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Behaviour problems and anxieties in dogs decrease their quality of life and may lead to relinquishment or euthanasia. Considering the large number of pet dogs and the commonness of these problematic behaviours, a better understanding of the epidemiology and related molecular and environmental factors is needed. We have here studied the prevalence, comorbidity, and breed specificity of seven canine anxiety-like traits: noise sensitivity, fearfulness, fear of surfaces and heights, inattention/impulsivity, compulsion, separation related behaviour and aggression with an online behaviour questionnaire answered by dog owners. Our results show that noise sensitivity is the most common anxiety-related trait with a prevalence of 32% in 13,700 Finnish pet dogs. Due to the high prevalence of noise sensitivity and fear, they were the most common comorbidities. However, when comparing the relative risk, the largest risk ratios were seen between hyperactivity/inattention, separation related behaviour and compulsion, and between fear and aggression. Furthermore, dog breeds showed large differences in prevalence of all anxiety-related traits, suggesting a strong genetic contribution. As a result, selective breeding focusing on behaviour may reduce the prevalence of canine anxieties. Anxious animals may suffer from chronic stress and thus, modified breeding policies could improve the welfare of our companion dogs.
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Prevalence, comorbidity, and breed
dierences in canine anxiety in
13,700 Finnish pet dogs
Milla Salonen
1,2,3, Sini Sulkama1,2,3, Salla Mikkola1,2,3, Jenni Puurunen
1,2,3,
Emma Hakanen1,2,3, Katriina Tiira
1,2,3, César Araujo1,2,3 & Hannes Lohi
1,2,3*
Behaviour problems and anxieties in dogs decrease their quality of life and may lead to relinquishment
or euthanasia. Considering the large number of pet dogs and the commonness of these problematic
behaviours, a better understanding of the epidemiology and related molecular and environmental
factors is needed. We have here studied the prevalence, comorbidity, and breed specicity of seven
canine anxiety-like traits: noise sensitivity, fearfulness, fear of surfaces and heights, inattention/
impulsivity, compulsion, separation related behaviour and aggression with an online behaviour
questionnaire answered by dog owners. Our results show that noise sensitivity is the most common
anxiety-related trait with a prevalence of 32% in 13,700 Finnish pet dogs. Due to the high prevalence of
noise sensitivity and fear, they were the most common comorbidities. However, when comparing the
relative risk, the largest risk ratios were seen between hyperactivity/inattention, separation related
behaviour and compulsion, and between fear and aggression. Furthermore, dog breeds showed large
dierences in prevalence of all anxiety-related traits, suggesting a strong genetic contribution. As a
result, selective breeding focusing on behaviour may reduce the prevalence of canine anxieties. Anxious
animals may suer from chronic stress and thus, modied breeding policies could improve the welfare
of our companion dogs.
Problematic behaviours can be a threat to dog welfare. Anxious dogs may be more vulnerable to diseases and
show decreased lifespan1. Satisfaction with the dog’s behaviour may increase attachment to the dog2 and prob-
lematic behaviours, especially aggressiveness, destructiveness, fearfulness and hyperactivity are a common reason
for relinquishment to shelters3,4. Problematic behaviours can even lead to euthanasia at a young age5,6. Behaviour
problems, especially aggressiveness, may be public health concerns7. Some of these behaviour problems have
been suggested to be analogous, or possibly even homologous to human anxiety disorders8, and the study of these
spontaneous behaviour problems arising in a shared environment with people may reveal important biological
factors underlying many psychiatric conditions. For example, canine compulsive disorder resembles human OCD
on both phenotypic and neurochemical level8.
Behaviour problems are common in our companion canines9. e most common reported behaviour prob-
lems include excessive barking, inappropriate elimination, destructiveness, aggression and fearfulness6,1013. e
prevalence of noise sensitivity in previous studies has varied between 20% and 50%1418. Around 20–25% of
dogs show fearfulness of strangers, dogs or situations15,16 and separation anxiety occurs in 14–20% of dogs15,16,19.
Furthermore,comorbidity between noise sensitivity and separation anxiety has also been observed15,20.
Behaviour has a major genetic component2127 and many traits are both phenotypically and genetically cor-
related22. For example, relatives of compulsive dogs are oen also aected28. Some genomic areas and loci are
associated with problematic behaviour, including compulsion29, fear and noise sensitivity30. Furthermore, prob-
lematic behaviour may be inuenced by many environmental factors, including, for example, maternal care,
owner experience, training and exercise3133. Behaviours are complex traits aected by several genes with small
eects, multiple environmental factors varying in eect, and intricate interactions between them34.
Many epidemiological cross-sectional studies of behaviour are hindered by a small sample size and breed
coverage or limited number of traits for a comprehensive overview. For example, our earlier anxiety survey
included only three traits and reached ~3,300 participants16. erefore, we developed here a more comprehensive
1Department of Medical and Clinical Genetics, University of Helsinki, Helsinki, Finland. 2Department of Veterinary
Biosciences, University of Helsinki, Helsinki, Finland. 3Folkhälsan Research Center, Helsinki, Finland. *email: hannes.
lohi@helsinki.
OPEN
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owner-completed online questionnaire to collect a large data set of companion dogs in the study of canine anxi-
ety. We expanded the study from three to seven anxiety-related traits and acquired four times as many records to
assess the prevalence of dierent anxiety-like behaviour problems in a large home living population of companion
canines, comorbidity between the anxiety-related traits, and trait-specic behavioural variation between breeds,
which could indicate genetic inheritance.
