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Breed differences in social cognition, inhibitory control, and spatial problem-solving ability in the domestic dog (Canis familiaris)

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The extraordinary genetic and behavioural diversity of dog breeds provides a unique opportunity for investigating the heritability of cognitive traits, such as problem-solving ability, social cognition, inhibitory control, and memory. Previous studies have mainly investigated cognitive differences between breed groups, and information on individual dog breeds is scarce. As a result, findings are often contradictory and inconsistent. The aim of this study was to provide more clarity on between-breed differences of cognitive traits in dogs. We examined the performance of 13 dog breeds (N = 1002 dogs) in a standardized test battery. Significant breed differences were found for understanding of human communicative gestures, following a human’s misleading gesture, spatial problem-solving ability in a V-detour task, inhibitory control in a cylinder test, and persistence and human-directed behaviour during an unsolvable task. Breeds also differed significantly in their behaviour towards an unfamiliar person, activity level, and exploration of a novel environment. No significant differences were identified in tasks measuring memory or logical reasoning. Breed differences thus emerged mainly in tasks measuring social cognition, problem-solving, and inhibitory control. Our results suggest that these traits may have come under diversifying artificial selection in different breeds. These results provide a deeper understanding on breed-specific traits in dogs.
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Breed dierences in social
cognition, inhibitory control,
and spatial problem‑solving
ability in the domestic dog (Canis
familiaris)
Saara Junttila
1*, Anna Valros
1, Katariina Mäki
2, Heli Väätäjä
3, Elisa Reunanen
4 &
Katriina Tiira
5,6
The extraordinary genetic and behavioural diversity of dog breeds provides a unique opportunity
for investigating the heritability of cognitive traits, such as problem‑solving ability, social cognition,
inhibitory control, and memory. Previous studies have mainly investigated cognitive dierences
between breed groups, and information on individual dog breeds is scarce. As a result, ndings are
often contradictory and inconsistent. The aim of this study was to provide more clarity on between‑
breed dierences of cognitive traits in dogs. We examined the performance of 13 dog breeds (N = 1002
dogs) in a standardized test battery. Signicant breed dierences were found for understanding of
human communicative gestures, following a human’s misleading gesture, spatial problem‑solving
ability in a V‑detour task, inhibitory control in a cylinder test, and persistence and human‑directed
behaviour during an unsolvable task. Breeds also diered signicantly in their behaviour towards an
unfamiliar person, activity level, and exploration of a novel environment. No signicant dierences
were identied in tasks measuring memory or logical reasoning. Breed dierences thus emerged
mainly in tasks measuring social cognition, problem‑solving, and inhibitory control. Our results
suggest that these traits may have come under diversifying articial selection in dierent breeds.
These results provide a deeper understanding on breed‑specic traits in dogs.
Cognitive abilities (such as learning, memory, inhibitory control, problem-solving, and social cognition) are
important traits in almost all aspects of an animal’s life, from nding food to cooperating with conspecics.
Despite this, the heritability of cognitive traits in non-human animals is still a largely unknown topic1. e
domestic dog (Canis familiaris), with its extraordinary genetic and phenotypic diversity, provides a unique oppor-
tunity for advancing our understanding of this subject. Behavioural variation between dog breeds is substantial2,
and there is evidence that many of these breed dierences are attributable to genetic factors3 (but see Morrill
etal.4).
Cognitive dierences between breeds have been investigated to some extent, but the results seem to be largely
contradictory and inconsistent. For example, studies have demonstrated dierences in breeds’ understanding of
human referential cues, such as pointing or gazing58, and genetic relatedness between breeds seems to account
for a substantial portion of variation in this trait9. Several other studies, however, have not been able to replicate
these results1013. Similarly, studies have failed to nd any signicant dierences between breeds for logical rea-
soning ability13, inhibitory control13, or memory12,13, although Gnanadesikan etal.9 demonstrated that genetic
relatedness among breeds accounted for a signicant proportion of variation in these traits.
e diculty in interpreting previous results lies partly in the fact that most studies have grouped breeds
together, either based on original function or genetic relatedness. What further complicates interpretation
is that across studies, groups are oen composed of dierent breeds, and the criteria for categorizing breeds
OPEN
1Department of Production Animal Medicine, University of Helsinki, 00014 Helsinki, Finland. 2International
Partnership for Dogs, Helsinki, Finland. 3Digital Solutions, Lapland University of Applied Sciences, Jokiväylä 11C,
96300 Rovaniemi, Finland. 4Department of Finnish and Finno-Ugric Languages, University of Turku, 20014 Turku,
Finland. 5smartDOG Ltd, 05800 Hyvinkää, Finland. 6Department of Equine and Small Animal Medicine, University
of Helsinki, 00014 Helsinki, Finland. *email: saara.junttila@helsinki.
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dier greatly. Some authors have questioned the use of breed group classications based on the breeds’ original
purpose1417, mainly because selection pressures may have changed drastically. For instance, Svartberg17 found no
associations between behavioural proles of breeds and their original purpose; instead, breed-typical behaviour
was correlated with the current use of the breed. Even grouping breeds based on their genetic relatedness is not
without its faults. For example, both Svartberg17 and Turcsán etal.16 used cluster analysis to group breeds based
on behavioural similarity, and found that these clusters corresponded poorly with the genetic categorization of
breeds. ese results suggest that when breeds are divided into groups, important dierences between breeds
may be missed.
Only a handful of studies have compared cognitive dierences between individual breeds instead of breed
groups7,8,12,14,1820. Unfortunately, most of these have involved restricted sample sizes and a limited number
of breeds. Moreover, very little empirical research has targeted non-social cognitive traits such as memory,
inhibitory control, spatial problem-solving, and logical reasoning. e aim of our study was to provide a more
complete picture of cognitive dierences between dog breeds. We explored breed dierences not only in dogs’
socio-cognitive abilities, but also in several cognitive traits not involving a social aspect. In addition, we com-
pared breeds regarding their exploration in a novel space, greeting an unfamiliar person, and activity level, since
these may also be linked to cognitive performance2124. Our large sample size allowed us to look at dierences
between individual breeds rather than having to resort to breed group classications. Our aim was to provide
a more comprehensive understanding of how breeds vary cognitively and behaviourally, which improves our
ability to predict how individual dogs are likely to behave. Our results may also further our understanding of
the heritability and genetics of cognition, both in humans and in other animals.
Methods
Subjects. A total of 2,352 adult dogs participated in a smartDOG™ test battery25 between March 2016 and
February 2022. Participating dogs were required to be interested in working for food, and to not be overly
aggressive to people. We limited the analysis to dogs between the ages of 1 and 8years, since cognitive traits
may not have fully developed in younger dogs26,27, while older dogs may experience cognitive decline28. We
included only dog breeds with a minimum of 40 individuals tested per breed. is resulted in a nal sample size
of 1,002 dogs representing 13 breeds, including one category consisting of mixed breeds. Any individual which
had parents belonging to dierent breeds was classied as mixed breed, except for the Labradoodle. Details of
participant dogs’ ages, sexes, and breeds are presented in Table1.
Participant dogs took part in a test battery involving multiple tests, but not all dogs had results for every test
included in this study (see Supplementary TableS1 for number of participant dogs in each test section). Most
of the results for each test came from the same dogs, with some results missing, mostly because 17% of the dogs
took part in a shortened version of the test battery (see Supplementary Materials S1 for details).
Most participant dogs were privately owned pet dogs. We did not have information on the training history of
most dogs, apart from 31 police dogs (see Suppl. TableS2). A large proportion of the pet dogs were actively used
in various dog sports (e.g., agility, scent work, obedience, etc.), based on discussions the testers have had with
participating owners. Most dogs (including the police dogs) lived inside the house with their owners.
Cognitive test battery. Dogs included in the study were participants in a commercial cognitive test bat-
tery (smartDOG™)25, which was developed by one of the authors (KT) based on previous scientic publications.
Tests were performed by eight trained female smartDOG licence testers (including KT) at testing sites across
Finland. We included 10 tests, seven of which measured cognitive traits and three of which measured behaviour.
Descriptions of included tests are outlined in Table2.
All tests involved solving various problems with food rewards. e owners were asked to bring the dog’s
favourite treats, which were then used as rewards. In some cases, a toy was also used, if the dog was more
Table 1. Number of dogs within each breed and their sexes and median ages.
