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Dogs aren’t jealous – they are just asking for accurate information

Authors:
Overall, Karen L. (2018) Dogs aren’t jealous – they are just asking for accurate
information.
Animal Sentience
22(17)
DOI: 10.51291/2377-7478.1351
This article has appeared in the journal
Animal Sentience
,
a peer-reviewed journal on animal cognition and feeling. It
has been made open access, free for all, by WellBeing
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Animal Sentience 2018.141: Overall on Cook et al. on Dog Jealousy
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Dogs aren’t jealous – they are just asking for accurate information
Commentary on Cook et al. on Dog Jealousy
Karen L. Overall
Biology Department, University of Pennsylvania
Abstract: Awake fMRI offers us a unique opportunity to view and understand how dogs see the
world and use the information in it. Given the limitations of behavioral assays and the small sample
sizes inherent in these studies, labeling of patterns of canine behaviors using pop psychology terms
may actually interfere with our understanding of canine brains and obscure for us a more
parsimonious but exciting interpretation of canine behavior. We should use this window into how
dogs think wisely.
Karen Overall, senior researcher,
Biology Department, University of
Pennsylvania, and Editor-in-Chief of
Journal of Veterinary Behavior: Clinical
Applications and Research, is author of
multiple text books and hundreds of
scientific articles. Her research focuses
on behavioral genetics of anxiety and
other disorders in dogs, on dogs as
natural animal models, and on problem-
solving behavior and other
performance assays in pet and working
dogs.
The target article by Cook et al. (2018) adds to the growing compendium of information about
regional brain activity for a group of pet dogs taught to cooperate with and participate in imaging
using awake fMRI. This body of work (Berns et al., 2012, 2013, 2017; Cook et al., 2014, 2016 a-c)
has suggested that dogs perform similarly to humans with respect to stimulus-dependent activity
in the amygdala and caudate nuclei. These findings are not unexpected given the convergent
patterns of evolution of candidate genes associated with neurotransmitter function shared by
dogs and humans (Saetre et al., 2004; Li et al., 2013).
Because of the detailed methodology used to validate this type of imaging, and the
conditions under which measurements are made, there is no doubt that there is variation in
regional signals and scan parameters within this small population of trained dogs (here, 13 dogs
are studied). The interpretation of the attendant variation in behavior is, unfortunately, not
assessed with a similarly rigorous methodology. This is no fault of the authors but is instead a
reflection of the state of the field of assessment in veterinary behavior and behavioral medicine
and it warrants discussion.
Phenotyping behavior. The behavioral assay used here is the C-BARQ, a questionnaire that asks
owners to rate their dogs’ behaviors in a variety of situations. There are two concerns here. The
first is that the C-BARQ is invariably represented as a validated tool. However, the study that is
Animal Sentience 2018.141: Overall on Cook et al. on Dog Jealousy
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always cited as having validated it (Hsu and Serpell, 2003) used two sets of clinical populations for
which published, detailed, semi-quantitative, questionnaires (Overall, 1997) were used in
conjunction with clinical behavioral evaluations. As such, the version of C-BARQ published as part
of this study represented a dramatic change in content from its earlier version (Serpell and Hsu,
2001). So what may have occurred during this process is that the questionnaires used to assess
the patients were actually validated for a clinical population. This matters because clinical
populations are not opportunistic samples, as is the case for most studies published using C-BARQ.
Clinical populations have greatly diminished population variation compared with the
range of dogs without behavioral concerns and are comprised of dogs whose owners already
believe that their dog needs specialist help. As such, the owners are not evaluating the dogs
wholly by themselves but are providing semi-quantitative, objective information, targeted by the
clinicians, which correlates with known patterns of behavioral associations within behavioral
diagnoses (Overall, 2001, 2005). As part of the process of making such diagnoses, owners’ answers
to questionnaires are themselves assessed for accuracy to ensure that what the clinician intended
by a word or phrase is understood and evaluated within that content. Such etymological care is
necessary because words used to talk about behaviors are all in the popular domain and
unprotected by scientific terminology. It is for this reason that behavioral medicine specialists use
such questionnaires in the context of an in-person behavioral evaluation of the patient, targeted
video and, sometimes, provocative tests. Only in this multi-tiered information context can
diagnostic criteria be consistently and accurately applied. As part of the process, questionnaire
information may change or be considered not relevant, as objective and quantitative information
from the behavioral evaluation emerges. When more subjective, owner-completed
questionnaires are used, there are no such standards or tests for accuracy. Simply, the population
for which the C-BARQ claims validation differs and is held to a different standard with respect to
validity.