Results
Demography. We examined the epidemiology of seven anxiety-like traits in dogs: noise sensitivity, fear, fear
of surfaces, inattention/impulsivity, compulsive behaviour, aggression and separation related behaviour with a
comprehensive owner-answered online questionnaire. We collected 13,715 responses in 264 dog breeds. In total,
51.5% of the dogs were female and the age of the dogs varied between 10 weeks and 17 years 10 months (mean 4.7
years). We received more than 200 responses from mixed breed dogs and from 14 breeds: Bernese Mountain Dog,
Border Collie, Finnish Lapponian Dog, German ShepherdDog, Labrador Retriever, Lagotto Romagnolo,
Lapponian Herder, Miniature Schnauzer, Rough Collie, Shetland Sheepdog, Smooth Collie, So-Coated Wheaten
Terrier (labelled Wheaten Terrier), Spanish Water Dog and Staordshire Bull Terrier(labelled Sta. Bull Terrier).
ese breeds and mixed breed dogs made up 35% of all dogs in the data. e sex ratio did not signicantly dier
between these breeds (χ2 = 4.17, DF = 14, P = 0.994), but the mean age diered (Kruskal-Wallis χ2 = 77.53,
DF = 14, P < 0.001): Bernese Mountain Dogs (
x
= 3.8 years, SD = 2.7) were, on average, younger and Miniature
Schnauzers (
x
= 5.3 years, SD = 3.4) and Wheaten Terriers (
x
= 5.7 years, SD = 3.7) older than other breeds.
Prevalence. In total, 72.5% of dogs had some kind of highly problematic behaviour. Noise sensitivity was the
most common anxiety trait with 32% of dogs being highly fearful of at least one noise (Fig.1a, Supplementary
TableS1). Fear was the second most common trait with a prevalence of 29%. Separation related behaviour and
aggression were the most uncommon traits with prevalences of 5% and 14%, respectively.
e prevalence of the subtraits varied (Fig.1b, Supplementary TableS1). In noise sensitivity, fear of reworks
was the most common subtrait with a prevalence of 26%. When comparing subtraits of fear, fear of other dogs
Noise sensitivit
y
Fear
Fear of surfaces and heights
Inattention
Compulsive behaviour
Hyperactivity/impulsivity
Aggression
Separation related behaviour
0
10
20
30
40
Behavioural trait
%of dogs
Fear of fireworks
Fear of thunder
Fear of gunshot
Fear of dogs
Fear of strangers
Fear of novel situations
Fear of surfaces and heights
Inattention
Hyperactivity/impulsivity
Self-biting
Surface licking
Excessive drinking
Tail chasing
Pacing
Fly snapping/light chasing
Staring
Aggression toward family members
Aggression toward strangers
Destroy/urinate alone
Vocalize/salivate/pant alone
0
10
20
30
Behavioural trait
%of dogs
Noise sensitivity
a
b
Fear
Fear of surfaces andh
eights
Inattention/impulsivity
Compulsive behavior
Aggression
Separation anxiety
Figure 1. Prevalence of the traits (a) and subtraits (b) in a sample of 13715 dogs in 264 breeds.
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was the most common. Aggression toward human family members was slightly more common than aggression
toward strangers. From compulsion subtraits, self-biting was the most commonly reported.
Comorbidity. Many dogs exhibited comorbidities between dierent anxiety-related traits (Fig.2a). Most
common comorbidity was fear, especially in aggressive and hyperactive/impulsive dogs, and the second most
common was noise sensitivity, especially in fearful dogs. However, a somewhat dierent pattern emerged when
comparing the relative risks (Fig.2b, Supplementary TableS2). Dogs displaying separation related behaviour
were 4.1 times more oen hyperactive/impulsive and 3.4 times more oen inattentive than dogs not displaying
separation related behaviour. Similar comorbidities were seen between compulsion and hyperactivity/inattention,
and compulsion and separation related behaviour. Finally, aggressive dogs were 3.2 times more oen fearful, and
dogs showing separation related behaviour were 2.8 times more likely fearful.
Comorbidity between subtraits was also common but showed high variability (Supplementary TableS3). Of
the highly noise sensitive dogs, 53% displayed noise sensitivity toward more than one target. Similarly, 38% of
the highly fearful dogs were highly fearful toward more than one subtrait. However, only 9% of aggressive dogs
showed aggression toward both family members and strangers.
Age and sex dierences. Male dogs were more oen aggressive and hyperactive/impulsive, but female
dogs were more oen fearful (Fig.3b–d, Supplementary Fig.S1, Supplementary TableS4). Separation related
behaviour was slightly more common in male dogs. e prevalence of noise sensitivity increased with age, espe-
cially fear of thunder (Fig.3a, Supplementary Fig.S1, Supplementary TableS5). Similarly, fear of surfaces and
1.3
1.3
1.0
1.2
1.4
1.3
1.5
2.3
1.9
2.0
1.6
2.0
2.5
1.7
2.3
1.9
2.0
1.9
1.7
3.2
1.4
1.3
1.7
1.3
1.6
1.4
1.9
1.7
3.2
2.4
2.5
1.8
1.8
1.8
1.5
4.1
3.4
2.8
1.7
1.9
2.8
1.4
2.5
2.5
2.2
1.8
2.0
1.4
3.3
3.1
1.6
2.0
2.0
1.4
Noise sensitivity
Fear
Aggression
Fear of surfaces and heights
Compulsive behaviour
Separation related behaviour
Inattention
Hyperactivity/impulsivity
Hyperactivity/impulsivity
Inattention
Separation related behaviour
Compulsive behaviour
Fear of surfaces and heights
Aggression
Fear
Noise sensitivity
Comorbidityriskratio
1
2
3
4
17.3
22.1
5.7
18.2
29.0
16.9
35.7
23.7
27.3
9.2
21.5
33.2
25.2
40.3
23.7
27.2
8.7
24.4
29.2
51.0
38.5
17.8
25.9
6.6
20.9
17.6
40.4
39.9
27.9
31.3
10.2
30.7
21.6
38.6
36.8
32.2
36.6
29.9
28.3
22.6
48.3
34.1
10.5
26.2
31.9
20.3
41.0
37.5
11.7
29.8
28.0
22.5
45.4
37.4
Noise sensitivity
Fear
Aggression
Fear of surfaces and heights
Compulsive behaviour
Separation related behaviour
Inattention
Hyperactivity/impulsivity
Hyperactivity/impulsivity
Inattention
Separation related behaviour
Compulsive behaviour
Fear of surfaces and heights
Aggression
Fear
Noise sensitivity
Pairs comorbidity/diagnosis
10
20
30
40
50
a
b
Figure 2. Comorbidity heat maps. Proportion of dogs with pairs of diagnoses using the disorder listed in the
column header as the denominator (a) and relative risk (b). (a) Adapted from Goldstein-Piekarski et al.72.
