Going back to "basics": Harlow's learning set task with wolves and dogs.

To survive and reproduce, animals need to behave adaptively by adjusting their behavior to their environment, with learning facilitating some of these processes. Dogs have become a go-to model species in comparative cognition studies, making our understanding of their learning skills paramount at multiple levels, not only with regards to basic research on their cognitive skills and the effects of domestication, but also with applied purposes such as training. In order to tackle these issues, we tested similarly raised wolves and dogs in a serial learning task inspired by Harlow's "learning set." In Phase 1, different pairs of objects were presented to the animals, one of which was baited while the other was not. Both species' performance gradually improved with each new set of objects, showing that they "learnt to learn," but no differences were found between the species in their learning speed. In Phase 2, once subjects had learned the association between one of the objects and the food reward, the contingencies were reversed and the previously unrewarded object of the same pair was now rewarded. Dogs' performance in this task seemed to be better than wolves', albeit only when considering just the first session of each reversal, suggesting that the dogs might be more flexible than wolves. Further research (possibly with the aid of refined methods such as computer-based tasks) would help ascertain whether these differences between wolves and dogs are persistent across different learning tasks.

Selective responding to human ostensive communication is an early developing capacity of domestic dogs.

Ostensive communication, communication motivated not only by an informative intention, but also by an intention to make this informative intention overt, is thought to be restricted to human signalers and to rely on metacognitive skills. On the receiver side, human infants and dogs have been found to selectively respond to human ostensive communication, even if it remains unclear whether they are able to discern the underlying intentions. At present only animals with extensive experiences with humans have demonstrated a response to human attention calling, suggesting that this sensitivity may be an exaptation of communicative skills allowing for context-specific responding to intraspecific signals. We investigated whether dogs could respond to subtle cues of ostension without clear attention-calling and how this skill varies across age and with different human-given signals. Using a two-way object choice task, we evaluated whether dogs of different ages followed, with their gaze and with their choices, ostensive human pointing and gaze more than a formally similar but non-ostensive gesture and a directional gaze cue. Dogs followed pointing more than gaze, and older dogs followed the directional cues more accurately than younger dogs. Independent of cue type, we found that dogs of all ages responded to human ostensive signals more than similar directional cues motivated by no communicative intention. Given that dogs are sensitive to subtle cues of ostensive communication from an early age, and that this does not appear to change with age, we suggest that living close to humans may have selected for this skill. RESEARCH HIGHLIGHTS: Like human infants, dogs followed human ostensive cues more than non-ostensive cues both by their approach behavior and their differential looking behavior. Dogs differentiated between ostensive and non-ostensive human cues from an early age on, without their attention having been called explicitly in advance. Dogs followed human-given pointing cues more than human-given gaze cues, independent of age or intention (ostensive or non-ostensive). The early developing sensitivity of dogs to human ostension suggests that living close to humans may have selected for this skill.

How to improve data quality in dog eye tracking.

Pupil-corneal reflection (P-CR) eye tracking has gained a prominent role in studying dog visual cognition, despite methodological challenges that often lead to lower-quality data than when recording from humans. In the current study, we investigated if and how the morphology of dogs might interfere with tracking of P-CR systems, and to what extent such interference, possibly in combination with dog-unique eye-movement characteristics, may undermine data quality and affect eye-movement classification when processed through algorithms. For this aim, we have conducted an eye-tracking experiment with dogs and humans, and investigated incidences of tracking interference, compared how they blinked, and examined how differential quality of dog and human data affected the detection and classification of eye-movement events. Our results show that the morphology of dogs' face and eye can interfere with tracking methods of the systems, and dogs blink less often but their blinks are longer. Importantly, the lower quality of dog data lead to larger differences in how two different event detection algorithms classified fixations, indicating that the results of key dependent variables are more susceptible to choice of algorithm in dog than human data. Further, two measures of the Nyström & Holmqvist (Behavior Research Methods, 42(4), 188-204, 2010) algorithm showed that dog fixations are less stable and dog data have more trials with extreme levels of noise. Our findings call for analyses better adjusted to the characteristics of dog eye-tracking data, and our recommendations help future dog eye-tracking studies acquire quality data to enable robust comparisons of visual cognition between dogs and humans.

