Simple Mechanisms Can Explain Social Learning in Domestic Dogs (Canis familiaris)

Recent studies have suggested that domestic dogs (Canis familiaris) engage in highly complex forms of social learning. Here, we critically assess the potential mechanisms underlying social learning in dogs using two problem‐solving tasks. In a classical detour task, the test dogs benefited from observing a demonstrator walking around a fence to obtain a reward. However, even inexperienced dogs did not show a preference for passing the fence at the same end as the demonstrator. Furthermore, dogs did not need to observe a complete demonstration by a human demonstrator to pass the task. Instead, they were just as successful in solving the problem after seeing a partial demonstration by an object passing by at the end of the fence. In contrast to earlier findings, our results suggest that stimulus enhancement (or affordance learning) might be a powerful social learning mechanism used by dogs to solve such detour problems. In the second task, we examined whether naïve dogs copy actions to solve an instrumental problem. After controlling for stimulus enhancement and other forms of social influence (e.g. social facilitation and observational conditioning), we found that dogs’ problem solving was not influenced by witnessing a skilful demonstrator (either an unknown human, a conspecific or the dog’s owner). Together, these results add to evidence suggesting that social learning may often be explained by relatively simple (but powerful) mechanisms.     

Interaction between individual experience and social learning in dogs

We investigated the interaction between individual experience and social learning in domestic dogs,Canis familiaris . We conducted two experiments using detour tests, where an object or food was placed behind a transparent, V-shaped wire-mesh fence, such that the dogs could get the reward by going around the fence. In some groups, two open doors were offered as an alternative, easier way to reach the reward. In experiment 1 we opened the doors only in trial 1, then closed them for trials 2 and 3. In experiment 2 other dogs were first taught to detour the fence with closed doors after they had observed a detouring human demonstrator, then we opened the doors for three subsequent trials. In experiment 1 all dogs reached the reward by going through the doors in trial 1, but their detouring performance was poor after the doors had been closed, if they had to solve the task on their own. However, dogs in the experimental group that were allowed to watch a detouring human demonstrator after the doors had been closed showed improved detouring ability compared with those that did not receive a demonstration of detouring. In experiment 2 the dogs tended to keep on detouring along the fence even if the doors had been opened, giving up a chance to get behind the fence by a shorter route. These results show that dogs can use information gained by observing a human demonstrator to overcome their own mistakenly preferred solution in a problem situation. In a reversed situation social learning can also contribute to a preference for a less adaptive behaviour. However, only repeated individual and social experience leads to a durable manifestation of maladaptive behaviour. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

The effect of ostensive cues on dogs’ performance in a manipulative social learning task

In contrast to animal social learning (e.g. dogs learning from observing another dog), humans typically teach by attracting the attention of the learner. Also during the training of dogs, humans tend to attract their attention in a similar way. Here, we investigated dogs’ ability to learn both from a dog and a human demonstrator in a manipulative task, where the models demonstrated which part of a box to manipulate in order to get a food reward. We varied the communicative context both during the dog and during the human demonstration comparably: a second experimenter directed the attention of the subjects to the model (dog/human ostensive demonstration) or remained silent (dog/human non-ostensive demonstration). Moreover, we investigated whether the training level of the dogs (well-trained vs. untrained) affected how the dogs performed in the manipulative tasks after the different demonstrations.We found that better trained dogs showed significantly better problem solving abilities. They paid more attention to the human demonstration than to the dog model, whereas such a difference in attentiveness of the less trained dogs was not found. Despite slight differences in paying attention to the different demonstrators, the presence of human or the dog demonstrators exerted equally effectiveness on the test performance of the dogs. However, the effectiveness of the demonstrations was significantly reduced if ostensive cues were given during the demonstrations by a second experimenter. Analysis of attentiveness and activity of the observer dogs during the demonstrations indicates that the reason for this negative effect was a combination of distracted attention paid to the demonstration and a higher level of excitement in the ostensive than in the non-ostensive demonstrations.This study suggests that third party communication during demonstration attracts dogs’ attention to the communicator instead of paying close attention to the model. We suggest that precise timing and synchronization of attention-calling and demonstration is necessary to avoid this distracting effect.     