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heights increased with age, whereas hyperactivity/impulsivity and tail chasing decreased. Other traits did not
show clear linear changes.
Breed specicity. Large breed dierences were observed in all behavioural traits (Fig.4, Supplementary
Fig.S2, Supplementary TableS6). For example, 10.6% of Miniature Schnauzers were aggressive toward strangers,
whereas only 0.4% of Labrador Retrievers showed aggression (Fig.4). Similarly, 9.5% of Staordshire Bull Terriers
were reported to display tail chasing, but none of the Lagotto Romagnolo dogs chased their tails.
When focusing on individual breeds, these dierences in prevalences resulted in breed-specic patterns
(Fig.5). Border Collies displayed a very high prevalence of compulsive staring and y snapping, but only moder-
ate levels of other traits (Fig.5). In contrast, Miniature Schnauzers displayed high levels of aggression and social
fear (fear of strangers and other dogs), but few stereotypies. Lagotto Romagnolos displayed high levels of noise
sensitivity, social fear, and aggression. Staordshire Bull Terriers showed high levels of compulsive behaviour,
hyperactivity, and inattention. For more breeds, see Supplementary Fig.S3.
Discussion
We have here developed a comprehensive behavioural survey and applied a citizen science approach to collect a
large canine behavioural dataset of over 260 breeds of dogs. Our data replicates results from some earlier studies
and the study also shows a number of novel insights, including prevalences, comorbidities and breed dierences
of traits not described before. ese results improve the overall understanding of problematic behaviour.
Based on the results here and in previous studies, noise sensitivity stands out as the most common canine
anxiety with a prevalence of 32% in this study. Earlier, the prevalence has varied between 20% and 50%9,12,1418.
Based on our study and previous studies14,17,18, the most common noise sensitivity is the fear of reworks. Fear
was the second most common canine anxiety, with a prevalence of 29%. Specically, 17% of dogs showed fear of
other dogs, 15% fear of strangers and 11% fear of novel situations. Prevalence of total fearfulness12,16 and preva-
lence of fear subtraits9,15,16,35 were quite similar in previous studies as well. Fear of surfaces and heights appears to
be highly prevalent in our study population, as 23.5% of dog owners reported that their dogs were highly fearful
of dierent surfaces and heights.
Based on our results, every h dog displays high levels of inattention and 15% high levels of hyperactiv-
ity/impulsivity. Excessive activity has been reported in 12% to 34% of dogs9,12. Compulsive behaviour patterns
were observed in 16% of the dogs, agreeing with a previous study9. Based on our results and previous studies9,28,
<2 years
2-4 years
4-6 years
6-8 years
8-10 years
>10 years
0
10
20
30
40
Age
Male
Female
<2 years
2-4 years
4-6 years
6-8 years
8-10 years
>10 years
0
5
10
15
20
25
Hyperactivity/impulsivity
Male
Female
<2 years
2-4 years
4-6 years
6-8 years
8-10 years
>10 years
0
5
10
15
Age
%of dogs
Male
Female
<2 years
2-4 years
4-6 years
6-8 years
8-10 years
>10 years
0
5
10
15
20
%of dogs
Fear of strangers
Male
Female
a
b
c
d
Figure 3. Prevalences of fear of thunder (a), aggression towardfamily members (b), hyperactivity/impulsivity
(c) and fear of strangers (d) for both sexes and six age groups.
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self-mutilation is the most common compulsive behaviour. Self-mutilation may be a compulsion, but it may also
be caused by allergies, ectoparasites or other skin problems, possibly explaining the high prevalence of the subtrait.
e prevalence of aggression was 14%, with both aggression toward human family members and toward
strangers occurring in 6% of dogs. In our previous study, 16% of dogs had at least once displayed aggressive
behaviours toward family members and 45% toward strangers16. In previous studies, the prevalence of aggression
and its subtraits has varied between 2% and 30%9,12,15. Studies focusing on referrals to veterinary/behaviour clinics
oen report aggression as the most common behaviour problem10,11,13, possibly because owners nd aggressive-
ness more problematic than, for example, fear of reworks. Separation related behaviours were only displayed by
6% of the dogs. Previously, the prevalence of separation anxiety has been 2–3 times higher than in this present
study9,15,16,19, possibly because we only included dogs with high frequencies of separation related behaviour. Taken
together, our results are surprisingly similar compared to previous studies, even though the populations studied
and the criteria for anxiety-related traits dier in every study.