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

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 trials needed to reach a learning criterion and their pessimistic bias in a cognitive bias test. No difference between the two groups emerged in terms of learning speed; however, dogs that were partially rewarded during clicker training showed a more pessimistic bias than dogs that were continuously rewarded. Generally, emotional reactivity was positively associated with a more pessimistic bias. Partial rewarding does not improve training efficacy, but it is associated with a negatively valenced affective state, bringing support to the hypothesis that partial rewarding might negatively affect dogs' welfare.

Dogs wait longer for better rewards than wolves in a delay of gratification task: but why?

Self-control has been shown to be linked with being cooperative and successful in humans and with the g-factor in chimpanzees. As such, it is likely to play an important role in all forms of problem-solving. Self-control, however, does not just vary across individuals but seems also to be dependent on the ecological niche of the respective species. With dogs having been selected to live in the human environment, several domestication hypotheses have predicted that dogs are better at self-control and thus more tolerant of longer delays than wolves. Here we set out to test this prediction by comparing dogs' and wolves' self-control abilities using a delay of gratification task where the animals had to wait for a predefined delay duration to exchange a low-quality reward for a high-quality reward. We found that in our task, dogs outperformed the wolves waiting an average of 66 s vs. 24 s in the wolves. Food quality did not influence how long the animals waited for the better reward. However, dogs performed overall better in motivation trials than the wolves, although the dogs' performance in those trials was dependent on the duration of the delays in the test trials, whereas this was not the case for the wolves. Overall, the data suggest that selection by humans for traits influencing self-control rather than ecological factors might drive self-control abilities in wolves and dogs. However, several other factors might contribute or explain the observed differences including the presence of the humans, which might have inhibited the dogs more than the wolves, lower motivation of the wolves compared to the dogs to participate in the task and/or wolves having a better understanding of the task contingencies. These possible explanations need further exploration.

Importance of a species' socioecology: Wolves outperform dogs in a conspecific cooperation task.

A number of domestication hypotheses suggest that dogs have acquired a more tolerant temperament than wolves, promoting cooperative interactions with humans and conspecifics. This selection process has been proposed to resemble the one responsible for our own greater cooperative inclinations in comparison with our closest living relatives. However, the socioecology of wolves and dogs, with the former relying more heavily on cooperative activities, predicts that at least with conspecifics, wolves should cooperate better than dogs. Here we tested similarly raised wolves and dogs in a cooperative string-pulling task with conspecifics and found that wolves outperformed dogs, despite comparable levels of interest in the task. Whereas wolves coordinated their actions so as to simultaneously pull the rope ends, leading to success, dogs pulled the ropes in alternate moments, thereby never succeeding. Indeed in dog dyads it was also less likely that both members simultaneously engaged in other manipulative behaviors on the apparatus. Different conflict-management strategies are likely responsible for these results, with dogs' avoidance of potential competition over the apparatus constraining their capacity to coordinate actions. Wolves, in contrast, did not hesitate to manipulate the ropes simultaneously, and once cooperation was initiated, rapidly learned to coordinate in more complex conditions as well. Social dynamics (rank and affiliation) played a key role in success rates. Results call those domestication hypotheses that suggest dogs evolved greater cooperative inclinations into question, and rather support the idea that dogs' and wolves' different social ecologies played a role in affecting their capacity for conspecific cooperation and communication.

Training pet dogs for eye-tracking and awake fMRI.

In recent years, two well-developed methods of studying mental processes in humans have been successively applied to dogs. First, eye-tracking has been used to study visual cognition without distraction in unrestrained dogs. Second, noninvasive functional magnetic resonance imaging (fMRI) has been used for assessing the brain functions of dogs in vivo. Both methods, however, require dogs to sit, stand, or lie motionless while yet remaining attentive for several minutes, during which time their brain activity and eye movements are measured. Whereas eye-tracking in dogs is performed in a quiet and, apart from the experimental stimuli, nonstimulating and highly controlled environment, MRI scanning can only be performed in a very noisy and spatially restraining MRI scanner, in which dogs need to feel relaxed and stay motionless in order to study their brain and cognition with high precision. Here we describe in detail a training regime that is perfectly suited to train dogs in the required skills, with a high success probability and while keeping to the highest ethical standards of animal welfare-that is, without using aversive training methods or any other compromises to the dog's well-being for both methods. By reporting data from 41 dogs that successfully participated in eye-tracking training and 24 dogs IN fMRI training, we provide robust qualitative and quantitative evidence for the quality and efficiency of our training methods. By documenting and validating our training approach here, we aim to inspire others to use our methods to apply eye-tracking or fMRI for their investigations of canine behavior and cognition.

