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Facial expressions potentially serve as indicators of animal emotions if they are consistently present across situations that (likely) elicit the same emotional state. In a previous study, we used the Dog Facial Action Coding System (DogFACS) to identify facial expressions in dogs associated with conditions presumably eliciting positive anticipation (expectation of a food reward) and frustration (prevention of access to the food). Our first aim here was to identify facial expressions of positive anticipation and frustration in dogs that are context-independent (and thus have potential as emotion indicators) and to distinguish them from expressions that are reward-specific (and thus might relate to a motivational state associated with the expected reward). Therefore, we tested a new sample of 28 dogs with a similar set-up designed to induce positive anticipation (positive condition) and frustration (negative condition) in two reward contexts: food and toys. The previous results were replicated: Ears adductor was associated with the positive condition and Ears flattener, Blink, Lips part, Jaw drop, and Nose lick with the negative condition. Four additional facial actions were also more common in the negative condition. All actions except the Upper lip raiser were independent of reward type. Our second aim was to assess basic measures of diagnostic accuracy for the potential emotion indicators. Ears flattener and Ears downward had relatively high sensitivity but low specificity, whereas the opposite was the case for the other negative correlates. Ears adductor had excellent specificity but low sensitivity. If the identified facial expressions were to be used individually as diagnostic indicators, none would allow consistent correct classifications of the associated emotion. Diagnostic accuracy measures are an essential feature for validity assessments of potential indicators of animal emotion.

Dogs are renowned for ‘looking back’ at humans when confronted with a problem, but it has been questioned whether this implies help-seeking or giving up. We tested 56 pet dogs from two breed groups (herding dogs and terriers) in a modified unsolvable task paradigm. One reward type (food or toy) was enclosed in a box, while the respective other reward was accessible. With both reward types, human-directed gazing in relation to the box was significantly positively correlated with interaction with the box, as long as an alternative was available. This suggests that both behaviours served to attain the unavailable reward and reflected individual motivation for the inaccessible vs the accessible reward. Furthermore, we varied whether the owner or the experimenter was responsible for handling the rewards. In the owner-responsible group, dogs rarely gazed at the experimenter. In the experimenter-responsible group, dogs preferentially directed box-related gazing (prior to or after looking at or interacting with the box) at the owner. Still, they gazed at the experimenter significantly longer than the owner-responsible group. Conversely, toy-related gazing was directed significantly more at the experimenter. Thus, dogs adjust their gazing behaviour according to the people’s responsibility and their current goal (help-seeking vs play). Gaze duration did not differ between herding dogs and terriers. We conclude that dogs use gazing at humans’ faces as a social problem-solving strategy, but not all gazing can be classified as such. Dogs’ human-directed gazing is influenced by the social relationships with the persons, situational associations, and context (unsolvable problem vs play). Supplementary Information The online version contains supplementary material available at 10.1007/s10071-021-01595-0.

Although free-roaming domestic dogs (FRDD) constitute the majority of the dog population worldwide, many aspects of their ecology across habitats are little known. Activity budgets by these dogs may also inform management decisions for domestic dogs in human hands. Here we collected data on the activity patterns of owned FRDD from Guatemala (n = 58) and Indonesia (n = 37), and of farm dogs (n = 11) and family dogs (n = 20) in Switzerland. The FRDD from the two countries and the Swiss farm dogs shared the similarity that although they had owners, they spent most or all of the day outside without confinement. Conversely, activity in family dogs is largely controlled by their owners. This cross-continental study thus allowed us to tease apart environmental effects on dogs’ activity from effects due to different levels of control by humans. Dogs were collared with FitBark activity trackers, which measure 3D acceleration, for 2.4–7 days. Activity for each dog was defined as the sum of BarkPoints (a continuous activity metric recorded by the FitBark tracker), calculated for each hour in the 24-hour cycle. The proportion of time resting, in ‘moderate’ and ‘high’ activity (defined by fixed thresholds of BarkPoints) over 24 h was calculated for each dog. The activity patterns of all dogs that (partly) roam freely, i.e. owned FRDD in Guatemala and Indonesia and Swiss farm dogs, showed two peaks over 24 h during 5:00–7:00 h and, less pronounced, 16:00–19:00 h. Such a bimodal activity pattern, which is also observed in other canine species, could only be detected in 45% of the family dogs. Their activity is more dependent on the owners’ daily routines and predominantly showed one high mid-day peak that often changes from day to day. Swiss dogs spent significantly more time resting and less time with ‘moderate’ activity than the owned FRDD. However, family dogs were significantly more often highly active than all other dog groups and compensated with longer resting periods. Activity decreased significantly with age, neutering and increased body condition score, whereas sex did not have any significant influence on activity. Within this study, similarities, but also differences of the activity pattern between owned FRDD and pet dogs could be revealed. Although overall activity levels of the pet dog sample fall in the range of those observed in the less controlled FRDD, it would be of interest to investigate the potential benefit of a more structured daily schedule on pet dogs in future studies.