Breed n Males (%) Females (%) Median age in years ± IQR
Australian Kelpie 41 20 (48.8) 21 (51.2) 3.0 ± 2.8
Australian Shepherd 49 23 (46.9) 26 (53.1) 2.9 ± 2.6
Belgian Shepherd Malinois 49 34 (69.4) 15 (30.6) 3.6 ± 2.6
Border Collie 106 48 (45.3) 58 (54.7) 2.6 ± 2.3
English Cocker Spaniel 60 29 (48.3) 31 (51.7) 2.7 ± 2.7
Finnish Lapphund 59 28 (47.5) 31 (52.5) 2.8 ± 3.4
German Shepherd 82 34 (41.5) 48 (58.5) 2.6 ± 2.9
Golden Retriever 74 35 (47.3) 39 (52.7) 2.8 ± 2.5
Hovawart 50 23 (46.0) 27 (54.0) 2.7 ± 2.9
Labrador Retriever 163 69 (42.3) 94 (57.7) 2.6 ± 2.3
Mixed Breed 149 54 (36.2) 95 (63.8) 3.3 ± 2.8
Shetland Sheepdog 48 19 (40.4) 28 (59.6) 3.3 ± 2.9
Spanish Water Dog 72 22 (30.6) 50 (69.4) 2.8 ± 2.5
Tot a l 1 002 438 (43.8) 563 (56.2) 2.9 ± 2.7
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motivated by toys than food. e owner was advised not to feed their dog prior to the test to ensure food moti-
vation during the test. Fresh drinking water was available throughout the test. Testing took place indoors, and
the minimum size of the testing room was approximately 30 m2. Most oen only the tester, owner, and dog were
present during testing, but occasionally family members were present as observers. e dog was o the lead
throughout the test battery. A short break was included in the COGNITION test battery, during which the owner
was asked to walk the dog outside for a maximum of two minutes.
e order of tests was the same for all dogs to ensure consistency across subjects. Each test included several
trials. At the beginning of each trial, the tester always drew the dog’s attention, if necessary, by showing them a
treat and saying the dog’s name. A (Finnish) word, such as “ok” or similar, was used to indicate to the owner the
beginning of each trial. e owner was advised to release the dog upon hearing this word during the test battery.
Aer each trial, the owner called or moved the dog back to the starting position using the collar and held the
dog in place before releasing the dog for the next trial. e owner was advised to remain quiet and neutral when
the dog was focusing on a task, but they were allowed to praise the dog when the dog ate a treat. Only during the
V-detour was the owner allowed to encourage the dog to continue if the dog continuously stared at the owner.
Between test sections, the dog was free to walk around for about 1–2min.
A short description of each test is provided below, and more detailed information can be found in the Sup-
plementary Information S1. An example of each cognitive test can also be found in the supplementary videos S1.
Greeting. When the owner and dog rst entered the test room, the dog’s response to the tester (an unknown
person) was rated. e tester faced the dog while talking in a friendly voice and allowed the dog to approach
herself. If the dog was not fearful or aggressive, the tester bent down and attempted to stroke the dog. e tester
then continued stroking the dog as long as the dog was willing. is test lasted between 1–2min in total.
e tester rated the dog’s response according to a scale ranging from 1 to 7. For analysis purposes, these
scores were condensed into four groups: ‘fearful’ (score 1–3), ‘indierent’ (score 4–5), ‘friendly’ (score 6), and
overexcited’ (score 7). e largest category (‘friendly’) was used as the reference category.
Activity level. Aer the tester had greeted the dog, she attached a FitBark (FitBark Inc., Kansas City, MO,
USA) activity monitor to the dog’s collar or harness. FitBark generates ‘BarkPoints’ (from here on referred to as
‘activity points’), which represent a proxy measure for the average activity level of the dog during the test battery.
e monitor was kept on throughout testing and taken o when the test battery ended. Only dogs which had
taken part in the COGNITION test battery were analysed.
Exploration. Aer the FitBark had been attached, the dog was released and allowed to freely explore the test
room for approximately ve minutes. e tester rated the dog’s behaviour on a scale of 1–5. ese scores were
condensed into four groups: ‘low investigation’ (score 1–2), ‘moderate investigation’ (score 3), active investiga-
tion, walking (score 4), and very active investigation, running (score 5). e aim of this test was to measure the
extent to which the dogs were willing to investigate a novel environment, which is thought to be an indication of
Table 2. Description of each test included in the study, in the order in which they were presented to the dog
during the COGNITION test battery.
Tes t Trait(s) measured Short description
Greeting Greeting behaviour towards an unfamiliar person e dog’s behaviour upon rst greeting the tester was rated on a scale of 1–7
Activity level Activity level during the test e dog was supplied with a FitBark57 accelerometer for the duration of the
test battery, which provided an average activity level for the dog
Exploration Dog’s exploration of a novel environment e dog’s behaviour during its rst few minutes in the testing environment
was rated on a scale of 1–5
Cylinder test Inhibitory control, impulsivity Food is placed inside a transparent cylinder, and the dog has to inhibit their
response to reach directly for the food, and instead go around the barrier to
reach the food
Gestures Social cognition, dog’s ability to understand human communicative cues Object choice test: the human provides a gesture towards the bowl which
contains food. Gestures included: dynamic distal pointing, momentary distal
pointing, dynamic foot pointing, dynamic cross-forward pointing, and gaze
V-detour Inhibitory control, spatial problem-solving ability e dog is required to reach a food reward by detouring around a V-shaped
fence
Unsolvable task Social cognition, help-seeking behaviour, persistence, problem-solving
strategy
e dog is faced with a problem where food is visible but out of reach. e
dog’s response was measured as (a) human-directed behaviour, (b) independ-
ent behaviour, attempting to solve the task, and (c) abandoning the task;
engaging in behaviours not directed at a human or the apparatus
Logical reasoning Logical reasoning, ability to make inferences based on exclusion e dog can see that one of two bowls is empty and has to infer that the treat
is hidden under the other bowl
Memory vs gesture Dog’s tendency to choose based on human gesture vs visual information,
social cognition
e dog is required to choose between two bowls. e dog can see a human
placing a treat in one bowl, and the human gestures towards the other (empty)
bowl. e dog’s choice is then observed
Memory Spatial short-term memory e dog was required to remember the location of a food treat which was
hidden under one of three bowls, for an increasing duration of time (from 1
to 2.5min)
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curiosity, boldness, and activity level, whereas remaining by the owner’s side was thought to be a possible indica-
tion of fear, anxiety, or neophobia.
The cylinder test. e cylinder test has been used extensively in animal cognition research to study impul-
sivity and inhibitory control29, more specically the motor inhibitory response. Inhibitory control is a core exec-
utive function, which involves suppression of a prepotent but inecient behaviour in favour of a more benecial
response. In this test, the dog is required to inhibit reaching directly for a visible food reward and instead detour
around the transparent barrier to reach the reward.
e owner and dog were positioned 2–2.5m away from the cylinder. During the training trials, the cylinder
was opaque, and the dog was taught to access a food reward from either of the open sides. e experimenter
stood directly behind the cylinder and placed a food reward inside while the dog was watching. e dog was then
released and allowed to eat the treat (Supplementary Video S1). Aer the dog fullled the learning criteria (4 out
of 5 trials without touching the outside of the cylinder), the test phase began. During the test phase the cylinder
was transparent, and the dog was required to inhibit reaching for the now visible food directly, and to instead
go around to the side of the cylinder to access the reward (Supplementary Video S2). Each trial during which
the dog touched the outside of the cylinder was marked as an incorrect trial. If the dog ate the food without rst
touching the outside of the cylinder, this was marked as a correct trial. Percentage of correct responses (out of a
total of 10 trials) was used as the response variable.
Human gestures. Dogs’ understanding of pointing and other human gestures is oen used as a measure of
social cognition30,31. e test battery included ve dierent gestures. Before the test phase, the dog was familiar-
ized with the test set-up over four training trials, during which the dog was simply taught that food is available
in either of the two bowls. (See Supplementary Video S3 for the training phase procedure.).
During the test phase, dogs took part in six trials for each gesture (30 trials in total). (See Supplementary
Video S4 for the test procedure for each of the ve gestures.) e percentage of correct responses (out of 30
trials) was calculated from all the gesture tests combined. e order of rewarded bowls was the same for each
dog, starting with the le side. Every other trial was rewarded to the right and every other to the le. In order
to ensure dogs were not learning this pattern, the percentage of correct responses from the nal 6 trials (gaze)
were compared to the percentage of correct responses from the rst 6 trials (dynamic distal pointing) using a
two-tailed paired t-test.
e trial always started with the dog and owner facing the tester, standing 2–2.5m away. e tester showed a
piece of food to the dog and placed it inside one of the bowls which she held in her hands. e bowls were placed
on the oor in front of the tester, 95cm apart from each other. Making sure the dog was watching, the tester
provided the gesture (these are described in more detail below). Aer this, the dog was released and allowed to
make a choice.
e procedure for each gesture was the same as described above. Each dog received the gestures in the same
order (6 trials each): (1) dynamic distal pointing: the tester pointed at the correct bowl with an extended arm
and index nger aer which the dog was released, and the tester kept her arm in the same position while the dog
made their choice, (2) momentary distal pointing: the tester pointed at the correct bowl with an extended arm
and index nger for a duration of 2s, aer which the tester lowered her arm and the dog was released while the
tester’s arms were ush at her sides, (3) dynamic proximal foot pointing: the tester placed the tip of her foot on
the ground directly behind the correct bowl, and the dog was released while the tester remained in this position,
(4) dynamic cross-forward pointing: THE tester used her contralateral arm to point at the correct bowl while
rotating her shoulders in the same direction, sustaining this position while the dog made their choice, (5) gaze:
the tester alternated her gaze between the dog and the bowl three times, and the dog was released while the tester
maintained her gaze on the correct bowl.