Lagotto Romagnol
o
Mixed Breed
Wheaten Terrier
Rough Collie
Finnish Lapponian Dog
Staff. Bull Terrier
Border Collie
Miniature Schnauzer
Smooth Collie
Shetland Sheepdog
Spanish Water Dog
Lapponian Herder
German Shepherd Dog
Bernese Mountain Dog
Labrador Retriever
0
10
20
30
40 Fear of thunder
% of dogs
Mixed Breed
German Shepherd Do
g
Spanish Water Dog
Staff. Bull Terrier
Border Collie
Smooth Collie
Bernese Mountain Dog
Lapponian Herder
Lagotto Romagnolo
Shetland Sheepdog
Finnish Lapponian Dog
Labrador Retriever
Wheaten Terrier
Miniature Schnauzer
Rough Collie
0
5
10
15
20
25 Hyperactivity/impulsivity
% of dogs
Staff. Bull Terrier
Finnish Lapponian Do
g
Mixed Breed
Lapponian Herder
German Shepherd Dog
Labrador Retriever
Shetland Sheepdog
Border Collie
Rough Collie
Wheaten Terrier
Spanish Water Dog
Miniature Schnauzer
Bernese Mountain Do
g
Smooth Collie
Lagotto Romagnolo
0
2
4
6
8
10
% of dogs
Tail chasing
Spanish Water Dog
Mixed Breed
Shetland Sheepdog
Miniature Schnauzer
Bernese Mountain Dog
Lagotto Romagnolo
German Shepherd Dog
Smooth Collie
Rough Collie
Lapponian Herder
Labrador Retriever
Border Collie
Finnish Lapponian Dog
Wheaten Terrier
Staff. Bull Terrier
0
10
20
30 Fear of strangers
% of dogs
Mixed Breed
Finnish Lapponian Dog
Lapponian Herder
Miniature Schnauzer
Bernese Mountain Dog
Staff. Bull Terrier
Wheaten Terrier
German Shepherd Dog
Smooth Collie
Shetland Sheepdog
Rough Collie
Lagotto Romagnolo
Labrador Retriever
Border Collie
Spanish Water Dog
0
10
20
30 Inattention
% of dogs
Border Collie
German Shepherd Do
g
Mixed Breed
Shetland Sheepdog
Rough Collie
Labrador Retriever
Spanish Water Dog
Finnish Lapponian Do
g
Lapponian Herder
Wheaten Terrier
Staff. Bull Terrier
Bernese Mountain Do
g
Miniature Schnauzer
Lagotto Romagnolo
Smooth Collie
0
1
2
3
4
5Fly snapping/light chasing
% of dogs
Rough Collie
Mixed Breed
Miniature Schnauzer
Labrador Retriever
Shetland Sheepdog
Wheaten Terrier
Staff. Bull Terrier
Spanish Water Dog
Lapponian Herder
Smooth Collie
Finnish Lapponian Dog
Bernese Mountain Dog
German Shepherd Dog
Lagotto Romagnolo
Border Collie
0
10
20
30
40
50 Fear of surfaces and heights
% of dogs
Miniature Schnauzer
Mixed Breed
German Shepherd Dog
Spanish Water Dog
Border Collie
Lagotto Romagnolo
Wheaten Terrier
Lapponian Herder
Rough Collie
Finnish Lapponian Dog
Bernese Mountain Dog
Shetland Sheepdog
Staff. Bull Terrier
Smooth Collie
Labrador Retriever
0
5
10
15 Aggression toward strangers
% of dogs
Wheaten Terrier
Miniature Schnauzer
Mixed Breed
Lagotto Romagnolo
Spanish Water Dog
Lapponian Herder
German Shepherd Dog
Shetland Sheepdog
Border Collie
Bernese Mountain Dog
Finnish Lapponian Do
g
Rough Collie
Labrador Retriever
Staff. Bull Terrier
Smooth Collie
0
2
4
6Vocalize/salivate/pant alone
% of dogs
a
b
c
d
g
Figure 4. Breed dierences in fear of thunder (a), fear of strangers (b), fear of surfacesand heights (c), hyperactivity/
impulsivity (d), inattention (e), aggression toward strangers (f), tail chasing (g), y snapping/light chasing (h) and
vocalization/salivation/panting alone (i). For other breed-wise dierences, see Supplementary Fig.S2.
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Male and female dogs displayed dierences in the prevalence of behaviour problems. Male dogs had a higher
prevalence of aggressiveness, separation related behaviour, inattention and hyperactivity/impulsivity. In contrast,
female dogs had a higher prevalence of fearfulness. Noise sensitivity, fear of surfaces and compulsive behaviour
occurred independent of sex. Previous studies have shown similar sex dierences in aggression9,13,3638, fearful-
ness13,32,3941, separation related behaviour36 and compulsion33. Reported sex dierences in noise sensitivity are
less clear (more common in females17,32, more common in males14, no dierence between sexes18).
We detected dierences between age groups in the prevalence of most behaviour problems. Younger dogs
had a higher prevalence of destroy/urinate when alone, inattention, hyperactivity/impulsivity, tail chasing and
self-biting. Older dogs had a higher prevalence of aggression, noise sensitivity and fear of surfaces. Fearfulness
was most common in dogs aged 4–8 years. Furthermore, vocalize/salivate/pant when alone and other compul-
sive behaviours (beside the aforementioned tail chasing and self-biting) occurred independent of age. Previous
studies are in agreement at least in noise sensitivity14,17,18,32, inattention42,43, hyperactivity/impulsivity12,4244 and
aggression12,37. Tail chasing, destroying and urinating indoors are typical behaviours for puppies, and this likely
explains the age dierences in these subtraits.
We observed large behavioural dierences between breeds. Noise sensitivity was the most common in Lagotto
Romagnolo, Wheaten Terrier and mixed breed dogs. Previous studies have also ranked these breeds high in
noise sensitivity14,1618. Among some other breeds, Miniature Schnauzers and Staordshire Bull Terriers were, in
contrast, noise sensitive less oen, as also observed in previous studies10,17. Fear was most common in Spanish
Water Dogs, Shetland Sheepdogs and mixed breeds. In contrast, Labrador Retrievers were seldomly fearful. ese
results are in agreement with previous studies, ranking mixed breed dogs high in fearfulness9,13,35,40 and Labrador
Retrievers and Staordshire Bull Terriers low in fearfulness35,41,45,46. Fear of surfaces and heights was the most
oen observed in Rough Collie and mixed breed dogs.
Self-biting
Surface licking
Excessive drinking
Pacing
Tail chasing
Staring
Fly snapping/
light chasing
Vocalize/salivate/
pant alone
Destroy/urinate
alone
Hyperactivity/
impulsivity Inattention Aggression toward family members
Aggression toward strangers
Fear of surfaces
and heights
Fear of novel situations
Fear of strangers
Fear of dogs
Fear of gunshot
Fear of thunder
Fear of fireworks
Border Collie Miniature Schnauzer
Lagotto Romagnolo Staffordshire Bull Terrier
Self-biting
Surface licking
Excessive drinking
Pacing
Tail chasing
Staring
Fly snapping/
light chasing
Vocalize/salivate/
pant alone
Destroy/urinate
alone
Hyperactivity/
impulsivity Inattention Aggression toward family members
Aggression toward strangers
Fear of surfaces
and heights
Fe
ar of novel situations
Fear of strangers
Fear of dogs
Fear of gunshot
Fear of thunder
Fear of fireworks
Self-biting
Surface licking
Excessive drinking
Pacing
Tail chasing
Staring
Fly snapping/
light chasing
Vocalize/salivate/
pant alone
Destroy/urinate
alone
Hyperactivity/
impulsivity
Inattention Aggression toward family members
Aggression toward strangers
Fear of surfaces
and heights
Fear of novel situations
Fear of strangers
Fear of dogs
Fear of gunshot
Fear of thunder
Fear of fireworksSelf-biting
Surface licking
Excessive drinking
Pacing
Tail chasing
Staring
Fly snapping/
light chasing
Vocalize/salivate/
pant alone
Destroy/urinate
alone
Hyperactivity/
impulsivity
Inattention Aggression toward family members
Aggression toward strangers
Fear of surfaces
and heights
Fear of novel situations
Fear of strangers
Fear of dogs
Fear of gunshot
Fear of thunder
Fear of fireworks
ab
cd
Figure 5. Radar chart representation of the behaviour of dog breeds: Border Collie (a), Miniature Schnauzer
(b), Lagotto Romagnolo (c) and Staordshire Bull Terrier (d). Colors represent the larger traits. Clockwise
from top: blue – noise sensitivity, lime green – fear, violet – fear of surfaces and heights, orange – aggression,
pine green – hyperactivity/inattention, purple – separation related behaviour, yellow – compulsive behaviour.