Guinea pigs are often involved in animal-assisted therapy (AAT) but there is little knowledge about the effects of human contact on guinea pigs involved in AAT. The aim of this study was to investigate effects of availability of a retreat, presence of conspecifics, prior experience with AAT, and human interaction on indicators of welfare in guinea pigs involved in AAT. Guinea pigs of both sexes and different ages (n=20) were assigned to a randomized, controlled within-subject trial with repeated measurements. Each guinea pig was tested in four settings: (I) therapy with retreat possibility with conspecifics, (II) therapy with retreat possibility without conspecifics, (III) therapy without retreat possibility, and (IV) setting without human interaction. We measured changes in eye temperature, as a proxy to infer stress levels, at 5-s intervals with a thermographic camera. All sessions were video recorded and the guinea pigs' behavior was coded using continuous recording and focal animal sampling. For the statistical analysis we used generalized linear mixed models, with therapy setting as a fixed effect and individual guinea pig as a random effect. We observed a temperature increase relative to baseline in settings (I) therapy with retreat with conspecifics present and (III) therapy without retreat. The percentage of time a guinea pig was petted was positively correlated with a rise in the eye temperature independent of the setting. Time spent eating was reduced in all therapy settings (I-III) compared to the setting without HAI (human animal interaction) (IV). In the setting with retreat (I), guinea pigs showed more active behaviors such as locomotive behavior or startling compared to the setting without retreat (III) and the setting without HAI (IV). When no retreat was available (III), they showed more passive behaviors, such as standing still or freezing compared to therapy with retreat (I). Based on our results we identified the behaviors "reduced eating", "increased startle" and "increased freezing" as indicators of an increased stress level. Petting the guinea pigs was correlated with a rise in the eye temperature and might be a factor which can cause stress. Our results support the suggestion that guinea pigs involved in AAT should have a retreat possibility, should have access to conspecifics, and should be given time to adapt to a new setting. In this way, stress might be reduced.

A high proportion of pet dogs show fear-related behavioural problems, with noise fears being most prevalent. Nonetheless, few studies have objectively evaluated fear expression in this species. Using owner-provided video recordings, we coded behavioural expressions of pet dogs during a real-life firework situation at New Year’s Eve and compared them to behaviour of the same dogs on a different evening without fireworks (control condition), using Wilcoxon signed ranks tests. A backwards-directed ear position, measured at the base of the ear, was most strongly associated with the fireworks condition (effect size: Cohen’s d = 0.69). Durations of locomotion (d = 0.54) and panting (d = 0.45) were also higher during fireworks than during the control condition. Vocalisations (d = 0.40), blinking (d = 0.37), and hiding (d = 0.37) were increased during fireworks, but this was not significant after sequential Bonferroni correction. This could possibly be attributed to the high inter-individual variability in the frequency of blinking and the majority of subjects not vocalising or hiding at all. Thus, individual differences must be taken into account when aiming to assess an individual’s level of fear, as relevant measures may not be the same for all individuals. Firework exposure was not associated with an elevated rate of other so-called ‘stress signals’, lip licking and yawning.

Successive negative contrast (SNC) effects are changes in anticipatory or consummatory behaviour when animals unexpectedly receive a lower value reward than they have received previously. SNC effects are often assumed to reflect frustration and appear to be influenced by background affective state. However, alternative explanations of SNC, such as the functional-search hypothesis, do not necessarily imply an aversive affective state. We tested 18 dogs in a SNC paradigm using a patch foraging task. Dogs were tested in two conditions, once with the low value reward in all of five trials (unshifted) and once when reward value was altered between high and low (shifted). Following a reward downshift, subjects showed a SNC effect by switching significantly more often between patches compared to the unshifted condition. However, approach latency, foraging time and quantity consumed did not differ between conditions, suggesting non-affective functional search behaviour rather than frustration. There was no relationship between strength of SNC and anxiety-related behaviours as measured in a novel object test and a personality questionnaire (C-BARQ). However, associations with the C-BARQ scores for Trainability and Stranger directed aggression suggest a possible link with behavioural flexibility and coping style. While reward quality clearly affects incentive motivation, the relationship between SNC, frustration and background affective state requires further exploration.