The Kintamani dog: genetic profile of an emerging breed from Bali, Indonesia

The Kintamani dog is an evolving breed indigenous to the Kintamani region of Bali. Kintamani dogs cohabitate with feral Bali street dogs, although folklore has the breed originating 600 years ago from a Chinese Chow Chow. The physical and personality characteristics of the Kintamani dog make it a popular pet for the Balinese, and efforts are currently under way to have the dog accepted by the Federation Cynologique Internationale as a recognized breed. To study the genetic background of the Kintamani dog, 31 highly polymorphic short tandem repeat markers were analyzed in Kintamani dogs, Bali street dogs, Australian dingoes, and nine American Kennel Club (AKC) recognized breeds of Asian or European origin. The Kintamani dog was identical to the Bali street dog at all but three loci. The Bali street dog and Kintamani dog were most closely aligned with the Australian dingo and distantly related to AKC recognized breeds of Asian but not European origin. Therefore, the Kintamani dog has evolved from Balinese feral dogs with little loss of genetic diversity.     

Breed-typical behaviour in dogs—Historical remnants or recent constructs?

Dogs show considerable variation in morphology, genetics and behaviour caused by long periods of artificial selection. This is evident in the large number of breeds we have today. Behavioural differences among breeds have often been regarded as remnants from past selection during the breeds’ origin. However, the selection in many breeds has, during the last decades, gone through great changes, which could have influenced breed-typical behaviour. In order to investigate this, breed differences were studied using data from a standardized behavioural test from 13,097 dogs of 31 breeds from the Swedish dog population. Based on the test results, breed scores were calculated for four behavioural traits: playfulness, curiosity/fearlessness, sociability and aggressiveness. These traits have previously been found to be stable and valid, and hence regarded as personality traits in the dog. The present results suggested large differences between breeds in all of the investigated traits, even though there were within-breed variations. No relationships between breed-characteristic behaviour and function in the breeds’ origins were found. Instead, there were correlations between breed scores and current use of the breeding stocks, which suggest that selection in the recent past has affected breed-typical behaviour. The breeds’ use in dog shows, the dominating use in general, was negatively correlated with all investigated traits, both in sires and in dams. In contrast, use in Working dog trials was positively correlated with playfulness and aggressiveness in sires. Thus, these results suggest that selection for dog show use is positively correlated with social and non-social fearfulness, and negatively with playfulness, curiosity in potentially threatening situations and aggressiveness, whereas selection for Working dog use is positively correlated with playfulness and aggressiveness. Furthermore, correlation analyses show that popular breeds have higher sociability and playfulness scores than less popular breeds, suggesting that a positive attitude towards strangers is an important characteristic of a functional pet dog and desirable by dog owners. This indicates that selection towards use in dog shows may be in conflict with pet dog selection. Furthermore, these results suggest that basic dimensions of dog behaviour can be changed when selection pressure changes, and that the domestication of the dog still is in progress. A standardized behavioural test, like the one used in this study, is suggested to be highly useful as a tool in dog breeding programs.     

Links between play and dominance and attachment dimensions of dog-human relationships

It is often claimed that certain behavioral problems in domestic dogs can be triggered by the games played by dog and caregiver (owner). In this study, we examine possible links between the types of games played and dimensions of the dog-owner relationship that are generally considered to affect such problems. Fifty dog-owner partnerships were filmed during 3-min play sessions in which the owner was allowed to choose the games played. All partnerships then undertook a 1-hr test designed to measure elements of behavior commonly ascribed to “dominance ”and “attachment. ”Principal components analysis of the data produced 2 dominance-related factors (Amenability and Confident Interactivity) and 4 factors describing aspects of attachment (Nonspecific Attention Seeking, Preference for Owner, Preference for Unfamiliar Person, and Separation-Related Behavior). Amenability, in particular, varied significantly between breeds. In the study, we then compared types of games played to each of these factors. Dogs playing rough-and-tumble scored higher for Amenability and lower on Separation-Related Behavior than did dogs playing other types of games. Dogs playing tug-of-war and fetch scored high on Confident Interactivity. Winning or losing these games had no consistent effect on their test scores. If the dog started the majority of the games, the dog was significantly less amenable and more likely to exhibit aggression. The results suggest that how dogs play reflects general attributes of their temperament and relationship with their owner. This study provides no evidence that games play a major deterministic role on dominance dimensions of dog-human relationships, but the results suggest that playing games involving considerable body contact may affect attachment dimensions.     