Radar charts for other breeds in Supplementary Fig.S3. e minimum in all traits is the breed-wise minimum
prevalence and the maximum in all traits is the breed-wise maximum prevalence. For the minimum and
maximum prevalences, see Supplementary TableS7.
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Inattention was most oen reported in mixed breed dogs, Finnish Lapponian Dogs and Lapponian Herders,
and rarely reported in Spanish Water Dogs and Border Collies. Although Finnish Lapponian Dogs and Lapponian
Herders have not been studied before, our results agree with previous results43. As the dog breeds showing high
prevalence of inattention are breeds that are oen regarded as “hard to train, owners may rate dogs not easily
motivated by food or petting as inattentive. Hyperactivity and impulsivity were the most common in mixed breed
dogs, German Shepherds, Spanish Water Dogs and Staordshire Bull Terriers, and the least common in Rough
Collies and Miniature Schnauzers. Similar breed dierences were observed in a previous study44. Furthermore,
herding dogs (including, for example, Border Collie and German Shepherd) and terriers (including, for example,
Staordshire Bull Terriers) have ranked high in extraversion and, in contrast, toy dogs low in extraversion47. In
our study, Labrador Retrievers and Rough Collies had a low prevalence of hyperactivity/impulsivity, but other-
wise our results match previous studies. However, it seems that classication into traditional and genetic breed
groups poorly reect behavioural dierences48, possibly explaining the dierences in these results. Compulsive
behaviour was most oen reported by owners of German Shepherds, mixed breed dogs and Staordshire Bull
Terriers. However, the breed dierences varied highly between dierent compulsions. For example, Staordshire
Bull Terriers had a high prevalence of tail chasing, with nearly 10% of them chasing their tails. In contrast, light
chasing and staring were the most oen observed in Border Collies. Interestingly, Border Collies were bred to
herd livestock by staring at them intensely, and even though this method of herding can be perfected by training,
the behaviour itself seems to be innate49. Pacing and excessive drinking were oen performed by mixed breed
dogs and German Shepherds. In a previous study, German shepherds had high odds of being presented to a
behaviour clinic for obsessive behaviour10.
Mixed breed dogs and Miniature Schnauzers had the highest prevalence of aggression, whereas Labrador
Retrievers had the lowest prevalence of aggression. Aggression toward strangers was most prevalent in Miniature
Schnauzers, mixed breed dogs, German Shepherd Dogs and Spanish Water Dogs, and least prevalent in Labrador
Retrievers. Aggression toward human family members was most common in Miniature Schnauzers and Lagotto
Romagnolos. Our results agree with previous studies both in total aggression9,13 and in the subtraits, aggression
toward strangers3638,45 and aggression towards family members16,37. Separation related behaviour was most com-
mon in mixed breed dogs and Wheaten Terriers. Specically, mixed breed dogs were likely to destroy, urinate or
defecate when le alone, whereas Wheaten Terriers were likely to vocalize, salivate or pant. Based on our results
and a previous study36, mixed breed dogs may be more prone to show separation related behaviour. It is possible
that the high prevalence of separation distress and other anxieties in the mixed breed dogs is caused by a poor
early life environment and adverse experiences in life, as many mixed breed dogs in our data are likely rescues.
Within-trait comorbidity was common in noise sensitivity and fear: 53% of dogs that were fearful of one noise
were fearful of several noises, and 38% of fearful dogs were fearful of more than one target. is result was also
discovered in our previous study16. Based on previous studies, noise sensitivity is oen generalised and displayed
toward several dierent noises9,14,17,18. We discovered that dogs were seldomly aggressive toward both family
members and strangers, as reported before in some dog breeds50. In contrast, one previous study did report
a signicant comorbidity between stranger-directed and owner-directed aggression13. However, it seems that
aggression toward strangers and family members are genetically distinct traits51.
We discovered that noise sensitivity and fear were the most common comorbidities, likely due to their high
prevalence in our study population. However, when comparing the risk ratios in comorbid traits, the largest risk
ratios were seen between separation related behaviour, hyperactivity/impulsivity, inattention and compulsive
behaviour, and between fear and aggression. Fearful dogs were 3.2 times more oen aggressive than non-fearful
dogs, a relationship found in previous studies as well9,13,16,38. is indicates that aggression is commonly motivated
by fear. e connection between impulsivity, compulsive behaviour and separation related behaviour is an interest-
ing nding that demands further research. One previous study discovered that excitable dogs had 9.8 times higher
odds of separation distress52 and another study discovered a connection between compulsive behaviour and hyper-
activity9. Intriguingly, impulsivity and compulsion are related constructs, as both are proposed to be caused by a
failure of response control and mediated by basal ganglia53. We observed many trait connections detected in pre-
vious studies as well, including comorbidity between fear and noise sensitivity14,16,17, between fear and separation
related behaviour16, between separation related behaviour and aggression13,16 and between fear and compulsive
behaviour9,33. However, previous studies have detected a comorbidity between separation anxiety and noise sen-
sitivity9,13,15,17,19,20. We indeed discovered that separation related behaviour was 1.4 times more prevalent in noise
sensitive dogs. However, the opposite was not true, as dogs showing separation related behaviour were not fearful
of noises more oen than dogs not showing separation related behaviour. Furthermore, we discovered a positive
connection between compulsive behaviour and aggression, contrasting with the results of our previous study33.