Reinforcer effectiveness refers to the reinforcer’s ability to control the subject’s target behaviour and is therefore critical to training success. Yet animals’ preferences, and the effectiveness of different rewards to function as reinforcers, are often assumed without scientific investigation. Here we explored the influence of reward quality, quantity and changes in reward value on motivation in domestic dogs. Subjects were trained to traverse a runway for a food reward. In Study 1, the quantity of food was varied (1 vs 5 pieces of dry food), while in Study 2, food quality was varied (1 piece of sausage vs 1 piece of dry food). Dogs were tested in two conditions (counterbalanced). In the unshifted condition, they received the low value reward in all of ten trials; in the shifted condition, reward value was altered (high value: trials 1–4 and 9–10; low value: trials 5–8). While preliminary preference tests had confirmed the relative value of the presumed high and low value rewards for both quantity and quality, dogs' responses in the runway task differed between the quality and quantity studies. Dogs ran significantly faster for the higher quality food compared to the lower quality food, confirming greater reinforcer effectiveness of the preferred food type. In contrast, there was no significant effect of food quantity on running speed at any stage. Higher quality rewards therefore appear to entail greater incentive motivation in dogs than a greater quantity of a lower value reward, with reward-specific habituation needing to be considered.

Facial expressions are considered sensitive indicators of emotional states in humans and many animals. Identifying facial indicators of emotion is a major challenge and little systematic research has been done in non-primate species. In dogs, such research is important not only to address fundamental and applied scientific questions but also for practical reasons, since many problem behaviours are assumed to have an emotional basis, e.g. aggression based on frustration. Frustration responses can occur in superficially similar contexts as the emotional state of positive anticipation. For instance, the anticipated delivery of a food reward may induce the state of positive anticipation, but over time, if the food is not delivered, this will be replaced by frustration. We examined dogs’ facial expressions in contexts presumed to induce both positive anticipation and frustration, respectively, within a single controlled experimental setting. Using DogFACS, an anatomically-based method for coding facial expressions of dogs, we found that the “Ears adductor” action was more common in the positive condition and “Blink”, “Lips part”, “Jaw drop”, “Nose lick”, and “Ears flattener” were more common in the negative condition. This study demonstrates how differences in facial expression in emotionally ambiguous contexts may be used to help infer emotional states of different valence.

Recently, there has been a move towards positive reinforcement using food rewards in animal training. By definition, rewards function as reinforcers if they increase or maintain the frequency of behaviour that they follow. However, in operant conditioning tasks animals frequently show systematic changes in performance – in particular a reduction in responding over time. One suggested strategy to avoid such performance decrements is to provide a variety of food rewards, rather than the same food reward in all trials. The enhancement of appetitive behaviour and consumption by reward variation is referred to as ‘variety effect’. We investigated whether dogs preferred a variable or a constant food reward in a concurrent two-choice test. Of 16 dogs, six subjects showed a significant preference for the varied food reward and six for the constant food reward, while four dogs exhibited no significant preference for either option. At the group level, there was a significant effect of block: preference for the varied food reward increased across six blocks of ten trials each. Thus, although some individuals may prefer a single, favourite food reward in the short term, introducing variation in reward types may maintain dogs’ motivation in operant tasks over a longer time period.

When an anticipated food reward is unexpectedly reduced in quality or quantity, many mammals show a successive negative contrast (SNC) effect, i.e. a reduction in instrumental or consummatory responses below the level shown by control animals that have only ever received the lower-value reward. SNC effects are believed to reflect an aversive emotional state, caused by the discrepancy between the expected and the actual reward. Furthermore, how animals respond to such discrepancy has been suggested to be a sign of animals' background mood state. However, the occurrence and interpretation of SNC effects are not unequivocal, and there is a relative lack of studies conducted outside of laboratory conditions. Here, we tested two populations of domestic dogs (24 owned pet dogs and 21 dogs from rescue kennels) in a SNC paradigm following the methodology by Bentosela et al. (J Comp Psychol 123:125-130, 2009), using a design that allowed a within-, as well as a between-, subjects analysis. We found no evidence of a SNC effect in either population using a within- or between-subjects design. Indeed, the within-subjects analysis revealed a reverse SNC effect, with subjects in the shifted condition showing a significantly higher level of response, even after they received an unexpected reduction in reward quality. Using a within-, rather than a between-, subjects design may be beneficial in studies of SNC due to higher sensitivity and statistical power; however, order effects on subject performance need to be considered. These results suggest that this particular SNC paradigm may not be sufficiently robust to replicate easily in a range of environmental contexts and populations.