Stray Dogs in an Island Community: A Case Study From New Providence, The Bahamas

The “stray dog problem ”in New Providence, The Bahamas, has been a recurring social complaint throughout the 20th century. This article looks at some aspects of the problem's background and reports the findings of a resident perception study. The results suggest that it is unlikely that short-term policies will be successful in eradicating free-roaming dogs from the island.     

Molecular structure in peripheral dog breeds: Portuguese native breeds as a case study

Genetic variability in purebred dogs is known to be highly structured, with differences among breeds accounting for ∼30% of the genetic variation. However, analysis of the genetic structure in non-cosmopolitan breeds and local populations is still limited. Nine Portuguese native dog breeds, and other peripheral dog populations (five) with regional affinities, were characterized using 16 microsatellites and 225 amplified fragment length polymorphism (AFLP) markers, and the pattern of genetic differentiation was investigated. Although the level of breed differentiation detected is below that of other dog breeds, there is in most cases a correlation between breed affiliation and molecular structure. AFLP markers and Bayesian clustering methods allowed an average of 73.1% of individuals to be correctly assigned to source populations, providing robust genotypic assessment of breed affiliation. A geographical genetic structure was also detected, which suggests a limited influence of African dogs on the Iberian breeds. The sampling effect on the estimation of population structure was evaluated and there was a 2.2% decrease in genetic differentiation among breeds when working animals were included. Genetic diversity of stray dogs was also assessed and there is no evidence that they pose a threat to the preservation of the gene pool of native dog breeds.     

Breed differences in dogs sensitivity to human points: A meta-analysis

The last decade has seen a substantial increase in research on the behavioral and cognitive abilities of pet dogs, Canis familiaris. The most commonly used experimental paradigm is the object-choice task in which a dog is given a choice of two containers and guided to the reinforced object by human pointing gestures. We review here studies of this type and attempt a meta-analysis of the available data. In the meta-analysis breeds of dogs were grouped into the eight categories of the American Kennel Club, and into four clusters identified by Parker and Ostrander [Parker, H.G., Ostrander, E.A., 2005. Canine genomics and genetics: running with the pack. PLoS Genet. 1, 507-513] on the basis of a genetic analysis. No differences in performance between breeds categorized in either fashion were identified. Rather, all dog breeds appear to be similarly and highly successful in following human points to locate desired food. We suggest this result could be due to the paucity of data available in published studies, and the restricted range of breeds tested.     

Imitation and emulation by dogs using a bidirectional control procedure

A successful procedure for studying imitative behavior in non-humans is the bidirectional control procedure in which observers are exposed to a demonstrator that responds by moving a manipulandum in one of two different directions (e.g., left vs. right). Imitative learning is demonstrated when observers make the response in the direction that they observed it being made. This procedure controls for socially mediated effects (the mere presence of a demonstrator), stimulus enhancement (attention drawn to a manipulandum by its movement), and if an appropriate control is included, emulation (learning how the environment works). Recent research with dogs has found that dogs may not demonstrate imitative learning when the demonstrator is human. In the present research, we found that when odors were controlled for, dogs imitated the direction of a screen-push demonstrated by another dog more than in a control condition in which they observed the screen move independently while another dog was present. Furthermore, we found that dogs would match the direction of screen-push demonstrated by a human and they were equally likely to match the direction in which the screen moved independently while a human was present.     