V‑detour. e V-detour has been used in canine cognitive research to investigate spatial problem-solving
ability29. e dog has to detour around a transparent V-shaped fence to access a food reward which is placed on
the other side (Supplementary Video S5). Since the dog is required to move away from the visible treat to access
it, the task is also oen considered to measure inhibitory control.
e V-shaped fence was made out of compost fence panels, which were attached at an angle of approximately
70°. e owner and dog waited about 40cm away from the intersecting angle of the V-shaped fence. e tester
showed the dog several treats (or a toy) and placed them inside the fence while standing outside the fence. e
owner released the dog while the dog was looking at the food. e number of seconds taken to solve the task
was measured using a stopwatch. If the dog was not able to solve the task within 3min, the trial was terminated.
Unsolvable task. e unsolvable task has been used in canine cognitive research to assess persistence,
problem-solving behaviour, human-directed communication, and social cognition32. In our version of this task,
the dog was presented with four solvable trials, aer which the task became impossible to solve (Supplementary
Video S6).
e test involved a plastic or wooden box with a transparent lid, which had small holes to allow the dog to
smell the food inside. During the four training trials, the dog was taught to access a treat placed inside the box by
moving the plastic lid o. e diculty of the trials increased gradually. Once the dog was successful in opening
the lid, the test trial was begun. With the dog watching, the tester placed several treats inside the box. She then
secured the lid in place so that it could not be opened, aer which the owner released the dog. Both owner and
experimenter remained quiet and still, looking only at the box during the subsequent 2-min period. e tester
measured the time the dog spent on each behaviour: (a) independent problem-solving: attempting to solve the
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task independently, (b) human-directed behaviour: initiating social contact with either the tester or the owner,
or (c) abandoning the task: not directing their behaviour toward the task or a person.
ree variables were used to measure the dogs’ behaviour during the unsolvable task. (1) Complete inde-
pendence (comparing dogs which did not spend any time on human-directed behaviour to those which spent
any amount of time on human-directed behaviour), (2) Percentage of time spent on human-directed behaviour,
and (3) Abandoning task (comparing dogs which abandoned the task to those which never abandoned the task).
Logical reasoning. is test aimed to measure the dog’s ability to make inferences based on exclusion33.
e dog could see that one of two bowls was empty, and it had to infer that the treat was hidden under the other
bowl. e tester sat on a chair or on the oor, about 1m away from the dog. Two opaque bowls were placed
upside down in front of the tester at arm’s length, one on the le and one on the right side. In each trial, a piece
of food was placed under one of the bowls. e order of baiting was the same for each dog; rst the le-hand
bowl was baited, aer which every other trial was baited to the right and every other trial to the le. e dog
rst took part in a training phase consisting of four trials, the aim of which was to familiarize the dog with the
set-up and learn that there is always food hidden under one of the bowls. (See Supplementary Video S7 for the
procedure of the training phase.).
When the dog correctly performed all four training trials, the test phase was initiated. is consisted of six
trials. e tester held a writing pad as a visual barrier in front of the le-hand bowl and placed a treat under the
bowl. She then placed the writing pad in front of the right-hand bowl and sham-baited the bowl. e writing
pad was then removed, and the tester lied the empty bowl up about 30cm above the oor, keeping it there for
about 1–2s while the dog was watching. At the same time, the tester held her other hand on top of the baited
bowl. e bowl was then placed back down, the tester placed her hands on her lap, and the dog was released. If
the dog approached the correct bowl, they were allowed to eat the treat. e trial was then repeated another ve
times, every other trial rewarded on the right and every other on the le. (See Supplementary Video S8 for the
procedure of the test phase.).
Two variables were used to measure logical reasoning of dogs: (a) percentage of correct responses and (b)
understanding of the task. For percentage of correct responses, dogs were divided into three groups: (1) 0–50%
of trials correct, (2) 51–82% of trials correct, and (3) 83–100% of trials correct. e tester also evaluated whether
the dog had understood the task and was successfully making inferences based on exclusion. e dog was con-
sidered to have understood the task if at least 4 out of 6 trials were correct or the nal 2–3 trials were correct.
Memory vs gesture. Previous studies have shown that dogs are more likely to choose an empty bowl out of
two choices if a human points towards it, even when they have seen that the other bowl has food in it3437. Similar
to the gesture tests, this test aims to measure social cognition. e procedure for this test was also similar to the
gesture test, but instead of gesturing towards the baited bowl, the tester gestured towards the empty bowl. e
tester used the gesture which the dog had been most successful with—this was oen the dynamic distal point.
e bowls remained on the oor throughout the test, and the treat was placed in one of the bowls while the dog
was watching. e test consisted of two trials, with the rst trial always baited to the le and the second trial
baited to the right. (See Supplementary Video S9 for the procedure.) Dogs were divided into two groups: those
which chose the baited container on both trials (relied on their memory), and those which chose the empty
container on 1–2 trials (relied on the human’s gesture).
Memory. e aim of this test was to measure the duration of the dogs’ short-term memory38. ree identical
opaque bowls were placed upside down on the oor in a straight line, about 1m apart from each other. A piece
of food (or a toy) was placed under one of the bowls in each trial. e owner sat on a chair 3m away from the
middle bowl with the dog in front of her. During 7 training trials the dog learned to nd a food reward from
under one of the three bowls. Only a very short delay between hiding the food and releasing the dog was in place
during this phase.
Once the dog had passed the training phase, the four test trials began. e tester placed a treat under one
bowl while the dog was watching, aer which a visual barrier was placed in front of the dog. e tester waited
an increasing duration of time behind the owner and the dog (1st trial: 1min, 2nd trial: 1.5min, 3rd trial: 2min,
4th trial: 2.5min). Aer the waiting period, the tester removed the barrier, and the dog was released and allowed
to make a choice. e order of baiting the bowls was the same for each dog: trial 1: middle, trial 2: le, trial 3:
right, and trial 4: middle. (See Supplementary Video S11 for the procedure of the test phase.) Number of correct
trials (out of a total of 4) was calculated for each dog. Since a very small number of dogs had a score of 0, these
dogs were combined into a group with dogs that had a score of 1.
Data analysis. All statistical analyses were performed using IBM SPSS Statistics Version 28. An alpha level
of 0.05 was used for all statistical tests. Multiple and logistic regression analyses were used with the enter method
to analyse dierences between breeds for each variable. For ordinal variables, cumulative odds ordinal logistic
regression with proportional odds was used. In each model, we included age and sex of the dogs as control vari-
ables, since previous research suggests these may aect various measures of cognition and behaviour3942. Each
model therefore included the predictors breed, age, and sex (apart from success in the V-detour, which only
included breed as a predictor). e Labrador Retriever, one of the most popular breeds worldwide, was used
as the reference breed, since it had the largest number of individuals tested out of all included pedigree breeds.
For the greeting variable, the data failed the assumption of proportional odds according to the full likelihood
ratio test, and therefore a multinomial logistic regression was conducted instead of ordinal logistic regression. For
success in the V-detour task, Fisher’s Exact test using the Monte Carlo method was run with 10,000 simulations,
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since an insucient number of individuals failed to solve the task for a binary logistic regression to be reliable. For
latency (s) to solve the V-detour task, the dependent variable was log-transformed to normalize its distribution.
Since the cylinder test variable ‘percentage of correct responses’ was negatively skewed, a reect and square root
transformation was applied (i.e., each data point was subtracted from the maximum value plus 1, and a square
root transformation was then applied to these scores). Due to the transformation, the variable was inverted, and
therefore named ‘percentage of incorrect responses’ to aid with interpretability. For all transformed variables, the
original values are included in the gures, whereas the transformed variables are reported in the tables and text.
Ethical statement. We conrm that the procedures comply with national and EU legislation. Research was
performed in accordance with the Declaration of Helsinki. e study was approved by the University of Helsinki
Viikki Campus Research Ethics Committee (Statement 12/2021, accepted on 18/05/2021). Before participating
in the cognitive test battery, each dog owner gave informed written consent for using their dogs’ test results in
research. Reporting of results follows the recommendations of the ARRIVE guidelines. Informed consent was
given by each subject for publication of identifying images/videos in an online-access publication.
Results
Greeting. A total of 934 dogs had results for the greeting test. Out of these, 12.8% were fearful when meeting
the unknown person, 28.2% were indierent, 41.1% responded in a friendly manner, and 11.1% of dogs had an
overexcited greeting.
e multinomial regression model signicantly predicted greeting scores over and above the intercept-
only model (χ2(42) = 174.1, p < 0.001). e model explained 18.8% (Nagelkerke R2) of the variance in greeting
behaviour. Sex did not have a signicant eect on the prediction of greeting (χ2(3) = 2.65, p = 0.45), but age did
(χ2(3) = 20.77, p < 0.001). An increase in age was associated with a decrease in the odds of having an overexcited
greeting compared to a friendly greeting (which was used as the reference group) (χ2(1) = 9.94, p = 0.002).