The second concern is the accuracy of owner-based descriptions and assessments. The
extent to which owners are willing to report behavioral traits that they think are perceived as
unfavourable depends on the confidentiality of the survey (Segurson et al., 2005). Subjects may
not all be dealt with equitably. There are also data showing that owners may recognize or
understand only the easiest to detect signs of any fear- or anxiety-related condition and so under-
estimate the presence of the condition if their dogs show different signs (Mariti et al., 2012). The
extent to which owners recognize anything other than overtly happy behaviour depends on their
professional training with dog behaviour, their personal experience with dogs, and which body
part(s) they are observing (Wan et al., 2012). Even after being given training to recognize
behavioural signs of concern or aggression in dogs, when quizzed using specific risk photo
scenarios, owners consistently under-estimate risk to children, and err in the interpretation of
canine behaviors (Arhant et al., 2016). When asked to view the scenarios as reflections of their
own dogs, owners further discount behavioural signs.
Even in studies where clients have received information defining specific behavioural
patterns, some behavioral signs used by researchers are simply less apparent than others if
constant monitoring of the dog does not occur. If dogs are not exposed to certain provocative
stimuli, or not observed to react in any given context, false negatives can be reported, a sequela
that can be minimized by behavioural testing and rigorous, objective quantitative or semi-
quantitative questionnaires (Overall et al., 2006; Tiira and Lohi, 2014; Overall et al., 2016; Bellamy
Animal Sentience 2018.141: Overall on Cook et al. on Dog Jealousy
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et al., 2018). Furthermore, dogs may change with age, exposure, and level and type of provocative
stimulus, all of which may have affected how the owner evaluated the dog the day any survey
was completed.
I go into this depth with respect to concerns about the use of a questionnaire as a solitary
behavioural measure because behaviour is more complex than this, and the assessment of scores
as a measure of dog-dog or dog-human aggression may be optimistic. Barking and growling
are not isomorphic with aggression. “Aggression” can be normal or pathological, and
determining this involves more than vocal signals. Barking, growling, et cetera can be viewed as
provocative signals designed to gain information about potential interactions. As such they may
tell us more about awareness, vigilance and tendencies to react, regardless of outcome. This
sounds like a minor distinction, but labelling a dog aggressive or an interaction as aggression
is not a trivial matter.
How do dogs inform others that they are aroused? Given the distribution of scores reported by
Cook et al. (2018), the dogs with the greatest extremes in average amygdala activation had the
lowest behavioral scores. Only 6/13 dogs had a score of greater than 1, but less than 2, suggesting
at most a mild behavioural response. Finally, in the best-case scenario, the C-BARQ scores
accounted for only 22% one fifth of the variance in a small population.
So, are we actually studying an aggressive response, or any kind of aggression, let alone
dog-dog aggression or human resource guarding? Here again, the pop culture labelling
appears to have gotten ahead of the science something the authors acknowledge in their
discussion.
The data for increased differential arousal are actually quite compelling. Consider that
dogs are not all the same with respect to their arousal levels (e.g., the level to which they will
react) or reactivity profile (e.g., the number of situations over which they react and number of
behaviors they use as part of their reaction) (Overall, 2013). Some dogs will need more
information than others to form a response in an uncertain circumstance. Rather than postulate
covert arousal, the authors may wish to consider the role played by signals involved in putative
aggressive responses. The point of signalling behaviour is to make clear something that was
previously unknown or hidden (Smith, 1981). When the signal is not clear or understood, the
receiver (here, the dogs in the scanner) may send reciprocal signals back to the original sender
(the fake dog) and may further act to seek information and clarity (Smith, 1981). Under such
circumstances, dogs who are more easily aroused or more reactive may exhibit provocative
behaviors or experience provocative arousal in the absence of more overt behaviors, especially if
the arousal is mild and they have been taught to behave in a specific manner within the given
context (e.g., the scanner). This profile is consistent with the Cook et al. results.