is study has limitations. Although the fear section of the questionnaire was validated and the test-retest
reliability of the fear and noise sensitivity sections was good54 and that the results we have obtained from the data
collected with it32,33,55 replicate many previous results, the psychometric properties of the rest of the question-
naire have not been formally evaluated. Future studies should aim to assess the reliability and validity of these
additional components. Secondly, the categorisation into low, moderate, and high categories was mostly based on
the frequency of signs and not the severity, except in aggression, separation anxiety, and impulsivity/inattention.
us, in the high groups, the severity of the symptoms can be variable. irdly, our sample is a self-selected con-
venience sample, and may not be representative of the overall Finnish dog population. Although the most com-
mon breeds in our sample are also common in Finland56, the representativeness of our sample is still unknown.
We are currently working on a separate study to understand the participant proles and details of the sample
demographics.
Our ndings on breed dierences indicate that canine anxieties likely have a genetic basis. In previous stud-
ies, many behavioural traits have been indeed shown to have small to moderate heritabilities22,26,57 and recently
we mapped two loci for generalized fear and noise sensitivity30. erefore, it could be possible to decrease the
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prevalence of canine anxieties by selecting non-anxious animals for breeding. Our results also show that these
canine anxieties are phenotypically correlated. Some of these traits, like many behaviour traits21,22, may also be
genetically correlated, and therefore selection for one trait may inuence other traits as well. Interestingly, a
genomic region associated with noise sensitivity in German Shepherd Dogs30 contains the oxytocin receptor gene
(OXTR). e gene is associated with social behaviour58 (but see a contrasting study with a smaller sample size59)
and most likely has been under strong selection during domestication. is could explain the high prevalence
of noise sensitivity in many study populations12,1418 and could also indicate that breeding eorts to reduce the
prevalence of noise sensitivity may prove dicult.
Based on our results, canine anxieties and behaviour problems are common across breeds. ere are around
77 million dogs in the United States60 and 85 million in Europe61, and therefore these behaviour problems can
aect millions of animals. As anxiety can impair welfare1 and problematic behaviour may be an indication of
poor welfare62, eorts should be made to decrease the prevalence of these canine anxieties. Breeding policies may
help to improve dog welfare, as could changes in the living environment14,19,32,33,37. Our ongoing eorts aim to
identify environmental and genetic risk factors behind these canine anxiety-related traits using the large survey
data collected here.
Methods
Questionnaire. A comprehensive online questionnaire was designed to collect extensive information about
behaviour and background informationof dogs (Supplementary information). e questionnaire consisted of
a background section and sections focusing on seven canine anxiety-related traits: noise sensitivity, fear, fear of
surfacesand heights, impulsivity/inattention, compulsive behaviour, aggression and separation related behaviour.
ese traits consisted of several subtraits (Fig.1b). e fear part of this questionnaire was previously shown to
have good validity and both the fear and noise sensitivity sections had good test-retest reliability54. A question-
naire can be a good method for collecting data, since the reliability of questionnaires has been good in behav-
ioural science and the answers are strongly linked to the behaviour of the animals54,6368. Furthermore, as the
owners have a long history with the dogs, it ispossible to study traits that would be dicult to study using other
methods (for example, behaviour test batteries), including compulsive behaviour69.
Subjects. Owners were mainly recruited from a social media (Facebook) channel. Furthermore, dog breed
organisations advertised the questionnaire on their own websites and Facebook pages. In total, we received ques-
tionnaire responses of 13,715 dogs in 264 breeds. To ensure reliable prevalence estimates in breed comparisons,
we calculated a minimum sample size per breed with the Epitools sample size calculator70. For this calculation,
we used the average prevalence of behaviour traits (19%), the average number of registrations per dog breed
within the last 10 years (1500), a desired precision of 0.05 and a condence level of 0.95. is calculation resulted
in a minimum sample size of 200 individuals per breed. We obtained more than 200 answers from 14 breeds
and mixed breed dogs (N = 418): Finnish Lapponian Dog (N = 538), Labrador Retriever (N = 465), German
Shepherd Dog (N = 461), Shetland Sheepdog (N = 411), So-coated Wheaten Terrier (N = 351), Lapponian
Herder (N = 294), Border Collie (N = 268), Miniature Schnauzer (N = 255), Spanish Water Dog (N = 251), Rough
Collie (N = 248), Lagotto Romagnolo (N = 248), Bernese Mountain Dog (N = 209), Smooth Collie (N = 203) and
Staordshire Bull Terrier (N = 200).
Informed consent was obtained from all participants. Participants agreed that all questionnaire answers could
be used for research. We emphasized that all data will be handled strictly condentially, and that individual dogs
and owners cannot be recognized from the published results.
Subject categorisation. Based on the questionnaire scores, dogs were categorised into low, moderate, and
high groups for each subtrait (Fig.1b, Supplementary TableS1) depending on the frequency of the behaviour.
e subtraits were, in turn, combined to form trait groups. Low trait groups consisted of dogs that fell into low
groups in all subtraits, and high trait groups consisted of dogs that had a high score in at least one subtrait. e
categorisation for dierent subtraits is explained below.
Noise sensitivity. Noise sensitivity consisted of three subtraits: fear of thunder, reworks, and gunshot. e
respondents were rst asked whether their dog showed fear toward these targets, and secondly, how oen their
dog shows fear, from rarely (0–20% of the time) to always (100% of the time). Dogs belonging to low group did
not show fear toward these targets and the owners did not report a frequency for noise sensitivity. Dogs belonging
to the high group showed fear at least oen (40–60% of the time).
Fear. e fear section of the questionnaire consisted of three fear subtraits: fear of strangers, other dogs, and
novel situations. Dogs that showed fear at least oen (40–60% of the time) constituted the high group and dogs
that never showed fear constituted the low group in all subtraits. Furthermore, low fear dogs never barked or
growled at strangers or other dogs. Behaviour tests have previously been conducted to validate fear of strangers
and fear of novel situations phenotypes54.