Selecting shelter dogs for service dog training

Service dogs are an essential aid to persons with disabilities, providing independence, mobility, and improved self-esteem. Because of these proven benefits, the growing use of service dogs is creating a demand and supply crisis. One major cause is the 50% average dropout rate for dogs selected for training. Weiss and Greenber (1997) recently found that a dog, successful on the most commonly used selection test items, was as likely to be either a poor or a good candidate for service work. The experiment presented here evaluated test items developed by the author in 15 years of experience with dogs. The test items were administered to 75 dogs from the Kansas Humane Society. Once tested, the dogs received obedience and retrieval training. The experiment assessed each dog on behavior over 5 weeks of training versus performance on each selection test item. A subset of the selection items, combined in a regression analysis, accounted for 36.4% of the variance with R = 0.603. This research also revealed a reliable test for dog aggression without risking injury to dog or tester. Items for testing included fear, motivation, and submission. Another set of selection items reliably predicted the trait of “high energy ”commonly described as “high strung. ”Future research should involve investigating the effectiveness of both cortisol levels and blood pressure in predicting traits to help strengthen the predictive value of the tool and then testing on dogs trained to be full service dogs.     

What’s in a Name? Effect of Breed Perceptions & Labeling on Attractiveness,Adoptions & Length of Stay for Pit-Bull-Type Dogs

Previous research has indicated that certain breeds of dogs stay longer in shelters than others. However, exactly how breed perception and identification influences potential adopters'' decisions remains unclear. Current dog breed identification practices in animal shelters are often based upon information supplied by the relinquishing owner, or staff determination based on the dog''s phenotype. However, discrepancies have been found between breed identification as typically assessed by welfare agencies and the outcome of DNA analysis. In Study 1, the perceived behavioral and adoptability characteristics of a pit-bull-type dog were compared with those of a Labrador Retriever and Border Collie. How the addition of a human handler influenced those perceptions was also assessed. In Study 2, lengths of stay and perceived attractiveness of dogs that were labeled as pit bull breeds were compared to dogs that were phenotypically similar but were labeled as another breed at an animal shelter. The latter dogs were called "lookalikes." In Study 3, we compared perceived attractiveness in video recordings of pit-bull-type dogs and lookalikes with and without breed labels. Lastly, data from an animal shelter that ceased applying breed labeling on kennels were analyzed, and lengths of stay and outcomes for all dog breeds, including pit bulls, before and after the change in labeling practice were compared. In total, these findings suggest that breed labeling influences potential adopters'' perceptions and decision-making. Given the inherent complexity of breed assignment based on morphology coupled with negative breed perceptions, removing breed labels is a relatively low-cost strategy that will likely improve outcomes for dogs in animal shelters.     

Dog Ownership in the West Indies: A Case Study From the Bahamas

ABSTRACT

Results from several studies in The Bahamas, which included 511 interviews with residents and data on 776 dogs, are reported. The composition and characteristics of the dog population are described. The majority of dogs, both owned and unowned, are local mongrels called “potcakes.” The median age of the dog population is three years. Potcakes are given a lower level of health care than dogs considered to resemble a recognised breed. Many dogs have no place in the household and are able to roam. The dynamics of the owned and unowned dog populations are derived, and abandoned dogs are shown to sustain the free-roaming population. The free-roaming dog population breeds throughout the year but it has a breeding cycle longer than six months, which is probably caused by its ill health. Despite the nuisance caused by unowned dogs, residents are tolerant towards them, and over half the households feed dogs they do not own. The changes in pet ownership required to increase the welfare of dogs are discussed.     

Assessment of the functionality of genome‐wide canine SNP arrays and implications for canine disease association studies

Domestic dogs share a wide range of important disease conditions with humans, including cancers, diabetes and epilepsy. Many of these conditions have similar or identical underlying pathologies to their human counterparts and thus dogs represent physiologically relevant natural models of human disorders. Comparative genomic approaches whereby disease genes can be identified in dog diseases and then mapped onto the human genome are now recognized as a valid method and are increasing in popularity. The majority of dog breeds have been created over the past few hundred years and, as a consequence, the dog genome is characterized by extensive linkage disequilibrium (LD), extending usually from hundreds of kilobases to several megabases within a breed, rather than tens of kilobases observed in the human genome. Genome‐wide canine SNP arrays have been developed, and increasing success of using these arrays to map disease loci in dogs is emerging. No equivalent of the human HapMap currently exists for different canine breeds, and the LD structure for such breeds is far less understood than for humans. This study is a dedicated large‐scale assessment of the functionalities (LD and SNP tagging performance) of canine genome‐wide SNP arrays in multiple domestic dog breeds. We have used genotype data from 18 breeds as well as wolves and coyotes genotyped by the Illumina 22K canine SNP array and Affymetrix 50K canine SNP array. As expected, high tagging performance was observed with most of the breeds using both Illumina and Affymetrix arrays when multi‐marker tagging was applied. In contrast, however, large differences in population structure, LD coverage and pairwise tagging performance were found between breeds, suggesting that study designs should be carefully assessed for individual breeds before undertaking genome‐wide association studies (GWAS).     