As the authors clearly note, none of these dogs was overtly aggressive, none of them had
given the owners cause to complain of any aggression, and all had been readily trained as part of
a group to cooperate with all aspects of the imaging process for these awake fMRIs, a scenario
that would probably have provoked dogs who had a propensity to react with aggression for any
reason. Furthermore and this is important the authors found significant habituation of the
amygdala response across experimental sessions but only in the aggressive dogs: the ones who
had amygdala activation in the first place. Notably, this activation was maximal in the first run but
effectively nonexistent in the second and third runs. Whether the decrease in amygdala response
Animal Sentience 2018.141: Overall on Cook et al. on Dog Jealousy
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is due to habituation or desensitization is difficult to determine, but a decreased physiological
response following repeated exposure is consistent with the classical definition of habituation
(page 9, Cook et al., 2018). This pattern of behavior is again consistent with provoking, obtaining
and using information about whether one should react. Clearly, these dogs gained sufficient data
across exposures to provide them with the information that further arousal was not warranted.
Fabulous fakes are still fake. Although the fake dog had been flavored with canine odorants,
dogs quickly read and integrate many more subtle signals over a variety of signaling modalities to
gain information. Dogs can distinguish between odorants from various body regions and can
discriminate scent pictures (Kalmus, 1955); they respond with incredibly different amplitudes and
directions of tail wags depending on the social stimulus (Quaranta et al., 2007), and they are able
to signal attention to human facial signaling with one set of muscles controlling their brow region
(Kaminski et al., 2017). Exposure would be sufficient to tell test subjects that these signaling
domains lacked congruence and that this dog was not a dog.
The problem with using models to represent a real, but potentially risky experience is at
the core of evaluations to test whether dogs will react adversely to food being taken, or to a child.
Data collected both in shelter environments (where food is taken with a fake plastic hand on a
stick) and home environments (where humans interact with dogs and food) have shown that the
test with the fake hand produces large numbers of false positives for a “problem” that people
who adopt the dogs do not perceive as a concern and that generally does not require further
intervention in a home environment (Mohan-Gibbons, 2012; Marder et al., 2013).
A similar concern has been raised for the use of dolls to test aggression toward children.
In a test situation, a doll or a fake dog elicited more social behaviors than did an ambiguous object,
but it lacked sufficient validity to use in a predictive assessment for any dogs tested when the
groups (control, aggressive to dogs, aggressive to children) were compared (Barnard et al., 2012).
Significant associations were found for barks, growls and snaps, signals used to elicit further
information about whether frank aggression is warranted.
These studies tell us is that dogs recognize models as “other”, i.e., not a real hand, not a
real dog, not a real child. The Cook et al. data support this contention.
The importance of labels. Behavior is dynamic. It’s the visceral representation of the integration
of all the body’s organ system responses to the immediate environment, within the boundary
conditions set by underlying genetic constraints and previous learning at the molecular level. This
means that behavior is difficult to define and measure in ways that are unambiguous. When such
concerns are raised, we should be mindful of attribution and should understand that how we label
something may affect how we think about it in which case, we should be careful, indeed. The
biggest risk is that we will think by that the application of a label, that we understand something
and we do not (Feymann, 1973).
We know that dogs recognize resource inequity, and having something or not having
something (Range et al., 2009), so need we label this as anything else? Jealousy is a concept in
human psychology that varies in how it is defined, and it is difficult to test. By using psychometric
testing to evaluate dependency, self-esteem, insecurity and trust, Rydell and Bringle (2007) found
that jealousy grouped into reactive and suspicious morphs, where insecurity and suspicious
jealousy co-occurred, but dependency did not. The pattern was reversed for reactive jealousy.
Animal Sentience 2018.141: Overall on Cook et al. on Dog Jealousy
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Reactive jealousy was more often associated with external events, whereas suspicious jealousy
was under endogenous control and associated with anxiety. The definitional and evaluative
criteria used in this and related studies are not met here, so a borrowed characterization of
jealousy is premature.