Cognitive Aging in Dogs.

A decline in the physical or mental health of older dogs can be a challenge for the owners, whose relationship with their dog is compromised by the cognitive and behavioral changes in their dogs. Although dog owners tend to consider many physiological and behavioral changes in old dogs as part of the normal aging process, it is important to differentiate between normal aging and pathologic aging, since behavioral changes may be the first indication of declining health and welfare in old dogs. Most reviews on cognitive aging in dogs have focused on translational approaches to human Alzheimer's disease; from a practical perspective, however, understanding normal cognitive aging in pet dogs and screening cognitively affected dogs are important in their own right. Here we review the literature on different cognitive functions that decline during aging, signs of cognitive dysfunction, screening methods, and preventive measures for age-related cognitive decline. Moreover, we discuss the drawbacks of using questionnaires as subjective measures of aging and propose the development of objective methods to distinguish normal cognitive aging from severe cognitive dysfunction. We suggest that multi-targeted approaches that combine owner-evaluated questionnaires with neuropsychological tests can be most effective in screening cognitively affected dogs from normally aging dogs. Regarding preventive measures, we conclude that combinations of dietary intervention and behavioral enrichment may be more beneficial than single-pathway manipulations in delaying cognitive aging or retaining various cognitive functions during aging.

DNA methylation patterns of behavior-related gene promoter regions dissect the gray wolf from domestic dog breeds.

A growing body of evidence highlights the relationship between epigenetics, especially DNA methylation, and population divergence as well as speciation. However, little is known about how general the phenomenon of epigenetics-wise separation of different populations is, or whether population assignment is, possible based on solely epigenetic marks. In the present study, we compared DNA methylation profiles between four different canine populations: three domestic dog breeds and their ancestor the gray wolf. Altogether, 79 CpG sites constituting the 65 so-called CpG units located in the promoter regions of genes affecting behavioral and temperamental traits (COMT, HTR1A, MAOA, OXTR, SLC6A4, TPH1, WFS1)-regions putatively targeted during domestication and breed selection. Methylation status of buccal cells was assessed using EpiTYPER technology. Significant inter-population methylation differences were found in 52.3% of all CpG units investigated. DNA methylation profile-based hierarchical cluster analysis indicated an unambiguous segregation of wolf from domestic dog. In addition, one of the three dog breeds (Golden Retriever) investigated also formed a separate, autonomous group. The findings support that population segregation is interrelated with shifts in DNA methylation patterns, at least in putative selection target regions, and also imply that epigenetic profiles could provide a sufficient basis for population assignment of individuals.

Is a local sample internationally representative? Reproducibility of four cognitive tests in family dogs across testing sites and breeds.

A fundamental precept of the scientific method is reproducibility of methods and results, and there is growing concern over the failure to reproduce significant results. Family dogs have become a favoured species in comparative cognition research, but they may be subject to cognitive differences arising from genetic (breeding lines) or cultural differences (e.g. preferred training methods). Such variation is of concern as it affects the validity and generalisability of experimental results. Despite its importance, this problem has not been specifically addressed to date. Therefore, we aimed to test the influence of three factors on reproducibility: testing site (proximal environment), breed and sex (phenotype). The same experimenter tested cognitive performance by more than 200 dogs in four experiments. Additionally, dogs' performance was tested in an obedience task administered by the owner. Breed of dog and testing site were found to influence the level of performance only mildly, and only in a means-end experiment and the obedience task. Our findings demonstrate that by applying the same test protocols on sufficiently large samples, the reported phenomena in these cognitive tests can be reproduced, but slight differences in performance levels can occur between different samples. Accordingly, we recommend the utilisation of well-described protocols supported by video examples of the whole experimental procedure. Findings should focus on the main outcome variables of the experiments, rather than speculating about the general importance of small or secondary performance outcomes which are more susceptible to random or local noise.

Individual and group level trajectories of behavioural development in Border collies.

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.

Testing the myth: tolerant dogs and aggressive wolves.