In order to assess dogs’ personality changes during ontogeny, a cohort of 69 Border collies was followed up from six to 18–24 months. When the dogs were 6, 12, and 18–24 months old, their owners repeatedly filled in a dog personality questionnaire (DPQ), which yielded five personality factors divided into fifteen facets. All five DPQ factors were highly correlated between the three age classes, indicating that the dogs’ personality remained consistent relative to other individuals. Nonetheless, at the group level significant changes with age were found for four of the five DPQ factors. Fearfulness, Aggression towards People, Responsiveness to Training and Aggression towards Animals increased with age; only Activity/Excitability did not change significantly over time. These changes in DPQ factor scores occurred mainly between the ages of 6 and 12 months, although some facets changed beyond this age. No sex differences were found for any of the tested factors or facets, suggesting that individual variation in personality was greater than male/female differences. There were significant litter effects for the factors Fearfulness, Aggression towards People and Activity/Excitability, indicating either a strong genetic basis for these traits or a high influence of the shared early environment. To conclude, from the age of six months, consistency in personality relative to other individuals can be observed in Border collies. However, at the group level, increases in fearful and aggressive behaviours occur up to 12 months and for some traits up to two years, highlighting the need for early interventions. Follow-up studies are needed to assess trajectories of personality development prior to six months and after two years, and to include a wider variety of breeds.

Visible and invisible displacement tasks have been used widely for comparative studies of animals' understanding of object permanence, with evidence accumulating that some species can solve invisible displacement tasks and, thus, reach Piagetian stage 6 of object permanence. In contrast, dogs appear to rely on associative cues, such as the location of the displacement device, during invisible displacement tasks. It remains unclear, however, whether dogs, and other species that failed in invisible displacement tasks, do so because of their inability to form a mental representation of the target object, or simply because of the involvement of a more salient but potentially misleading associative cue, the displacement device. Here we show that the use of a displacement device impairs the performance of dogs also in visible displacement tasks: their search accuracy was significantly lower when a visible displacement was performed with a displacement device, and only two of initially 42 dogs passed the sham-baiting control conditions. The negative influence of the displacement device in visible displacement tasks may be explained by strong associative cues overriding explicit information about the target object's location, reminiscent of an overshadowing effect, and/or object individuation errors as the target object is placed within the displacement device and moves along a spatiotemporally identical trajectory. Our data suggest that a comprehensive appraisal of a species' performance in object permanence tasks should include visible displacement tasks with the same displacement device used in invisible displacements, which typically has not been done in the past.

String pulling tasks are commonly used to investigate recognition of means-end connections. Previous studies suggested that dogs base their choice on proximity rather than connectivity (Osthaus, Lea, & Slater, 2005), nonetheless, dogs performed successfully in the related support problem (Range, Hentrup, & Viranyi, 2011). To reinvestigate dogs' means-end understanding, we tested 34 Border collies in string pulling tasks in which the proximity of the reward to the connected string's end was varied. First, subjects were presented with a four-string task (four parallel perpendicular strings, one baited, with the reward in line with the correct string's end). Dogs that performed above chance in this task were tested with a curved string task, involving one straight and one curved string. When the reward was attached to the curved string, it was equidistant from both strings' ends so that choosing by proximity was not possible. Although group level performance was significantly above chance, only three of 20 dogs met criterion individually, of which one dog subsequently solved a broken string task upon its first presentation. However, the dogs seemed to be unable to overcome their proximity bias in a parallel diagonal string task where proximity of the unconnected string's end to the reward was misleading. We conclude that although dogs may not demonstrate means-end understanding spontaneously, some can learn to pay attention to connectivity when proximity is not a confounding factor. This study supports the notion that animals may apply several alternative strategies to solve physical problems, which are influenced by the test-setup.

A wealth of comparative data has been accumulated over the past decades on how animals acquire and use information about the physical world. Domestic dogs have typically performed comparably poorly in physical cognition tasks, though in a recent study Kundey et al. (Anim Cogn 13:497-505, 2010) challenged this view and concluded that dogs understand that objects cannot pass through solid barriers. However, the eight subjects in the study of Kundey et al. may have solved the task with the help of perceptual cues, which had not been controlled for. Here, we tested dogs with a similar task that excluded these cues. In addition, unlike the set-up of Kundey et al., our set-up allowed the subjects to observe the effect of the solid barrier. Nevertheless, all 28 subjects failed to solve this task spontaneously and showed no evidence of learning across 50 trials. Our results therefore call into question the earlier suggestion that dogs have, or can acquire, an understanding of the solidity principle.