Dogs may not be “guarding” their humans as a “social resource” through any use of
aggressive signals when their humans interact with another dog; they may be asking for
clarification of the behavioral patterns, using signals that occur in agonistic events but that need
not by themselves entail true hostility or aggression. It’s far more likely that such events are an
essential part of defining, establishing, solidifying and maintaining social relationships for a
species that have a co-evolutionary history with humans but that differ in the repertoire of verbal
and non-verbal signals.
The role of opposable thumbs (which dogs lack) is essential to understanding differences
in how dogs signal. When dogs ask questions of and gain information from other dogs in active
interactions, tactile exchanges are frequent and complex, often involving mouthing (Normal et
al., 2015; Cordoni et al., 2016), an activity that humans both misunderstand and actively
discourage in interactions with dogs. Detailed evaluation of dogs approaching known vs. unknown
dogs in a controlled test situation reveals a discrepancy in behavioral patterns (Mariti et al., 2017).
The incidence and frequency of various behaviors differ depending on whether the interaction
was close or at a distance, whether the recipient of the signals was known or unknown, and what
signal preceded the response. The most common response to any aggressive signal from an
unknown dog was to freeze again, a finding consistent with the Cook et al. data.
There are other explanations for this study’s outcomes that do not involve evoking
aggression or guarding behavior. The handler, who was instructed to look at the dog in the
scanner only when feeding that dog, either put food into a hidden pouch attached to the fake
dog’s mouth or placed it into a bucket. Kaminski et al. (2017) have demonstrated that in such
circumstances the absence of attentiveness from humans dogs view food as a non-social
but arousing signal. Yet the handler knew whether they were putting food in the bucket or into
the fake dog’s pouch and so could have exhibited subtle intention cues that animals with
experience with humans learn and use to shape responses (Proops et al., 2010, 2013), again
providing information that may have caused differential attention across the study dogs. Both
outcomes pertain to the Cook et al. data.
We should also consider that pet dogs witness food of all types being put in various
containers as part of daily life. At a minimum, this activity signals to them that the food is
unavailable to them. Here, the fake dog could not exhibit any behavior, including appetitive
behavior. Such a contextually discordant situation would logically provoke information-seeking
behavior in dogs who were monitoring the reactions of others more; this may be what has been
described here. One would then hypothesize that with exposure and no further interaction on
the part of the fake dog, these dogs would attenuate their response once they received no input
from either the fake dog or the handler because there was no value of salience to them in future
interaction. This is exactly what happened here in an outcome consistent with the Cook et al.
results.
So there are potentially many things going on here; there is need to invoke jealous
behavior and human resource guarding.
Animal Sentience 2018.141: Overall on Cook et al. on Dog Jealousy
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Scary outcomes: Will dogs die? My final concern is about things that scare me. The possibility
that this study will be used to justify assertions that dogs exhibit jealous behavior and human
resource guardingis worrisome.
As someone who does research as well as sees patients as a specialist in veterinary
behavioural medicine, I see the damage that such labels can do and how virulent simplified
concepts from papers with sexy titles can be for my patients. The dog-owning public still has not
recovered from the popular myth that they must dominate their dogs, despite a large number
of publications showing that normal dogs don’t spend their days challenging people or other dogs
for status. There are compelling data showing that the misapplication of such concepts has
resulted in abusive and damaging treatment to pet dogs (reviewed in Ziv, 2017), and that the risk
of euthanasia is higher for dogs for whom such techniques were recommended by “behaviourists”
who were not veterinary specialists (Siracusa et al., 2017).
As Cook et al. note, there is always a concern about anthropomorphism. However, the
concern quickly rises to the level of anthropocentrism when dealing with sexy labels especially
if they fit with extant bias. When this happens, everything that the dog is, is lost in translation.
Anthropomorphism may demean the inherent nature of self for other species, but it can
make humans more compassionate. Anthropocentrism prohibits us from realizing the extent to
which another species lives within its own context. As result, we are far more interested in how
dogs and other species may resemble us than we are in understanding how they perceive and act
on various behaviors and emotions while they live in a world where they must accommodate
human mistakes in translation.
