuous boards were used. Unlike in the previous study, our subjects did not perform above chance level in the initial trials of the on–off condition, but 13 subjects learned to solve it. Their performance in the transfer tasks suggests that dogs can learn to solve the support problem based on perceptual cues, that they can quickly adopt new cues when old ones become unreliable, but also that some apparently inherent preferences are hard to overcome. Our study contributes to accumulating evidence demonstrating that animals typically rely on a variety of perceptual cues to solve physical cognition tasks, without developing an understanding of the underlying causal structure.