Fear of surfaces and heights. In this section, owners were asked whether their dog had diculties walking on
dierent surfaces, including on a metal grid, on shiny oors, or moving from one surface to another. Moreover,
owners were asked whether their dog had diculties in high places: climbing stairs where you can or cannot see
between steps, walking next to glass railings, climbing metal stairs, and walking over narrow bridges. In all ques-
tions, these answers were scored between 0 (never) and 4 (always). Dogs were categorised as high fear of surfaces
if at least one of the answers was 3 (oen) or 4, and low group consisted of dogs that never had diculties in any
of these tasks.
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Impulsivity/inattention. Dog owners were asked to rate their dog’s behaviour on a scale of 1 (never) to 4 (very
oen) in 13 questions, developed by Vas and colleagues42 and designed to measure inattention, hyperactivity,
and impulsivity. A principal component analysis with a promax rotation was run to divide the questions into two
components, inattention and hyperactivity/impulsivity (Supplementary TableS8), and component scores were
calculated for each dog. One question (Item 11) was removed from the analysis, as it loaded equally on both com-
ponents. e cut-o between low and moderate category was set at the rst dog having a rating of 3 (oen) in the
item that had the highest loading (Item 2 in inattention, Item 5 in hyperactivity/impulsivity). e cut-o between
the moderate and high category was set at the rst dog having a rating of 4 (very oen) in the highest loading item
and the same rating on one additional item.
Compulsive behaviour. Compulsive behaviour part of the questionnaire measured the occurrence of tail chas-
ing, y snapping/light chasing, surface licking, pacing, staring, excessive drinking, and self-biting. Tail chasing,
y snapping/light chasing, surface licking, pacing, and staring were scored on a scale of 0 (I’ve never noticed
this behaviour) to 6 (several times per day). e dogs scoring from 4 (every other day-weekly) to 6 were placed
in the high group, and dogs scoring 0 or 1 (a few times during the dog’s lifetime) were placed in the low group.
Furthermore, owners were asked to estimate the time dog spent drinking or near the water bowl from less than
5 minutes to 1 hour or more. Dogs that spent less than 5 minutes near the water bowlformed the low group in
excessive drinking, and dogs spending more than 15 minutes near it formed the high group. Finally, the preva-
lence of self-biting was rated from 0 (never) to 3 (several hours per day). Dogs scoring 2 (almost every day) or 3
constituted the high group whereas dogs scoring 0 constituted the low group.
Aggression. e aggression trait in the questionnaire consisted of two subtraits: aggression toward strangers and
toward family members. e respondents were asked to score the likelihood of their dog growling and trying to
snap/bite from 1 (never) to 5 (always or almost always) when a stranger tries to pet the dog in its home or outside
and when the owner handles the dog or tries to take a resource (food, bone, or toy) from the dog. If the dog tried
to snap or bite at least sometimes (3) or it growled at least oen (4), it was categorised as a high aggression dog in
each subtrait. e dogs that never showed aggression in any of these situations constituted the low group.
Separation related behaviour. Separation related behaviour was divided into two parts: destroy/urinate alone
and vocalize/salivate/pant alone. Dog owners were asked whether their dog exhibited separation anxiety and how
oen did the dog perform the aforementioned behaviours from 0 (never) to 4 (very oen). High group dogs were
reported to exhibit separation anxiety and performed at least one of these behaviours oen (3) or very oen. With
the subtrait vocalize/salivate/pant, high group dogs were reported to salivate or pant at least oen (3–4), or barked
at least oen (3–4) and salivated or panted at least rarely (1–4). Low group dogs did not show separation anxiety
and never performed these behaviours.
Statistical analyses. To estimate the prevalence of high anxiety dogs, the number of dogs in high group was
divided by the sum of dogs in all trait groups, hence obtaining the percentage of dogs in the high group for each
subtrait and trait in the population.
e same prevalence calculation was performed individually for dierent breeds (sample size > 200) and for
dened age groups and both sexes. e dogs were divided into six age groups: less than 2 years, 2–4 years, 4–6
years, 6–8 years, 8–10 years, and more than 10 years. Chi-squared tests were run in R71 for breed, sex, and age
group dierences in all traits and subtraits to reveal the signicant associations.
Comorbidities were calculated between all pairs of behaviour problems. In the pairs comorbidity/diagnosis
adapted from Goldstein-Piekarsi et al.72, the number of dogs belonging to the high group in both of the traits was
divided by the number of all dogs in the high group in the denominator trait, hence obtaining the percentage of
dogs in each denominator trait that also had the comorbid trait. e relative risk was calculated by rstly calcu-
lating the same percentage and secondly calculating the percentage of dogs that belonged to the low group in the
denominator trait but in the high group in the other trait. irdly, the rst percentage was divided by the second.
Two-proportions z-tests were run in R71 for all risk ratios to see whether any two traits have statistically signicant
comorbidity.
All P-values were corrected for false discovery rate (FDR) to decrease the probability of type I error. e sig-
nicance cut-o P-value was set at P < 0.05.
Data availability
e anonymised data is available as a supplementary datasheet.
Received: 17 November 2019; Accepted: 29 January 2020;
Published: xx xx xxxx
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Acknowledgements
We thank all the dog owners who participated in the study. is study was partially funded by the Academy of
Finland (308887), the ERCStG (260997), ERA-NET NEURON and the Jane and Aatos Erkko Foundation. H.L. is
a member of HiLIFE. M.S., S.M., E.H. & H.L. are members of the Helsinki One Health.
Author contributions
M.S., S.S. & H.L. designed the study. S.S., K.T., & H.L. developed the questionnaire. S.S. & H.L. piloted and edited
the questionnaire and collected the data. M.S., S.S., & S.M. handled and processed the data with help from J.P. &
E.H. using the scripts planned by M.S., S.S., H.L. and developed by C.A. M.S. performed the statistical analyses.
Manuscript was planned by M.S., S.S. and H.L., draed by M.S., and revised and accepted by all.
Competing interests
e authors declare no competing interests.
Additional information
Supplementary information is available for this paper at https://doi.org/10.1038/s41598-020-59837-z.