Cooperation is thought to be highly dependent on tolerance. For example, it has been suggested that dog-human cooperation has been enabled by selecting dogs for increased tolerance and reduced aggression during the course of domestication ('emotional reactivity hypothesis'). However, based on observations of social interactions among members of captive packs, a few dog-wolf comparisons found contradictory results. In this study, we compared intraspecies aggression and tolerance of dogs and wolves raised and kept under identical conditions by investigating their agonistic behaviours and cofeeding during pair-wise food competition tests, a situation that has been directly linked to cooperation. We found that in wolves, dominant and subordinate members of the dyads monopolized the food and showed agonistic behaviours to a similar extent, whereas in dogs these behaviours were privileges of the high-ranking individuals. The fact that subordinate dogs rarely challenged their higher-ranking partners suggests a steeper dominance hierarchy in dogs than in wolves. Finally, wolves as well as dogs showed only rare and weak aggression towards each other. Therefore, we suggest that wolves are sufficiently tolerant to enable wolf-wolf cooperation, which in turn might have been the basis for the evolution of dog-human cooperation (canine cooperation hypothesis).

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.

Cross-modal perception of puppies and adult conspecifics in dogs (Canis familiaris).

Understanding conspecifics' age classes is crucial for animals, facilitating adaptive behavioral responses to their social environment. This may include gathering and integrating information through multiple modalities. Using a cross-modal preferential-looking paradigm, we investigated whether dogs possess a cross-modal mental representation of conspecific age classes. In Experiment 1, dogs were presented with images of an adult dog and a puppy projected side by side on a wall while a vocalization of either an adult dog or a puppy was played back simultaneously. To test the effect of relative body size between adult dog and puppy images, two size conditions (natural size and same size) were employed for visual stimuli. We examined dogs' looking behavior in response to cross-modally matched versus mismatched stimuli. We predicted that if dogs have cross-modal representations of age classes, they would exhibit prolonged attention toward matched images compared to mismatched ones. In Experiment 2, we administered the same paradigm within an eye-tracking experiment to further improve the measurement quality of dogs' looking times. However, dogs' looking times in either experiment did not demonstrate significant differences based on the match or mismatch between image and vocalization. Instead, we observed a size effect, indicating dogs' increased attention toward larger adult dog images compared to smaller puppy images. Consequently, we found no evidence of cross-modal representation of age class in dogs. Nonetheless, we found increased looking time and pupil size upon hearing puppy vocalizations compared to adult vocalizations in Experiment 2, suggesting that dogs exhibited heightened arousal when hearing puppy whining. (PsycInfo Database Record (c) 2024 APA, all rights reserved).

Choice of conflict resolution strategy is linked to sociability in dog puppies.

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.

The Power of Discourse: Associations between Trainers' Speech and the Responses of Socialized Wolves and Dogs to Training.

In a previous study, we found that Positive Reinforcement Training reduced cortisol of wolves and dogs; however, this effect varied across trainer-animal dyads. Here we investigate whether and how the trainers' use of speech may contribute to this effect. Dogs' great interest in high-pitched, intense speech (also known as Dog Directed Speech) has already been reported, but whether and how wolves respond similarly/differently to voice characteristics has never been studied before. We analyzed 270 training sessions, conducted by five trainers, with nine mixed-breed dogs and nine wolves, all human-socialized. Through Generalized Linear Mixed Models, we analyzed the effects of (a) three speech categories (nice, neutral, reprehensive) and laugh; and (b) acoustic characteristics of trainers' voices on animals' responses (correct responses, latency, orientation, time at less than 1 m, non-training behaviors, tail position/movements, cortisol variation). In both subspecies, tail wagging occurred more often in sessions with longer durations of nice speech, and less often in sessions with reprehensive speech. For dogs, the duration of reprehensive speech within a session was also negatively related to correct responses. For wolves, retreat time was associated with more reprehensive speech, whereas duration of nice speech was positively associated with time spent within one meter from the trainer. In addition, most dog behavioral responses were associated with higher average intonations within sessions, while wolf responses were correlated with lower intonations within sessions. We did not find any effects of the variables considered on cortisol variation. Our study highlights the relevance of voice tone and speech in a training context on animals' performances and emotional reactions.

Object permanence in adult common marmosets (Callithrix jacchus): not everything is an "A-not-B" error that seems to be one.