The cost of such mistakes is high. My fear is that when the popular press grabs hold of the
concepts of jealousy and resource guarding of humans, without any understanding of
whether such concepts are ones that dogs would choose and use themselves, owners will view
canine comments as a signs of jealousy and will punish normal dogs; or they will blithely dismiss
pathological dogs as merely jealous or resource guarding their human, and so fail to seek
timely, humane treatment that will make their and their dog’s lives better and safer.
These fears are not unfounded. “Resource guarding” has become the training
community’s default catchphrase for many behaviors involving dog-dog and dog-human
interactions, stunningly demonstrating how by using a label, we obscure what we could know. It
now takes a 10-to-15-minute disquisition during consultations to disentangle this phrase from
what the dog is actually doing that concerns the clients. This is not progress.
Dogs tell us what they value. Aggression and the signals that go with it can be a normal behavior
in dogs. Some dogs do not like some dogs. Some dogs don’t like or trust some people. These can
be normal behaviors about which dogs should be allowed to signal honestly. In the absence of
labels that obscure our ability to understand what such behaviors mean to the dog, we can learn
their value from the dog’s perspective.
And it is here that the excitement in the Cook et al. study is found. What is fascinating in
this study and should not be lost in any of my complaints about labels, process, and sexy taglines,
is what this differential response tells us about how dogs get and use information. Dogs ask
questions and work for accurate information.
The types of imaging studies pursued by Berns and colleagues, especially when coupled
with the imaging studies of behavioral pathologies (Peremans et al., 2003, 2005, 2006; Vermeire
Animal Sentience 2018.141: Overall on Cook et al. on Dog Jealousy
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et al., 2011, 2012) and interventional studies that show how dogs with specific suites of behaviors
respond to psychopharmacological interventions (Overall, 1994, 2001; King et al., 2000; Simpson
et al., 2007; Korpivaara et al., 2017; Taylor et al., 2017), can be our window into dogs in
translation, as translated by native speakers. For this to happen, we need to let the dogs speak
without anthropomorphism or anthropocentrism and we need to listen without labels.
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L H, Dobbeleir A, Peremans K. Serotonin 2A receptor, serotonin transporter and dopamine
transporter alterations in dogs with compulsive behaviour as a promising model for human
obsessive-compulsive disorder. Psychiatry Research: Neuroimaging, 2012;201:78-87.
Wan M, Bolger N, Champagne FA. Human perception of fear in dogs varies according to
experience with dogs. PLoS One, 2012;7(12):e51775.
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ISC 2018 Summer School
in Cognitive Sciences
June 26 - July 6, 2018
Montreal (Canada)
The Other Minds
Problem: Animal
Sentience and Cognition
Overview. Since Descartes, philosophers know there is no way to know for sure what or whether others feel (not
even if they tell you). Science, however, is not about certainty but about probability and evidence. The 7.5 billion individual
members of the human species can tell us what they are feeling. But there are 9 million other species on the planet (20
quintillion individuals), from elephants to jellyfish, with which humans share biological and cognitive ancestry, but not one
other species can speak: Which of them can feel and what do they feel? Their human spokespersons the comparative
psychologists, ethologists, evolutionists, and cognitive neurobiologists who are the world’s leading experts in “mind-
reading" other species will provide a sweeping panorama of what it feels like to be an elephant, ape, whale, cow, pig,
dog, chicken, bat, fish, lizard, lobster, snail: This growing body of facts about nonhuman sentience has profound
implications not only for our understanding of human cognition, but for our treatment of other sentient species.
Gregory Berns: Decoding the Dog's Mind with Awake Neuroimaging
Gordon Burghardt: Probing the Umwelt of Reptiles
Jon Sakata: Audience Effects on Communication Signals
PANEL 1: Reptiles, Birds and Mammals
WORKSHOP 1: Kristin Andrews: The "Other" Problems: Mind,
Behavior, and Agency
Sarah Brosnan: How Do Primates Feel About Their Social Partners?