Breed had a signicant eect on the prediction of greeting score (χ2(36) = 152.86, p < 0.001). See Fig.1a for the
proportions of greeting scores for each breed, and Supplementary TableS3 for parameter estimates for each
greeting score.
Activity level. A total of 759 dogs had activity level results available. Dogs received a mean of 29.4 FitBark
activity points, ranging from 9.9 to 93.4 activity points. e multiple regression model signicantly predicted
activity levels of dogs (F (14) = 7.91, p < 0.001, adj. R2 = 0.12). ere were 10 outliers but none of these were inu-
ential according to Cook’s Distance. Age and sex did not signicantly predict activity level in dogs aged 1–8years
(age β = −0.04, p = 0.23, sex β = −0.03, p = 0.42). Six breeds diered signicantly from the Labrador Retriever
(which was used as the reference breed for all analyses). See Fig.1b for mean activity points for each breed, and
Supplementary TableS4 for parameter estimates.
Exploration. A total of 820 dogs had results for exploration in a novel environment. Out of these, 8.3%
showed low investigation, 15.8% showed moderate investigation, 39.7% showed active investigation (walking),
and 36.3% showed very active investigation (running).
e ordinal regression model signicantly predicted exploration scores over and above the intercept-only
model (χ2(14) = 61.98, p < 0.001). e model explained 8% (Nagelkerke R2) of variance in exploratory behav-
iour. Age and sex did not have a signicant eect on the prediction of exploration in dogs aged 1–8years (age
χ2(1) = 1.9, p = 0.17; sex χ2(1) = 0.05, p = 0.82), but breed did (χ2(12) = 57.68, p < 0.001). See Fig.1c for percentage
of each exploration score for each breed, and Supplementary TableS5 for parameter estimates.
Cylinder test. A total of 992 dogs took part in the cylinder test. e median success rate of dogs was 80%
(IQR = 30%). e multiple regression model signicantly predicted the percentage of incorrect trials in the cylin-
der test (F (14, 961) = 6.96, p < 0.001, adj. R2 = 0.08). No outliers were detected. Age was a signicant predictor of
performance in the cylinder test, β = 0.13, p < 0.001, with increasing age associated with an increasing percentage
of incorrect trials (in dogs aged 1–8-years old). Sex was also a signicant predictor of cylinder test performance,
β = −0.09, p = 0.004, with females making fewer mistakes than males. Five breeds had signicantly lower per-
centages of incorrect trials compared to the Labrador Retriever. See Fig.2 for the mean percentage of correct
trials for each breed, and Supplementary TableS6 for parameter estimates.
Human gestures. A total of 831 dogs took part in each of the ve gesture tests. e mean percentage of
correct trials for these dogs was 79% (SD = 12.9%). A two-tailed paired samples t-test showed that there was no
signicant change from the rst 6 object choice trials to the last 6 trials, with a mean decrease of 0.84% in cor-
rect responses (95% CI [−2.65, 0.96], t(831) = −0.92, p = 0.36, d = -0.03). erefore, it is unlikely that dogs were
learning the pattern for baiting the bowls.
e multiple regression model signicantly predicted the percentage of correct trials for the gesture tests
(F (14, 804) = 3.41, p < 0.001, adj. R2 = 0.04). ere were two outliers, but these were not inuential according
to Cook’s Distance. Age and sex did not signicantly predict performance in dogs aged 1–8years (age β = 0.03,
p = 0.45; sex β = −0.06, p = 0.12). ree breeds had signicantly lower scores than the Labrador Retriever. See
Fig.3 for the mean percentage of correct trials for each breed, and Supplementary TableS7 for parameter
estimates.
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Figure1. Breed dierences of greeting unfamiliar person, activity level, and exploration in novel environment. Signicant
P-values (Bonferroni-corrected) are indicated with asterisks: ***p ≤ 0.001, **p 0.01, *p ≤ 0.05. e Labrador Retriever was
used as the reference breed. (a) Percentage of dogs within each breed receiving each of the four greeting scores (n = 934).
Breeds have been ordered based on odds ratios, from most fearful (highest odds ratio for score 1–3) on the le to least fearful
(lowest odds ratio for score 1–3) on the right. P-values have been indicated for fearful response using asterisks. (b) Mean
activity level scores (measured in FitBark activity points) for each breed (N = 759). Error bars represent 95% condence
intervals. Breeds are ordered based on B-values, with lowest activity levels on the le and highest activity levels on the right.
(c) Percentage of dogs within each breed with each exploration score (N = 820). Breeds are ordered based on B-values, with
lowest exploration on the le and highest exploration on the right. Score 1–2: low investigation (stays close to owner). Score 3:
moderate investigation. Score 4: active investigation (walking). Score 5: very active investigation (running).
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Figure2. Mean percentage of correct trials for each breed in the cylinder test are presented, using the
untransformed, original data (n = 992). Error bars represent 95% condence intervals. Breeds are ordered
based on B-values, with lowest success (low inhibitory control) on the le and highest success (high inhibitory
control) on the right. e Labrador Retriever was used as the reference breed. Signicant P-values (Bonferroni-
corrected) are indicated with asterisks: ***p ≤ 0.001, **p 0.01, *p ≤ 0.05.
Figure3. Mean percentage of correct trials in the gesture tests within each breed (n = 831). Error bars represent
95% condence intervals. Breeds have been ordered based on B-values, with lowest success on the le and
highest success on the right. e Labrador Retriever was used as the reference breed. Signicant P-values
(Bonferroni-corrected) are indicated with asterisks: ***p ≤ 0.001, **p 0.01, *p ≤ 0.05.
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V‑detour. A total of 993 dogs took part in the V-detour test. Out of these, 88.5% successfully solved the
V-detour within 3min. ere was no signicant dierence in the proportions of dogs within each breed suc-
ceeding compared to those failing (p = 0.13, 99% CI [0.12, 0.14]). See Fig.4a for the proportion of successful
dogs in each breed.
e median time for solving the V-detour task (when unsuccessful dogs were removed from the analysis)
was 12s (IQR = 29s). e multiple regression model signicantly predicted the time taken to solve the task
(n = 863, F (14, 848) = 2.52, p = 0.002, adj. R2 = 0.02). No outliers were detected. Age and sex did not signicantly
Figure4. Performance of dogs during the V-detour task. e Labrador Retriever was used as the reference
breed. Signicant P-values (Bonferroni-corrected) are indicated with asterisks: ***p ≤ 0.001, **p 0.01, *p ≤ 0.05.
(a) Percentage of dogs within each breed succeeding vs failing in the V-detour task (n = 993). Breeds have been
ordered based on B-values, with lowest succeeding breeds on the le and highest succeeding breeds on the right.
(b) Mean latency (s) to solve the V-detour for each breed are presented using the untransformed, original data
(n = 863). Error bars represent 95% condence intervals. Breeds are ordered based on B-values, with lowest
succeeding breeds (took a long time to solve the task) on the le and highest succeeding breeds (solved the task
quickly) on the right.
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predict performance in the V-detour in dogs aged 1–8years (age β = 0.05, p = 0.14; sex β = −0.05, p = 0.12). e
Border Collie had a signicantly lower predicted mean log score compared to the Labrador Retriever (β = −0.14,
p = 0.01). See Fig.4b for mean time taken to solve the task for each breed, and Supplementary TableS8 for
parameter estimates.
Unsolvable task. A total of 969 dogs took part in the unsolvable task. Only 57 (5.7%) of these were com-
pletely independent during the unsolvable task (i.e., they spent 0% of their time on human-directed behav-
iour). When comparing these dogs to those which spent over 0% of their time on human-directed behav-
iour, the binomial logistic regression model was statistically signicant (χ2(14) = 34.28, p = 0.002). e model
explained 9.7% (Nagelkerke R2) of the variance. Age was not a signicant predictor of independence in dogs
aged 1–8years (χ2(1) = 0.59, p = 0.44), but females had lower odds of being completely independent compared
to males (χ2(1) = 5.22, p = 0.02). Breed was also a signicant predictor of independence (χ2(12) = 23.13, p = 0.03),
but no breed was signicantly dierent from the Labrador Retriever. See Fig.5a for the percentage of completely
independent dogs within each breed, and Supplementary TableS9 for parameter estimates.
When completely independent dogs had been removed from the analysis, the median amount of time spent
on human-directed behaviour was 54.2% (65s out of a total of 120s) (IQR = 40%). e multiple regression model
signicantly predicted the percentage of time spent on human-directed behaviour (n = 912, F (14, 885) = 5.7,
p < 0.001, adj. R2 = 0.07). No outliers were detected. Age signicantly predicted time spent on human-directed
behaviour (β = 0.09, p = 0.009), with older dogs spending a larger proportion of their time on human-directed
behaviour. Sex did not signicantly predict time spent on human-directed behaviour (β = 0.002, p = 0.95). e
Golden Retriever and Australian Kelpie had signicantly higher scores than the Labrador Retriever. See Fig.5b
for the percentage of time spent on human-directed behaviour for each breed, and Supplementary TableS10
for parameter estimates.