In this paper, we describe a behaviour pattern similar to the "A-not-B" error found in human infants and young apes in a monkey species, the common marmosets (Callithrix jacchus). In contrast to the classical explanation, recently it has been suggested that the "A-not-B" error committed by human infants is at least partially due to misinterpretation of the hider's ostensively communicated object hiding actions as potential 'teaching' demonstrations during the A trials. We tested whether this so-called Natural Pedagogy hypothesis would account for the A-not-B error that marmosets commit in a standard object permanence task, but found no support for the hypothesis in this species. Alternatively, we present evidence that lower level mechanisms, such as attention and motivation, play an important role in committing the "A-not-B" error in marmosets. We argue that these simple mechanisms might contribute to the effect of undeveloped object representational skills in other species including young non-human primates that commit the A-not-B error.

Domestic dogs (Canis familiaris) flexibly adjust their human-directed behavior to the actions of their human partners in a problem situation.

Domestic dogs (Canis familiaris) have been shown to actively initiate triadic communicative interactions by looking at a human partner or by alternating their gaze between the human and an object when being faced with an out-of-reach reward or an unsolvable problem. It has hardly been investigated, however, whether dogs flexibly adjust their human-directed behavior to the actions of their partners, which indicate their willingness and abilities to help them when they are faced with a problem. Here, in two experiments, we confronted dogs-after initially allowing them to learn how to manipulate an apparatus-with two problem situations: with an empty apparatus and a blocked apparatus. In Experiment 1, we showed that dogs looked back at their owners more when the owners had previously encouraged them, independently from the problem they faced. In Experiment 2, we provided dogs with two experimenters and allowed them to learn through an initial phase that each of the experimenters could solve one of the two problems: the Filler re-baited the empty apparatus and the Helper unblocked the blocked apparatus. We found that dogs could learn to recognize the ability of the Filler and spent time close to her when the apparatus was empty. Independently from the problem, however, they always approached the Helper first. The results of the present study indicate that dogs may have a limited understanding of physical problems and how they can be solved by a human partner. Nevertheless, dogs are able to adjust their behavior to situation-specific characteristics of their human partner's behavior.

Does the A-not-B error in adult pet dogs indicate sensitivity to human communication?

Recent dog-infant comparisons have indicated that the experimenter's communicative signals in object hide-and-search tasks increase the probability of perseverative (A-not-B) errors in both species (Topál et al. 2009). These behaviourally similar results, however, might reflect different mechanisms in dogs and in children. Similar errors may occur if the motor response of retrieving the object during the A trials cannot be inhibited in the B trials or if the experimenter's movements and signals toward the A hiding place in the B trials ('sham-baiting') distract the dogs' attention. In order to test these hypotheses, we tested dogs similarly to Topál et al. (2009) but eliminated the motor search in the A trials and 'sham-baiting' in the B trials. We found that neither an inability to inhibit previously rewarded motor response nor insufficiencies in their working memory and/or attention skills can explain dogs' erroneous choices. Further, we replicated the finding that dogs have a strong tendency to commit the A-not-B error after ostensive-communicative hiding and demonstrated the crucial effect of socio-communicative cues as the A-not-B error diminishes when location B is ostensively enhanced. These findings further support the hypothesis that the dogs' A-not-B error may reflect a special sensitivity to human communicative cues. Such object-hiding and search tasks provide a typical case for how susceptibility to human social signals could (mis)lead domestic dogs.

The absence of reward induces inequity aversion in dogs.

One crucial element for the evolution of cooperation may be the sensitivity to others' efforts and payoffs compared with one's own costs and gains. Inequity aversion is thought to be the driving force behind unselfish motivated punishment in humans constituting a powerful device for the enforcement of cooperation. Recent research indicates that non-human primates refuse to participate in cooperative problem-solving tasks if they witness a conspecific obtaining a more attractive reward for the same effort. However, little is known about non-primate species, although inequity aversion may also be expected in other cooperative species. Here, we investigated whether domestic dogs show sensitivity toward the inequity of rewards received for giving the paw to an experimenter on command in pairs of dogs. We found differences in dogs tested without food reward in the presence of a rewarded partner compared with both a baseline condition (both partners rewarded) and an asocial control situation (no reward, no partner), indicating that the presence of a rewarded partner matters. Furthermore, we showed that it was not the presence of the second dog but the fact that the partner received the food that was responsible for the change in the subjects' behavior. In contrast to primate studies, dogs did not react to differences in the quality of food or effort. Our results suggest that species other than primates show at least a primitive version of inequity aversion, which may be a precursor of a more sophisticated sensitivity to efforts and payoffs of joint interactions.