Alexander Ophir: The Cognitive Ecology of Monogamy
Michael Hendricks: Integrating Action and Perception in a Small
Nervous System
PANEL 2: Primates, Voles and Worms
WORKSHOP 2: Jonathan Birch: Animal Sentience and the
Precautionary Principle
Malcolm MacIver: How Sentience Changed After Fish Invaded Land
385 Million Years Ago
Sarah Woolley: Neural Mechanisms of Preference in Female
Songbird
Simon Reader: Animal Social Learning: Implications for
Understanding Others
PANEL 3: Sea to Land to Air
WORKSHOP 3: Steven M. Wise: Nonhuman Personhood
Tomoko Ohyama: Action Selection in a Small Brain (Drosophila
Maggot)
Mike Ryan: "Crazy Love": Nonlinearity and Irrationality in Mate Choice
Louis Lefebvre: Animal Innovation: From Ecology to
Neurotransmitters
PANEL 4: Maggots, Frogs and Birds: Flexibility Evolving
SPECIAL EVENT: Mario Cyr: Polar Bears
Colin Chapman: Why Do We Want to Think People Are Different?
Vladimir Pradosudov: Chickadee Spatial Cognition
Jonathan Balcombe: The Sentient World of Fishes
PANEL 5: Similarities and Differences
WORKSHOP 5 (part 1): Gary Comstock: A Cow's Concept of Her
Future
WORKSHOP 5 (part 2): Jean-Jacques Kona-Boun: Physical and
Mental Risks to Cattle and Horses in Rodeos
Joshua Plotnik: Thoughtful Trunks: Application of Elephant Cognition
for Elephant Conservation
Lori Marino: Who Are Dolphins?
Larry Young: The Neurobiology of Social Bonding, Empathy and
Social Loss in Monogamous Voles
Panel 6: Mammals All, Great and Small
WORKSHOP 6: Lori Marino: The Inconvenient Truth About Thinking
Chickens
Andrew Adamatzky: Slime Mould: Cognition Through Computation
Frantisek Baluska & Stefano Mancuso: What a Plant Knows and
Perceives
Arthur Reber: A Novel Theory of the Origin of Mind: Conversations
With a Caterpillar and a Bacterium
PANEL 7: Microbes, Molds and Plants
WORKSHOP 7: Suzanne Held & Michael Mendl: Pig Cognition and
Why It Matters
James Simmons: What Is It Like To Be A Bat?
Debbie Kelly: Spatial Cognition in Food-Storing
Steve Phelps: Social Cognition Across Species
PANEL 8: Social Space
WORKSHOP 8: To be announced
Lars Chittka: The Mind of the Bee
Reuven Dukas: Insect Emotions: Mechanisms and Evolutionary
Biology
Adam Shriver: Do Human Lesion Studies Tell Us the Cortex is
Required for Pain Experiences?
PANEL 9: The Invertebrate Mind
WORKSHOP 9: Delcianna Winders: Nonhuman Animals in Sport
and Entertainment
Carel ten Cate: Avian Capacity for Categorization and Abstraction
Jennifer Mather: Do Squid Have a Sense of Self?
Steve Chang: Neurobiology of Monkeys Thinking About Other
Monkeys
PANEL 10: Others in Mind
WORKSHOP 10: The Legal Status of Sentient Nonhuman Species
... If an owner's anthropomorphic relationship perception is related to social expectations of the cat, this could influence the owner's behavior towards the cat and its living environment. Anthropomorphism has been related to both negative [38][39][40] and positive [41][42][43][44] outcomes for animals. Anthropomorphism can influence feline welfare in a negative way when unrealistic expectations result in disappointment and misinterpretation of behavior results in punishment or inadequate care [45]. ...
Article
Full-text available
Describing the relationship with one’s cat in human terms might reflect an underlying anthropomorphic view of the relationship which might be associated with an owner’s behavior towards their cat and the cat’s living environment. Owners self-categorized the relationship with their cat as either a ‘member of the family’, ‘as a child’, ‘best friend’, or ‘a pet animal’. The extent to which owner- and cat-related factors influence these four relationship descriptions are examined in survey data of approximately 1800 cat owners. Differences in outdoor access, care during absence of the owner, and access to the bedroom are examined between the four relationship perceptions. The owner’s age and household composition, ideas about their cat’s equality, support, and dependency, and whether their cat is a pedigree were significantly associated with relationship description and explained 46% of the variance. Owners who perceive their cat as a child or best friend see their cat as loyal, empathetic, equal to family, and dependent on them for love and care. Their cats are less often left in the care of others, are allowed more often in the bedroom and have less often (unrestricted) outdoor access. Moreover, cats perceived as children are more likely to live in a multi-cat household. Our results provide insight in the factors that are related to different (anthropomorphic) perceptions of the human–cat relationship and how perceptions relate to the living environment of cats.