Do disrupted early attachments affect the relationship between guide dogs and blind owners?

The study investigated the affectional bond developed by dogs (Canis familiaris) towards their human companions during the selection process to become guide dogs and compared this bond with that formed by pet dogs with their owners. One hundred and nine dog-owner pairs were tested using a modified version of the Strange Situation Test: custody dogs-puppy walkers (n = 34), apprentice dogs-trainers (n = 26), guide dogs-blind owners (n = 25) and pet dogs-owners (n = 24).

Twenty-six behaviours were scored using a 5 s point sampling method and two vocal behaviours were recorded as bouts. Factor analysis carried out on 24 mutually exclusive behaviours highlighted two different profiles of response. A relaxed reaction characterised by a high play activity was distinctive of custody and apprentice dogs, whereas an anxious reaction characterised by a high degree of proximity seeking behaviours was distinctive of pet dogs. Guide dogs were intermediate between these two extremes, expressing their attachment to the owners but showing a more controlled emotional reaction. This finding suggests that guide dogs can be viewed as “working pets”. Furthermore, the experimental set-up, characterised by the presence of a frightening stimulus, revealed that untrained dogs (pets and custody dogs) were more fearful than trained dogs (guide dogs and apprentice dogs). Finally, differences in temperament emerged between retrievers: Golden retrievers showed a higher level of affection demand while Labrador retrievers were more playful. Overall, these findings show that in spite of separations from previous attachment figures, guide dogs established with their blind owner a rather good and secure affectional bond.     

Studies of modern Italian dog populations reveal multiple patterns for domestic breed evolution

Through thousands of years of breeding and strong human selection, the dog (Canis lupus familiaris) exists today within hundreds of closed populations throughout the world, each with defined phenotypes. A singular geographic region with broad diversity in dog breeds presents an interesting opportunity to observe potential mechanisms of breed formation. Italy claims 14 internationally recognized dog breeds, with numerous additional local varieties. To determine the relationship among Italian dog populations, we integrated genetic data from 263 dogs representing 23 closed dog populations from Italy, seven Apennine gray wolves, and an established dataset of 161 globally recognized dog breeds, applying multiple genetic methods to characterize the modes by which breeds are formed within a single geographic region. Our consideration of each of five genetic analyses reveals a series of development events that mirror historical modes of breed formation, but with variations unique to the codevelopment of early dog and human populations. Using 142,840 genome‐wide SNPs and a dataset of 1,609 canines, representing 182 breeds and 16 wild canids, we identified breed development routes for the Italian breeds that included divergence from common populations for a specific purpose, admixture of regional stock with that from other regions, and isolated selection of local stock with specific attributes.     

Comparison of Adoption Agency Breed Identification and DNA Breed Identification of Dogs

Governmental and other agencies may require dog caregivers (owners) to provide breed identification of their dogs. This study compares breed identification by adoption agencies with identification by DNA analysis in 20 dogs of unknown parentage. Of the 20 dogs who had been adopted from 17 different locations, the study identified 16 dogs as having (or probably having) 1 or 2 specific breed(s) in their ancestry. DNA analysis of these dogs indicated that 25% (4/16) did in fact contain genetic evidence of an adoption agency's identified breed as one of the predominant breeds in a dog's ancestry. DNA analysis did not detect all specified breeds in 14 of these dogs. That is, 87.5% of the dogs identified by an adoption agency as having specific breeds in their ancestry did not have all of those breeds detected by DNA analysis. The discrepancies between opinions of adoption agencies and identification by DNA analysis suggest that it would be worthwhile to reevaluate the reliability of breed identification as well as the justification of current public and private policies pertaining to specific dog breeds.     