Based on their high sociability and their capabilities in social cognition, we should consider that dogs experience rich emotions in the social domain. I discuss the importance of dog-human attachment and some of the controversial evidence for so-called secondary emotions, which leaves many questions to answer.

Human infants develop an understanding of their physical environment through playful interactions with objects. Similar processes may influence also the performance of non-human animals in physical problem-solving tasks, but to date there is little empirical data to evaluate this hypothesis. In addition or alternatively to prior experiences, inhibitory control has been suggested as a factor underlying the considerable individual differences in performance reported for many species. Here we report a study in which we manipulated the extent of object-related experience for a cohort of dogs (Canis familiaris) of the breed Border Collie over a period of 18 months, and assessed their level of inhibitory control, prior to testing them in a series of four physical problem-solving tasks. We found no evidence that differences in object-related experience explain variability in performance in these tasks. It thus appears that dogs do not transfer knowledge about physical rules from one physical problem-solving task to another, but rather approach each task as a novel problem. Our results, however, suggest that individual performance in these tasks is influenced in a complex way by the subject’s level of inhibitory control. Depending on the task, inhibitory control had a positive or a negative effect on performance and different aspects of inhibitory control turned out to be the best predictors of individual performance in the different tasks. Therefore, studying the interplay between inhibitory control and problem-solving performance will make an important contribution to our understanding of individual and species differences in physical problem-solving performance.

Humans in a negative emotional state are more likely to judge ambiguous stimuli as negative. In recent years, similar judgement biases have been found in some non-human animals that were exposed to long-term or short-term treatments aimed at influencing their affective states. Here we tested pet dogs in the presence and absence of their owners in a judgement bias test with an established go/no-go procedure. Even though owner absence is thought to induce a state of anxiety in dogs that have formed an attachment bond with their primary caretakers, we found no difference between the dogs' responses to ambiguous stimuli in the presence or absence of their owners. This result may be explained by the absence of anxiety in dogs that are accustomed to brief periods of separation from their owners, or by a sensitivity limit of the customary judgement bias tests in non-human animals when only a moderate, short-term state of anxiety is induced. In addition, we found significant differences between individuals and populations in the responses to ambiguous stimuli, which give impetus for further research.

Highlights • Pet dogs were tested in a brief separation test and filmed remotely using thermography. • Temperature was analyzed from selected patches of both ear pinnae simultaneously. • Social isolation was associated with a significant decrease in ear pinnae temperature. • Temperature of the two ears did not differ significantly from each other. • Long distance thermography is a useful tool in non-invasive stress monitoring. Abstract Infrared thermography can visualize changes in body surface temperature that result from stress-induced physiological changes and alterations of blood flow patterns. Here we explored its use for remote stress monitoring (i.e. removing need for human presence) in a sample of six pet dogs. Dogs were tested in a brief separation test involving contact with their owner, a stranger, and social isolation for two one-minute-periods. Tests were filmed using a thermographic camera set up in a corner of the room, around 7 m from where the subjects spent most of the time. Temperature was measured from selected regions of both ear pinnae simultaneously. Temperatures of both ear pinnae showed a pattern of decrease during separation and increase when a person (either the owner or a stranger) was present, with no lateralized temperature differences between the two ears. Long distance thermographic measurement is a promising technique for non-invasive remote stress assessment, although there are some limitations related to dogs' hair structure over the ears, making it unsuitable for some subjects.

Measures that are likely to increase sociability in dog puppies, such as appropriate socialisation, are considered important in preventing future fear or aggression related problems. However, the interplay between sociability and conflict behaviour has rarely been investigated. Moreover, while many studies have addressed aggression in domestic dogs, alternative, non-aggressive conflict resolution strategies have received less scientific attention. Here we tested 134 Border collie puppies, aged 40-50 days, in a personality test which included friendly interactions with an unfamiliar person, exposure to a novel object, and three brief restraint tests. Considering the latter to be mild ‘conflict’ situations, we analysed whether the puppies’ behaviour in the restraint tests was related to their sociability or to their boldness towards the novel object. Strategies employed by the puppies during restraint tests included trying to interact socially with the experimenter, remaining passive, and attempting to move away. In line with findings from humans and goats, puppies scoring high on sociability were more likely to adopt an interactive conflict resolution strategy, while those with low sociability scores tended to react passively. In contrast, avoidance behaviours were unrelated to sociability, possibly reflecting inconsistency in the flight strategy in dogs. Boldness towards a novel object was not related to sociability or to puppies’ reactions in restraint tests. This is one of the first studies to demonstrate a link between sociability and conflict resolution strategies in non-human animals.