A total of 37.2% of dogs spent over 0% of their time abandoning the task. When comparing these dogs to those
which persisted with the task (0% of time spent on abandoning the task), the binomial logistic regression model
was statistically signicant (χ2(14) = 40.89, p < 0.001). e model explained 5.7% (Nagelkerke R2) of the variance.
Age was a signicant predictor (χ2(1) = 16.34, p < 0.001), with older dogs being less likely to abandon the task.
Breed was a signicant predictor of abandoning the task, χ2(12) = 24.16, p = 0.02, but sex was not (χ2(1) = 0.32,
p = 0.57). No breed signicantly diered from the Labrador Retriever. See Fig.5c for the percentage of dogs within
each breed abandoning the task, and Supplementary TableS11 for parameter estimates.
Logical reasoning. A total of 826 dogs took part in the logical reasoning task. Out of these, 33.4% were
rated as having understood the logical reasoning task. When comparing dogs which understood the task to
dogs which failed to understand the task, the binomial logistic regression model was not statistically signicant
(χ2(14) = 20.95, p = 0.1). See Supplementary Fig.S5a for the percentage of dogs within each breed which under-
stood the task.
For percentage of correct trials, 60.9% of dogs were correct on 0–50% of trials, 22.6% of dogs were correct
on 51–82% of trials, and 16.5% of dogs were correct on 83–100% of trials. e ordinal regression model did
not signicantly predict success in the logical reasoning task above the intercept-only model, χ2(14) = 20.03,
p = 0.13. See Supplementary Fig.S5b for the percentage of dogs within each breed with dierent rates of success.
Gesture vs memory. A total of 823 dogs took part in the gesture vs memory test. Out of these, 40.5% fol-
lowed the human gesture to the baited container in at least one trial. e binomial logistic regression model
for trusting the human gesture was statistically signicant (χ2(14) = 29.23, p = 0.01). e model explained 4.8%
(Nagelkerke R2) of the variance in the gesture vs memory test. Age and sex were not signicant predictors in
dogs aged 1–8years, but breed was a signicant predictor of trusting the gesture, χ2(12) = 25.42, p = 0.01. See
Fig.6 for percentage of dogs within each breed which trusted the human gesture vs their own memory, and Sup-
plementary TableS12 for parameter estimates.
Memory. A total of 822 dogs took part in the memory test. Out of these, 14.1% were correct on 0–1 trials,
30.4% were correct on two out of four trials, 35.3% were correct on three trials, and 20.2% were correct on all
four trials. e ordinal logistic regression model did not signicantly predict the number of correct trials in the
spatial memory task above the intercept-only model (χ2(14) = 19.26, p = 0.16). See Supplementary Fig.S6 for the
percentage of dogs within each breed with dierent success rates.
Discussion
We found signicant dierences between individual dog breeds for ve of the seven cognitive tests included in
the test battery. Breed dierences were found for measures of social cognition, persistence, inhibitory control,
and spatial problem-solving ability. Dierences were also evident for activity level, greeting of an unfamiliar
person, and exploration of a novel environment. In contrast, no breed dierences were found for short-term
memory or logical reasoning.
Both inhibitory control and social cognition are likely to be especially important traits during articial selec-
tion of dog breeds, both historically and in the present day. For example, inhibitory control may be a valued
trait in herding dogs, which are required to inhibit their predatory responses. e Border Collie and Australian
Shepherd were among the highest-scoring breeds in the cylinder test, indicating high inhibitory control. In
contrast, the Malinois and German Shepherd were some of the lowest-scoring breeds. ese breeds are oen
used in working roles requiring high responsiveness, which is oen associated with low inhibitory control and
high impulsivity43,44. Human-directed behaviour and socio-cognitive abilities may be highly valued in pet dogs
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Figure5. Performance of breeds in the unsolvable task (n = 969). e Labrador Retriever was used as the reference breed.
Signicant P-values (Bonferroni-corrected) are indicated with asterisks: ***p ≤ 0.001, **p ≤ 0.01, *p ≤ 0.05. (a) Percentage
of dogs within each breed belonging to the human-directed compared to the completely independent groups during the
Unsolvable Task. Breeds have been ordered based on odds ratios, with most independent breeds (most likely to spend 0%
of their time on human-directed behaviour) on the le and least independent breeds (least likely to spend 0% of their time
on human-directed behaviour) on the right. (b) Mean percentage of time (out of a total of 120s) spent on human-directed
behaviour during the unsolvable task for each breed. Error bars represent 95% condence intervals. Breeds are ordered based
on B-values, with lowest amount of time spent on human-directed behaviour on the le and highest amount of time on the
right. (c) Percentage of dogs within each breed abandoning the task (over 0% of time spent on abandoning task) vs persisting
(0% of time spent on abandoning task) with the unsolvable task. Breeds are ordered based on odds ratios, with the most
persistent breeds (least likely to abandon task) on the le and the least persistent breeds (most likely to abandon task) on the
right.
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and breeds required to work closely with people, such as herding dogs and retrievers45,46. In line with this, the
Kelpie, Golden Retriever, Australian Shepherd, and Border Collie spent the largest proportion of their time on
human-directed behaviour during the unsolvable task. In contrast, the ability to work independently may be
important for various working dogs, such as detection dogs47,48. In our study, the two breeds which were most
likely to be completely independent during the unsolvable task (spending 0% of their time on human-directed
behaviour) were the German Shepherd and Malinois.
In conclusion, many of our results seem to reect the breeds’ original or current function, but several breed
dierences could not be easily explained by breed function alone. For example, many of the breeds in our study
belonged to the herding group, but there was oen large variation in their results in several cognitive tests. e
Finnish Lapphund received the lowest score in the gesture tests, whereas the Kelpie and Malinois had the highest
scores for this test, despite all three being herding breeds. Similarly, during the unsolvable task, the Australian
Shepherd was the least likely breed to abandon the task, whereas the Kelpie was among the breeds most likely to
abandon the task. It seems evident that breeds can vary behaviourally from each other even within their breed
groups, since dierent traits may have been (both intentionally and unintentionally) selected for in dierent
breeds, despite the breed group they belong to.
Our results replicate the ndings of various studies which have investigated dierences between individual
breeds in the V-detour task19, understanding of human gestures7,8, and the unsolvable task8,18. In contrast, most
studies which have failed to nd breed dierences have compared breed groups10,11,13,19,49,50. It is possible that
potential dierences between breeds may have been missed in these studies, since (as shown by our current
study) behavioural variation within breed groups can be substantial. For example, the Golden Retriever diered
signicantly from the Labrador Retriever in the unsolvable task and the gesture test, even though both belong
to the retriever group. Our ndings therefore highlight the importance of investigating behavioural dierences
between individual breeds rather than only relying on breed group categorizations.
We found no signicant breed dierences for logical reasoning or memory, and these results seem to be in
line with previous studies12,13. However, it is important to note that even though we did not nd dierences
between these specic breeds, this does not mean they do not exist between other breeds. For example, we did
not include ancient breeds, hounds, or terriers in our analysis. erefore, more research on individual breed
dierences on these cognitive traits is warranted.
It is important to note that the population used in this study is not representative of the entire dog population,
since ndings may dier across countries and cultures. In addition, since we used a commercial test battery, only
certain types of owners and dogs were inevitably self-selected. e breeds included in our study were mostly
breeds used in dog sports, and most owners were active in various dog sports or competitions. Even though this
limits the generalization of our results, it is noteworthy that signicant breed dierences emerged despite the
similarity of participant dogs and their training histories.
Figure6. Percentage of dogs within each breed which trusted the human gesture compared to those which
trusted their own memory (n = 823). Breeds have been ordered based on odds ratios, with breeds which are
most likely to trust their own memory on the le and breeds which are most likely to trust the human’s gesture
on the right. Signicant P-values (Bonferroni-corrected) are indicated with asterisks: ***p 0.001, **p 0.01,
*p ≤ 0.05. e Labrador Retriever was used as the reference breed.
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ere isa possibility that the dierences seen in our study were not based on genetic dierences between
breeds but rather due to variation in life experiences or training, since these have also been found to inuence
behaviour in cognitive tests8,13,49,51,52. Unfortunately, we were not able to control for the possible eects of train-
ing, environment, life experiences, or background of the dogs, since this information was not available to us.
erefore, the extent of their possible eects on breed dierences in our study is not known; however, this is
something we will investigate next. Previous studies suggest that environmental eects are unlikely to be the
only explanation for breed dierences. Genetic relatedness between dog breeds has been shown to account for
a substantial portion of variation in understanding human gestures and inhibitory control9. In addition, when
training history of participating dogs has been controlled for, signicant breed dierences have still been found
in the unsolvable task18,27,53, problem-solving tasks20,49, inhibitory control9, and point-following ability9,54. How-
ever, this topic warrants further investigation to determine the extent of heritable dierences between breeds as
opposed to the eect of life experiences.