Correspondence and requests for materials should be addressed to H.L.
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Fear is an emotion needed to survive, but when prolonged and frequent, causes suffering in both humans and animals. The most common forms of canine anxiety are as follows: general fearfulness, noise sensitivity, and separation anxiety are responsible for a large proportion of behavioral problems. Information on the prevalence and comorbidity of different anxieties is necessary for breeding, veterinary behavior, and also for behavioral genetic research, where accurate information of the phenotype is essential. We used a validated owner-completed questionnaire to collect information on dogs' fearfulness (toward unfamiliar people, dogs, in new situations), noise sensitivity, separation anxiety, as well as aggressive behavior. We received 3284 answers from 192 breeds. The prevalence estimate for noise sensitivity was 39.2 %, 26.2% for general fearfulness, and 17.2% for separation anxiety. The owner reported the median onset age for noise sensitivity to be 2 years and varied between 8 weeks and 10 years (N = 407). High comorbidity was observed between different anxieties: fearful dogs had a significantly higher noise sensitivity (P < 0.001) and separation anxiety (P < 0.001) compared with nonfearful dogs. Fearful dogs were also more aggressive compared with nonfearful dogs (P < 0.001). Prevalence estimates of fearfulness, noise sensitivity, and separation anxiety are in agreement with earlier studies. Previous studies have suggested early onset of noise sensitivity during the first year of life; however, we found a later onset with large variation in the onset age. High comorbidity between anxieties suggests a genetic overlap. Fearful personality may predispose to specific anxieties such as noise sensitivity or separation anxiety.
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Variation across dog breeds presents a unique opportunity to investigate the evolution and biological basis of complex behavioural traits. We integrated behavioural data from more than 14 000 dogs from 101 breeds with breed-averaged genotypic data (n = 5697 dogs) from over 100 000 loci in the dog genome. We found high levels of among-breed heritability for 14 behaviour-al traits (the proportion of trait variance attributable to genetic similarity among breeds). We next identified 131 single nucleotide polymorphisms associated with breed differences in behaviour, which were found in genes that are highly expressed in the brain and enriched for neurobiological functions and developmental processes, suggesting that they may be functionally associated with behavioural differences. Our results shed light on the herit-ability and genetic architecture of complex behavioural traits and identify dogs as a powerful model in which to address these questions.
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Aggression in dogs is a safety concern both for humans and animals, and can lead to decreased animal welfare in affected dogs due to potential abuse, neglect, relinquishment or euthanasia. We examined risk factors associated with stranger-directed aggression in dogs using the previously validated, Canine Behavioural Assessment and Research Questionnaire (C-BARQ). Results are based on participant reports of dog behaviour. Data were analyzed using mixed logistic regression, with participant ID and country as random effects. Dogs (n = 14,310) were more likely to demonstrate stranger-directed aggression if the participant rated them as mildly or severely fearful of strangers, or mildly, but not severely, fearful in non-social situations, when compared to dogs with no fear. There was an interaction between sex and neuter status, with neutered males being more likely to be aggressive than any other group. Furthermore, adult dogs were more likely to be aggressive compared to adolescents or seniors, and dogs were less likely to be aggressive if acquired as an adult when compared to being acquired as a puppy or adolescent. The random effects for country and participant were significant (p. <. 0.001) with ICCs of 0.01 (CI: 0.00-0.08) and 0.40 (CI: 0.35-0.46), respectively, indicating that there was some correlation in behaviour among dogs within the same country and owned by the same person. The moderate effect of participant suggests that household effects need to be examined further. When looking only at dogs categorized as aggressive towards strangers (n = 11,240), dogs were significantly more likely to be categorized as having severe aggression if they were male, and if the owner rated them as mildly or severely fearful of strangers, or mildly, but not severely, fearful in non-social situations, when compared to dogs with no fear. Breed group and where the dog was acquired also had an association with severe aggression. The random effects for country and participant were significant (p. <. 0.001) with ICCs of 0.06 (CI: 0.02-0.15) and 0.34 (CI: 0.22-0.48), respectively, indicating once again that there was some correlation in behaviour among dogs within the same country and owned by the same person. These results suggest that variables related to the environment, owner experience and the dog's level of fearfulness are associated with aggressive behaviour towards strangers in dogs. Therefore, it might be possible to identify dogs at risk of developing stranger-directed aggression and implement plans to prevent behavioural issues from developing.
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Variation in the oxytocin receptor (OXTR) gene has been linked to dog social behavior in some studies, while others have failed to replicate this relationship (in dogs and other species). Ratings of observed behavior are often used to assess personality, however owner-based behavior/personality assessments have been shown to be comparably consistent. The relationship between OXTR gene variation and behavior as measured by owner-based personality questionnaires has yet to be explored in dogs. The aims of our current study were three-fold: (1) to investigate the relationship between two OXTR SNPs (rs8679684 and 19131AG) and three owner-based behavior/personality measures in dogs (the MCPQ-R, a history of aggression questionnaire, and an “eagerness to please” scale), (2) to explore the relationships of personality assessments and OXTR polymorphisms with genetic breed clusters, purebred versus mixed breed status, and how the dog was acquired (shelter versus other means), and (3) to examine the relationships among the three owner-based assessments. We found no relationship between any of the three personality measures and genotype at either SNP. This may be due to the relationships between OXTR gene SNPs and behaviors being breed-specific, in which case our genetically varied sample would not detect them. In line with kennel club breed descriptions and prior research, herding breeds scored higher in “eagerness to please” than scent hound breeds, and scent hounds’ Training Focus scores were lower than those of both herding and working breeds. Purebred and mixed breed dogs did not score differently in any of the three personality assessments, and dogs adopted from shelters were rated lower in Extraversion and “eagerness to please” than those acquired by other means, suggesting that early shelter experience may affect adult behavior. Dogs with no history of aggression towards household members, unfamiliar children, unfamiliar men/women, and unfamiliar dogs scored higher in Amicability, and “eagerness to please” scores were positively correlated with Training Focus and Amicability. These personality measures appear to evaluate different facets of the same personality concepts. Future research on OXTR gene variation in dogs should be mindful of the possibly limited implications of behavioral assessments that are context-specific.