... In this regard, Cook et al.'s research may not be making all the requisite distinctions. Like other commentators (Bräuer & Amici, 2018;Denson, 2018;Jiang, Huttunen, & Platt, 2018;Overall, 2018;Prato-Previde & Valsecchi, 2018;Vonk, 2018), we wonder how the subject in this study perceived the fake dogespecially in light of recent evidence that dogs do not perceive fake dogs as real social rivals (Prato-Previde et al., 2018b). Perhaps equally concerning is the fact that the relationship between the caregiver and the fake dog is stagnant -never varying in the extent to which it poses a threat to the existing bond between the subject dog and its caregiver. ...
Article
Full-text available
Whereas the feelings of other beings are private and may always remain so, emotions are simultaneously manifested in behavior, physiology, and other observables. Nonetheless, uncertainty about whether emotions can be studied adequately across species has promoted skepticism about their very presence in other parts of the animal kingdom. Studying social emotions like jealousy in the context of the social relationships in which they arise, as has been done in the case of animal empathy, may help dispel this skepticism. Empathy in other species came to be accepted partly because of the behavioral similarities between its expression in nonhuman animals and humans, and partly because of the neurological parallels. Non-invasive brain imaging results like those reported in the target article can thus help integrate human and animal emotions within an evolutionary framework-but the social context underlies precise definitions of the phenomenon.
Article
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It is gratifying and significant that so many scientists from diverse fields are arguing in-depth regarding a particularly complex set of social emotions in a non-human animal. Emotions play a fundamental role in decision making and information processing. Neuroimaging is important in understanding the cognitive and emotional worlds of non-human animals and can help measure covert emotions lacking clear behavioral correlates. Various experimental approaches could clarify the relative importance of attachment and aggression in jealousy and whether the phenomenon we measured is more akin to human envy or jealousy. Reverse inference from amygdala activation is probably justified because behavior is "degenerate": there are fewer behavioral programs than brain states that give rise to them. Individual differences are also important.
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Domestic dogs are highly social and have been shown to be sensitive not only to the actions of humans and other dogs but to the interactions between them. We used the C-BARQ scale to estimate dogs' aggressiveness, and we used noninvasive brain imaging (fMRI) to measure activity in their amygdala (an area involved in aggression). More aggressive dogs had more amygdala activation data while watching their caregiver give food to a realistic fake dog than when they put the food in a bucket. This may have some similarity to human jealousy, adding to a growing body of evidence that differences in specific brain activities correlate with differences in canine temperament. The amygdala response habituates when an interaction is observed repeatedly, suggesting that repeated exposures may be a useful behavioral intervention with potentially aggressive dogs.
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Most mammalian species produce facial expressions. Historically, animal facial expressions have been considered inflexible and involuntary displays of emotional states rather than active attempts to communicate with others. In the current study, we aimed to test whether domestic dog facial expressions are subject to audience effects and/ or changes in response to an arousing stimulus (e.g. food) alone. We presented dogs with an experimental situation in which a human demonstrator was either attending to them or turned away, and varied whether she presented food or not. Dogs produced significantly more facial movements when the human was attentive than when she was not. The food, however, as a non-social but arousing stimulus, did not affect the dogs’ behaviour. The current study is therefore evidence that dogs are sensitive to the human’s attentional state when producing facial expressions, suggesting that facial expressions are not just inflexible and involuntary displays of emotional states, but rather potentially active attempts to communicate with others.