It could be argued that the large proportion of police dogs within the Malinois breed (45%) could have biased
breed dierences in our study. However, our sample also consisted of other working dogs as well as several pet
dogs which participated extensively in training and competitions, but we did not have information about these
individuals. erefore, it would not have made much sense to exclude police dogs. We nevertheless analysed
the cylinder test, V-detour, and unsolvable task (which were the only tests which police dogs took part in) with
police dogs excluded. As a result, breed dierences in the proportions of completely independent dogs in the
unsolvable task became non-signicant, and the proportion of completely independent Malinois dropped from
14.6 to 7.7%. For all other variables breed dierences were still signicant and the scores of Malinois changed
only slightly (results not shown). erefore, it seems that the inclusion of police dogs only aected complete
independence during the unsolvable task but did not inuence other results to a great extent.
Another factor which could aect breed dierences is the brain size and skull shape of the breeds13,5456.
However, the breeds included in our study all had similar skull shapes, and there were no extremes in body size.
e Shetland Sheepdog and Cocker Spaniel were the only breeds which markedly diered in size from the others,
but they did not seem to consistently dier from the larger breeds in their results.
In conclusion, we found signicant breed dierences for various behavioural and cognitive traits in dogs.
is is one of the few studies investigating individual breed dierences in dogs, especially in non-social cogni-
tive traits which are rarely studied in this context. Our results provide a more complete picture of breed-typical
behaviour in dogs.
Data availability
e datasets analysed during the current study are not publicly available due to privacy restrictions but are avail-
able from the corresponding author on reasonable request.
Received: 20 September 2022; Accepted: 22 December 2022
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Acknowledgements
We thank all the owners and their dogs who participated in this research, as well as the smartDOG licence testers:
Katja Kontu, Liisa Tikka, Sanni Somppi, Kati Vierimaa, and Ulla-Marja Ruistola. We would also like to thank the
University of Helsinki and the Finnish Foundation of Veterinary Research for funding this study.Open access
funded by the Helsinki University Library.
Author contributions
K.T., H.V., and E.R. collected the data. A.V., K.M., and K.T. supervised the project. S.J. designed and performed
the analysis. S.J. wrote the manuscript. All authors discussed and reviewed the manuscript and contributed to
the nal manuscript.
Competing interests
KT is the founder and owner of smartDOG Ltd, and HV and ER are licence testers of the company. KM, SJ, and
AV have no aliation with smartDOG. SJ’s work has been funded by the the Finnish Foundation of Veterinary
Research. ere are no other conicts of interest.
Additional information
Supplementary Information e online version contains supplementary material available at https:// doi. org/
10. 1038/ s41598- 022- 26991-5.
Correspondence and requests for materials should be addressed to S.J.
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... Centuries of selective breeding have yielded a broad range of genetic and phenotypic variations in domestic dogs (Canis lupus familiaris) (Junttila et al., 2022;Sutter et al., 2004;Svartberg, 2006). This diversity has facilitated the involvement of dogs in a variety of roles within the human society, encompassing various functions such as herding, hunting, sledding, assistance to individuals with disabilities, and as family companions (Svartberg, 2006;Udell and Wynne, 2008). ...
... Variables with * are statistically significant. research studies (Gnanadesikan, Hare, Snyder-Mackler, Call, et al., 2020;Junttila et al., 2022), it is also plausible that genuine differences among the studied breed clades may manifest more within dogs' performances throughout the challenging, more cognitively demanding phases of the task (such as the Reversal learning phasesee Azadian and ...
... The majority of the owners in the current study declared food as the predominant type of reinforcer employed in their dog's training. This aligns with the prevailing trend in previous dog cognition studies, where food was frequently utilized as the primary reward in learning or cognitive tasks González-Martínez et al., 2013;Junttila et al., 2022;Khan et al., 2023;Piotti et al., 2018). However, a critical question still remains unanswered: whether the choice of reward in these experiments might have exerted any influence on dogs' task performance or potentially contributed to the exclusion of certain participants. ...
... These abilities, although debated [15,35], include an innate capacity to understand human pointing gestures with minimal learning time [34]. However, dog breeds vary widely in behavioural profiles, cognitive abilities [36][37][38] and capacity to understand and respond to human visual cues [17,36,37,39]. Some of these differences are reflected in breed function, with 'cooperative' breeds (i.e. ...
... However, recent neuroanatomical work detected differences in internal brain structure associated with specific breed functions, such as hunting, guarding and companionship [42]. Genetic [43] and behavioural work has also linked cognitive disparities among breeds to specific lineages and functions, as evidenced by patterns of breed-specific behaviours including playfulness, sociability and aggression [36,39,44]. Lastly, absolute brain size in dogs has been linked to higher 'executive function' and is suggestive of advanced cognition [38]. ...
... Lastly, absolute brain size in dogs has been linked to higher 'executive function' and is suggestive of advanced cognition [38]. Given the relevance of brain size in comparative studies of cognition [5,6,45] and the recent genetic, behavioural and neuroanatomical advances [39,42,44,46,47], the relationship between a dog's relative brain size and cognition warrants further investigation. ...
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Domestic dogs display a remarkable diversity of functions, morphologies and cognitive abilities. Using data from 1682 dogs representing 172 breeds, we tested for variation in relative endocranial volume (REV), a proxy for brain size and a basic measure of cognitive ability, in relation to breed function, phylogenetic classification, cranial shape, cooperative behaviour and temperament. Function, body size, phylogenetic clade and cranial shape correlate with REV. Toy dogs, functioning mainly as companions, have the largest endocranial volumes relative to their body size. Working dogs, bred to perform complex human-assistance skills and reportedly possessing higher cognitive abilities, have the smallest. Our results thus show that complex skills and cooperative behaviour—a hallmark of social cognition—do not predict larger REV in dogs. However, REV increases with fear and aggression, attention-seeking and separation anxiety and decreases with trainability. Significant correlations between REV and behavioural traits underscore the evolutionary plasticity of mammalian brain size under domestication and artificial selection and provide support for hypotheses linking the modulation of fear and aggression to brain size change under domestication.
... While the artificial breeding in domestic dogs has provided a unique opportunity for researchers to study the heritability of cognitive traits and breed-typical behaviour, 29 , previous research has yielded inconclusive and conflicting findings. A large body of research found substantial differences in behavioural traits across different breeds, suggesting genetic factors as one of the main elements causing these differences 19,29,63 . On the other hand, some studies found that breed has little predictive value for the behaviour of individual dogs 2,35,39 . ...
... As the capacity to learn may be genetically determined, behavioural differences across dog breeds can stem from not only differences in various innate expressions of behaviour (e.g., stalking behaviour) but also in learning capacity itself (e.g., how quickly something is learned or the value of a certain reward). In this regard, previous research findings indicated that cognitive abilities (e.g., problem-solving abilities) and working intelligence (i.e., the general ability to learn tasks) have strong potential to undergo selection and vary across breeds 19,29 , with this potentially attributed to differences in genetically determined physical skill performance, such as running speed, jumping height, and physical strength when it comes to working-related tasks 23 . ...
... Behavioural persistence and social cognition may have gone through diversifying artificial selection in different dog breeds 29,32 . In support of this, we found differences in learning and behavioural performances across distinct dog breed clades. ...
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Learning is crucial for shaping domestic dogs’ behaviour through life experiences, yet not all breeds exhibit the same learning aptitude towards a particular task. The current study’s objective was to identify differences in behaviour and learning performance across and within five breed clades and elucidate the underlying factors contributing into these variations. Dogs (n = 111) from five breed clades (UK Rural, Retrievers, Asian Spitz, European Mastiff, and New World) participated in a virtual learning task with their owners. Owners completed validated questionnaires of Impulsivity and Reward Responsiveness. The learning task comprised of reinforcing an arbitrary behaviour (hand-touch) through multiple sessions of Acquisition (reinforcing the hand-touch), Discrimination (reinforcing the hand-touch on one of two hands) and Reversal Learning (reinforcing the hand-touch on the opposite hand), followed by a single session of Extinction (hand-touch not reinforced). Results showed notable differences across the studied breed clades in certain learning and behavioural components. However, the observed disparities may not be entirely attributed to inherent cognitive differences among the breed clades but rather potentially influenced by contextual factors such as the human-dog communication dynamics associated with breeds’ cooperativity. Furthermore, breed clades differed in the contributing factors predicting individual learning performances, which could highlight the potential effect of breeds’ historical function.
... Domestic dogs exhibit substantial variation in their behavioral tendencies and cognitive abilities. This can be seen, for example, in the diversity of behavioral phenotypes that characterize dog breeds (6)(7)(8)(9). Even within a given breed, individual dogs exhibit extensive variation in their behavior and cognition (9)(10)(11). ...