Behavioral evaluation and demographic information in the assessment of aggressiveness in shelter dogs

Behavioral evaluations of 2017 shelter dogs were used for identifying dogs with aggressive tendencies and for predicting post-adoption behavior problems. Associations between failure of the behavioral evaluation and demographic factors (age, breed, and sex) and the dog's behavioral history, evaluated by logistic regression, were highly significant (P < 0.0005), except for “old” dogs (age > 72 months). Dogs that failed the behavior evaluation were not placed for adoption; therefore it was not possible to study prospectively the capability of behavioral evaluation to predict future aggressiveness in these dogs. Instead we developed tests for classifying dogs as aggressive or not aggressive based on their demographic factors and behavior evaluation outcomes. The results were compared retrospectively to the dogs’ known behavioral histories, which were obtained at intake to the shelter. This allowed estimation of the sensitivity, specificity, and accuracy of the classification tests. The most significant postdictor (i.e., “retrospective predictor”) of aggressiveness was failure of the behavioral evaluation (odds ratio 11.83, P < 0.0005). Among dogs that passed the behavioral evaluation, classification as “unsocial” by the behavioral evaluation was associated with nonaggressive behavior problems reported at 6-month follow-up (P = 0.005), and classification as “borderline” was associated with return to the shelter for aggressive behavior (P = 0.028). Behavioral evaluations reduced the rate of returns of adopted dogs from 19% in the year prior to their institution to 14% (P = 0.001) during the 2-year study period. Returns for aggression were reduced from 5% to 3.5% (P = 0.05), with many fewer incidents of serious aggression. The majority of dogs (86%) that failed the behavioral evaluation failed multiple component tests, indicating general aggressive tendencies in these dogs. Together with demographic information and behavioral history, behavioral evaluations can help to improve decisions regarding the disposition of shelter dogs.     

CRANIAL MORPHOLOGY OF DOMESTIC AND WILD CANIDS: THE INFLUENCE OF DEVELOPMENT ON MORPHOLOGICAL CHANGE

The domestic dog varies remarkably in cranial morphology. In fact, the differences in size and proportion between some dog breeds are as great as those between many genera of wild canids. In this study, I compare patterns of intracranial allometry and morphologic diversity between the domestic dog and wild canid species. The results demonstrate that the domestic dog is morphologically distinct from all other canids except its close relatives, the wolf-like canids. Following this, I compare patterns of static and ontogenetic scaling. Data on growth of domestic dogs are presented and used to investigate the developmental mechanisms underlying breed evolution. Apparently, most small breeds are paedomorphic with respect to certain morphologic characters. In dogs and other domestic animals, morphologic diversity among adults seems to depend on that expressed during development.     

The Effect of Domestication on Inhibitory Control: Wolves and Dogs Compared

Inhibitory control i.e. blocking an impulsive or prepotent response in favour of a more appropriate alternative, has been suggested to play an important role in cooperative behaviour. Interestingly, while dogs and wolves show a similar social organization, they differ in their intraspecific cooperation tendencies in that wolves rely more heavily on group coordination in regard to hunting and pup-rearing compared to dogs. Hence, based on the ‘canine cooperation’ hypothesis wolves should show better inhibitory control than dogs. On the other hand, through the domestication process, dogs may have been selected for cooperative tendencies towards humans and/or a less reactive temperament, which may in turn have affected their inhibitory control abilities. Hence, based on the latter hypothesis, we would expect dogs to show a higher performance in tasks requiring inhibitory control. To test the predictive value of these alternative hypotheses, in the current study two tasks; the ‘cylinder task’ and the ‘detour task’, which are designed to assess inhibitory control, were used to evaluate the performance of identically raised pack dogs and wolves. Results from the cylinder task showed a significantly poorer performance in wolves than identically-raised pack dogs (and showed that pack-dogs performed similarly to pet dogs with different training experiences), however contrary results emerged in the detour task, with wolves showing a shorter latency to success and less perseverative behaviour at the fence. Results are discussed in relation to previous studies using these paradigms and in terms of the validity of these two methods in assessing inhibitory control.