... In these tasks, the dog could reach the motivator if they performed the correct behavior(s), which was expected to reflect a particular ability. For example, tests that involved detouring around a barrier required the dog to inhibit the impulse of moving directly towards a motivator and instead to first move away from it to reach it, which may reflect their level of inhibitory control or impulsivity (7). In choice tasks, understanding information about where a motivator was hidden would allow them to reach it, for example by following a human's pointing gesture, which may reflect their ability to understand human communicative cues (85). ...
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Background Behavioral testing is widely used to measure individual differences in behavior and cognition among dogs and predict underlying psychological traits. However, the diverse applications, methodological variability, and lack of standardization in canine behavioral testing has posed challenges for researchers and practitioners seeking to use these tests. To address these complexities, this review sought to synthesize and describe behavioral testing methods by creating a framework that uses a “dog-centric” perspective to categorize the test stimuli used to elicit responses from dogs. Methods A scoping review was conducted to identify scientific literature that has reported behavioral testing to assess psychological traits in dogs. Five online databases were systematically searched. Following this, an inductive content analysis was conducted to evaluate and summarize the behavioral testing methods in the literature. Results A total of 392 publications met the selection criteria and were included in the analysis, collectively reporting 2,362 behavioral tests. These tests were individually evaluated and categorized. Our content analysis distinguished 29 subcategories of behavioral testing stimuli that have been used, grouped into three major categories: human-oriented stimuli; environmental stimuli; and motivator-oriented stimuli. Conclusion Despite the methodological heterogeneity observed across behavioral testing methods, our study identified commonalities in many of the stimuli used in test protocols. The resulting framework provides a practical overview of published behavioral tests and their applications, which may assist researchers in selecting and designing appropriate tests for their purposes.
... The modules range from the simplest forms of sensorimotor interference that provide the necessary feedback for differentiating the self from the environment [17] to more complex cognitive features such as episodic-like memory [18], or the formation of theory of mind [19]. Although domesticated species, especially dogs and cats (e.g., [20,21]), have recently been at the forefront of investigation of non-human sociocognitive capacities, apart from a few exceptions (e.g., [22,23]) they are seldom included in experiments about self-representation. ...
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Body awareness allows the individual to negotiate spatial tasks by referencing their own body. Here, we tested whether biologically meaningful factors, such as an alternative solution (detour around an obstacle) and learning from a human demonstrator, would affect dogs’ reliance on their body size in an aperture test. We hypothesized that the dogs would choose the socially reinforced solution over a shortcut, and they would choose the shortcut when the opening was comfortably large. We tested N = 45 adult, mixed-breed dogs, by using a 3 m long, transparent fence. The dogs had three trials with closed doors and then three trials with either a small (but passable) or a large open door. In the demonstration group the experimenter performed a detour before the first three trials, then we opened the large door. The dogs preferred the shortcut, where they relied on body awareness, as they either hesitated or opted for a detour when the small door was open. The dogs who watched the demonstration for longer in trial 3 chose the door less often in trial 4, which indicates social learning. By testing mixed-breed dogs, our results are minimally influenced by functional breed selection and serve as valuable baseline for testing complex socio-cognitive traits in companion dogs.
... Dogs are not merely well-trainable helping assets to humans, but they also seem to have developed a specific willingness to cooperate with us in joint actions, preferring human partners over their conspecifics with whom they rather maintain a competitive/neutral relationship [14]. Dogs (from this point on, we always refer to dogs which have an owner/handler) can show strong variability in their reliance on human-provided cueing and assistance depending on their genetic distance from the common ancestor [15], breed-specific selection [13], or level of training [16]. However, in general, dogs still show a more prevalent dependency on human contribution in difficult-to-solve situations than (socialised) wolves do [12]. ...
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Dogs are strongly dependent on humans, not only for sustenance, but they also form asymmetrical bonds with us where they rely on assistance from the human partner in the case of difficult situations. At the same time, cohabiting dogs form hierarchies, and their rank strongly influences their behaviour in various social interactions. In this study, we investigated whether high- and low-ranking dogs would behave differently in non-social and social contexts where a formerly available reward suddenly becomes inaccessible. We hypothesised that dominant and subordinate dogs would show different levels of human dependence; thus, they will show different levels and different signs of frustration depending on the social nature of the context, where the reward was locked either in a closed cage or withheld in the hand of the experimenter. The results showed that while the holistic rank (‘dominant’ vs. ‘subordinate’) of the dogs did not show a significant association with their reactions to frustrating situations, the rank components (‘agonistic’ and ‘leadership’ ranks) were better predictors of the dogs’ behaviour. In the non-social context, the highly resource-oriented ‘agonistic-dominant’ dogs were more persistent with their attempts of getting to the reward. However, in the social context, the dogs with high ‘leadership scores’ behaved more demandingly with the non-complying experimenter. This study provides a first-time indication that the various aspects of dominance in dogs can affect their adaptive reward-oriented behaviours differently, depending on the potentially available human assistance.
... Although dogs in general have a species-typical predisposition for affiliation with humans 29 , their sensitivity for human behavior differs between breed types 26,30 . Dog breeds bred for close cooperation with humans seem to be more sensitive to human behavior and emotions than more independent breeds. ...
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Behavioral and physiological synchrony facilitate emotional closeness in attachment relationships. The aim of this pseudorandomized cross-over study was to investigate the emotional and physiological link, designated as co-modulation, between dogs and their owners. We measured the heart rate variability (HRV) and physical activity of dogs belonging to co-operative breeds (n = 29) and their owners during resting baselines and positive interaction tasks (Stroking, Training, Sniffing, Playing) and collected survey data on owner temperament and dog–owner relationship. Although overall HRV and activity correlated between dogs and their owners across tasks, task-specific analyses showed that HRV of dogs and owners correlated during free behaving (Pre- and Post-Baseline), whereas the activity of dogs and owners correlated during predefined interaction tasks (Stroking and Playing). Dog overall HRV was the only predictive factor for owner overall HRV, while dog height, ownership duration, owner negative affectivity, and dog–owner interaction scale predicted dog overall HRV. Thus, the characteristics of dog, owner, and the relationship modified the HRV responses in dog–owner dyads. The physiology and behavior of dogs belonging to co-operative breeds and their owners were therefore co-modulated, demonstrating physiological and emotional connection comparable to those found in attachment relationships between humans.
... Currently, the differences between dog breeds have been highlighted in behavioural sciences and there are several approaches to the grouping and comparing of breeds. Some researchers use convenience samples (i.e. to compare the most popular/available breeds at a given location and time [41,42]), which are a favourable method for explorative research, but it makes the formulating of hypotheses about the potential factors behind the found differences difficult. Ancestry-based approaches, aided by the recent advances of molecular genetics and the subsequent construction of evolutionary 'trees' for dog breeds (e.g. ...
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Background While complex dog–human coexistence has been deeply investigated, there is a relative scarcity of similar knowledge regarding dog–dog interactions. Social learning, a fundamental synchronizing mechanism between dogs and humans, was recently found to be influenced by the functional breed selection of dogs: with the cooperative breeds being more effective learners from a human demonstrator than the independent working breeds were. Here, we investigated whether these differences would also be present when dogs had to learn from another dog and how to effectively perform a detour around a transparent V-shaped obstacle. We tested dogs from 28 independent and 19 cooperative breeds in three consecutive trials. In the control groups, all dogs had to detour on their own the obstacle. In the dog demonstration groups, in trial 1, the subjects had to detour on their own, but before the next two trials, a trained dog showed them the solution. Results We found that the performance of the two breed groups was the same in the without demonstration groups. However, after observing the dog demonstrator, the independent dogs learned the task more successfully than the cooperative breeds did. In the case of the independent working breeds, detour latencies significantly dropped along the consecutive trials, and these dogs also showed higher rate of successful detours after observing the demonstrator dog’s action than in the control group. Conclusions This is the first study where the consequences of functional breed selection were confirmed in a scenario that involved conspecific social learning in dogs. The results fit well to the ecologically valid framework of the evolutionary past of dog breed formation, in which cooperative breeds were selected for their interactivity with humans, whereas independent breeds often had to work together with their conspecifics. Graphical Abstract
... Apart from the bewildering array of morphological differences among the various breeds of dogs [3], traditionally they are also characterized by typical behavioral traits that are both mentioned in the official breed standards and utilized in work and sports-related activities [4]. Terms such as "herding dog, " "sighthound" or "sled dog" are telltale signs of the original utilization of the breeds, however, it is commonly assumed that dog breeds would also exhibit different traits of temperament [5] and behavior in everyday contexts such as problem-solving [6] and social interactions with other dogs and humans [7]. ...
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Background The ability to learn from humans via observation was considered to be equally present across properly socialized dogs. We showed recently that cooperative working breeds learned from a human demonstrator more effectively. We hypothesized that functional breed selection could affect sensitivity to human attention-eliciting behavior. Accordingly, we ran the first ever study on dogs that compared the effect of ostensive and neutral verbal communication in a social learning scenario. We used the detour paradigm around a transparent V-shaped fence with either ostensive (addressing the receiver both with words and specific, attention-eliciting prosody) or neutral speech (monotonous reciting of a short poem) demonstration. The other features (gestures, movement) of the demonstration sequence were kept identical between the two conditions. We tested (N = 70) companion dogs from 17 cooperative and 16 independent breeds in three 1-min trials. Subjects had to obtain the reward by detouring around the fence. Results Detour latencies of the cooperative dogs improved after both ostensive and neutral speech demonstrations. The independent dogs did not improve their detour latency in either of the conditions. Remarkably, ostensive verbal utterances elicited longer relative looking time towards the demonstrator, cooperative dogs looked longer at the demonstrator, and longer looking time resulted in more successful detours. Conclusions Our study provides the first indication that functional breed selection had a significant impact on dogs’ sensitivity to ostensive human communication, which, apart from being crucially important for social learning from humans, until now was considered as a uniformly present heritage of domestication in dogs. Graphical Abstract
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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).
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The communicating skills of dogs are well documented and especially their contact-seeking behaviours towards humans. The aim of this study was to use the unsolvable problem paradigm to investigate differences between breed groups in their contact-seeking behaviours towards their owner and a stranger. Twenty-four dogs of ancient breeds, 58 herding dogs, and 17 solitary hunting dogs were included in the study, and their behaviour when presented with an unsolvable problem task (UPT) was recorded for 3 min. All breed groups interacted with the test apparatus the same amount of time, but the herding dogs showed a longer gaze duration towards their owner compared to the other groups and they also preferred to interact with their owner instead of a stranger. Interestingly, the solitary hunting dogs were more in stranger proximity than the other groups, and they also showed a preference to make contact with a stranger instead of their owner. Hence, we found differences in contact-seeking behaviours, reflecting the dog–human relationship, between breed groups that might not only be related to their genetic similarity to wolves, but also due to the specific breeding history of the dogs.
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Animals of different taxa can read and respond to various human communicative signals. Such a mechanism facilitates animals to acquire social information and helps them react in a context-dependent manner. Dogs have garnered extensive attention owing to their socio-cognitive skills and remarkable sensitivity to human social cues. For example, dogs readily respond to different human pointing gestures to locate hidden food rewards. However, a general inclination towards testing highly socialized pet dogs has resulted in a dearth of information on other sub-populations of dogs. Free-ranging dogs are one of the least socialized dog populations yet exhibit point-following behaviour flexibly. As a consequence of frequent negative interspecific interactions, they are typically wary of unfamiliar humans; thus, contextual recognition of human actions is paramount for these dogs to avoid potential conflict. However, the mechanisms influencing their point-following behaviour remain unidentified. We asked to what extent the informative-deceptive nature of cues and positive human interactions influence the interspecific communicative behaviour of these minimally socialized dogs. Using a point-following experiment with a 2 × 2 design, we focused on adult free-ranging dogs’ behavioural adjustments. Dogs were randomly divided into two groups, with only one receiving brief social petting. Further, informative and deceptive cues were given to separate subsets within each group. Our findings suggest that brief social petting strongly affects the likelihood of free-ranging dogs’ point-following tendencies. Dogs who received petting followed the pointing cues regardless of their informative or deceptive nature, whereas dogs who did not receive petting discriminated between informative and deceptive pointing. This study highlights the contribution of positive human interaction and informative-deceptive quality of cues in modulating the behavioural responses of free-ranging dogs in an interspecific communicative context.
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Forming eye contact is important in dog–human communication. In this study we measured what factors affect dogs’ propensity for forming eye contact with an experimenter. We investigated the effect of [1] cephalic index (head shape’s metric, indicator of higher visual acuity at the centre of the visual field), [2] breed function (visual cooperativeness), [3] age and [4] playfulness with strangers in 125 companion dogs. Cephalic index was measured individually and analysed as a continuous variable. Results showed that [1] dogs with a higher cephalic index (shorter head) established eye contact faster. Since cephalic index is highly variable even within a breed, using artificial head shape groups or breed average cephalic index values is not recommended. [2] Breed function also affected dogs’ performance: cooperative breeds and mongrels established eye contact faster than dogs from non-cooperative breeds. [3] Younger dogs formed eye contact faster than older ones. [4] More playful dogs formed eye contact faster. Our results suggest that several factors affect dogs’ interspecific attention, and therefore their visual communication ability.
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Previously, we found that dogs belonging to the herding breed group, selected for human cooperation, synchronise their long-term stress levels with their owners. The aim of the current study was to investigate features that could influence long-term stress levels in ancient dog breeds, genetically closer to wolves, and dogs specifically selected to work independently of their owner. Twenty-four ancient breed dogs and 18 solitary hunting dogs were recruited and hair samples were obtained from both dogs and owners from which hair cortisol concentration (HCC) was analysed. Additionally, the owners completed lifestyle surveys, the Monash Dog Owner Relationship Scale (MDORS) on human–dog relationship, and both dog and owner personality questionnaires (Dog Personality questionnaire and Big Five Inventory survey). The results from the MDORS indicate that the subscale Perceived cost correlated to the dog HCC of tested breed groups: solitary hunting breeds (χ2 = 4.95, P = 0.026, β = 0.055), ancient breeds (χ2 = 2.74, P = 0.098, β = 0.027), and herding dogs included from a previous study (χ2 = 6.82, P = 0.009, β = − 0.061). The HCC of the solitary hunting dogs was also related to the owner personality traits Agreeableness (χ2 = 12.30, P < 0.001, β = − 0.060) and Openness (χ2 = 9.56, P = 0.002, β = 0.048) suggesting a more substantial influence of the owner on the solitary hunting dog’s HCC compared to the ancient breeds. No effect of owner HCC on dog HCC was found in either ancient or in solitary hunting breeds. Hence, the long-term stress synchronisation is likely to be a trait in breeds selected for human cooperation. In conclusion, dog HCC is often related to the owners’ personality, but is primarily influenced by the owner-dog relationship.
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We investigated how dog–owner relationship–with a focus on possible behavioural problems–might associate with the individual variability in dogs’ social learning performance. Dog owners first completed a questionnaire about their relationship with their dogs (N = 98). Then, dogs were tested in a detour test: a control group without demonstration, a group where the owner demonstrated the task and another group where the experimenter demonstrated the task. Finally, the dogs participated in two behaviour tests measuring their tractability and possessiveness. The two principal components from the questionnaire (called “overactive” and “irritable”) did not show significant association with dogs’ detour performance in the control group. “irritable” dogs performed better in the unfamiliar demonstrator group. These more persistent, goal-oriented dogs also looked back less at their owners during the detour. In the individual problem-solving context, the factor “overactive” had a similar effect on looking back at the owner, suggesting that the items of this component primarily are not connected to the dog–human relationship. Our results indicate that dog–human relationship has an integral role in the complex social behaviour of dogs, which warrants for the need of further empirical testing of the associations between social dynamics in dogs and their relationship with humans, including problem behaviours.
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Communication between dogs and humans is a topic of growing interest, and the “unsolvable task” is a common method used to measure human-directed communication. In this task, dogs learn how to solve a problem to obtain a reward. After a fixed number of trials, the reward becomes impossible to access, arguably leading to communicative attempts from the dog. Although useful to observe dogs’ communicative behaviors in a fairly naturalistic situation, the methodology varies among studies regarding apparatus, number of trials, and other factors. The proxies used, for instance, gaze duration or frequency of gaze alternation, also vary, and there are discrepancies and a debate regarding what the task actually measures. Therefore, in this study, we reviewed the usage of the unsolvable task in canids of the genus Canis, searching Web of Science and Scopus for the terms “dog*”, “Canis”, “dingo*”, “wolf” or “wolves” in the title and "unsolvable task" or "impossible task" in the topic. We included thirty-five studies in this review and discussed their different methodologies and proxies, such as different apparatuses, number of solvable trials, and different interpretations of “looking back”, pointing out how they can affect results and hinder comparisons. Lastly, we used current data to propose strategies to homogenize the use of this important paradigm, with an ethogram of possible behaviors and their interpretation and a predefined set of methodological aspects for future research.
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Interspecific communication between dogs and humans enables dogs to occupy significant roles in human society, both in companion and working roles. Dogs excel at using human communicative signals in problem-solving tasks, and solicit human contact when unable to solve a problem. Dogs’ sociocognitive behavior likely results from a selection for attention to humans during domestication, but is highly susceptible to environmental factors. Training for particular tasks appears to enhance dog–human communication, but effects may depend on the nature of the relationship determined by their role. Our aim was to examine two types of social cognition (responsiveness to human gestures, and human-directed communicative behavior in an unsolvable task) in pet dogs (n = 29) and detection dogs (n = 35). The groups did not differ in their ability to follow human signals, but pets were less responsive to signals given by a stranger than by their owner. Pets also exhibited more human-directed gazing in the unsolvable task, showing a bias for gazing at their owner compared with the stranger, whereas detection dogs showed greater persistence in attempting to solve the task compared with pets. Thus, different aspects of dogs’ sociocognitive behavior may differentially vary as a function of selection or training for particular roles.