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Although the favourable characteristics of escitalopram as being the most selective serotonin reuptake inhibitor and having an increased therapeutic efficacy via binding on an additional allosteric binding site of the serotonin transporter, its dosing regimen has not yet been optimized for its use in dogs. This study aimed to estimate the optimal dosing frequency and the required dose for achieving 80% occupancy of the serotonin transporters in the basal ganglia. The dosing frequency was investigated by determining the elimination half-life after a four day oral pre-treatment period with 0.83 mg/kg escitalopram (3 administrations/day) and a subsequent i.v. injection 0.83 mg/kg. Blood samples were taken up to 12 hours after i.v. injection and the concentration of escitalopram in plasma was analysed via LC-MSMS. The dose-occupancy relationship was then determined by performing two PET scans in five adult beagles: a baseline PET scan and a second scan after steady state conditions were achieved following oral treatment with a specific dose of escitalopram ranging from 0.5 to 2.5 mg/kg/day. As the elimination half-life was determined to be 6.7 hours a dosing frequency of three administrations a day was proposed for the second part of the study. Further it was opted for a treatment period of four days, which well exceeded the minimum period to achieve steady state conditions. The optimal dosing regimen to achieve 80% occupancy in the basal ganglia and elicit a therapeutic effect, was calculated to be 1.85 mg/kg/day, divided over three administrations. Under several circumstances, such as insufficient response to other SSRIs, concurrent drug intake or in research studies focused on SERT, the use of escitalopram can be preferred over the use of the already for veterinary use registered fluoxetine, however, in case of long-term treatment with escitalopram, regularly cardiac screening is recommended.
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The overall goal of this work was to measure the efficacy of fMRI for predicting whether a dog would be a successful service dog. The training and imaging were performed in 49 dogs entering service training at 17–21 months of age. 33 dogs completed service training and were matched with a person, while 10 were released for behavioral reasons (4 were selected as breeders and 2 were released for medical reasons.) After 2 months of training, fMRI responses were measured while each dog observed hand signals indicating either reward or no reward and given by both a familiar handler and a stranger. Using anatomically defined ROIs in the caudate, amygdala, and visual cortex, we developed a classifier based on the dogs’ subsequent training outcomes. The classifier had a positive predictive value of 94% and a negative predictive value of 67%. The area under the ROC curve was 0.91 (0.80 with 4-fold cross-validation, P = 0.01), indicating a significant predictive capability. The magnitude of response in the caudate was positively correlated with a successful outcome, while the response in the amygdala depended on the interaction with the visual cortex during the stranger condition and was negatively correlated with outcome (higher being associated with failure). These results suggest that, as indexed by caudate activity, successful service dogs generalize associations to hand signals regardless who gives them but without excessive arousal as measured in the amygdala.
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Full-text available
The aim of this randomised, double-blind, placebo-controlled, clinical-field study was to evaluate the effect of dexmedetomidine oromucosal gel at subsedative doses in alleviation of noise-associated acute anxiety and fear in dogs. On New Year's Eve, 182 dogs with a history of acute anxiety and fear associated with fireworks received treatment as needed up to five times: 89 dogs received dexmedetomidine and 93 dogs received placebo. For the primary efficacy variables, dog owners assessed the overall treatment effect as well as signs and extent of anxiety and fear. The overall treatment effect was statistically significant (P<0.0001). An excellent or good treatment effect was reported for a higher proportion of dogs treated with dexmedetomidine (64/89, 72 per cent) than those receiving placebo (34/93, 37 per cent). Additionally, dexmedetomidine-treated dogs expressed significantly (P<0.0314) fewer signs of fear and anxiety despite the noise of fireworks. No local tolerance or clinical safety concerns occurred during the study. This study demonstrated that oromucosal dexmedetomidine at subsedative doses alleviates noise-associated acute anxiety and fear in dogs.
Article
Behavioral problems occur frequently in dogs and represent a significant threat to dog welfare. Anxiety, phobias, and fears comprise most of the canine behavioral conditions. The identification of an association between specific behavioral phenotypes and genetic variants of candidate genes would be a valuable tool in selection for dogs less susceptible to anxiety and fear, which may improve animal welfare. The DRD2 gene encodes the dopamine receptor 2. In this study, we found 8 SNPs in the DRD2 gene of the Havanese, a breed that shows large variation in a behavioral phenotype that manifests itself as a tendency to react fearfully by withdrawing in social situations. Significant associations were detected between 2 SNPs in exon 2 of the DRD2 gene and increased social fear in Havanese dogs (n = 158), as evaluated through observation by an external evaluator (respective allelic odds ratio: 4.35, 4.07) and through owner questionnaires (respective allelic odds ratio: 1.96, 2.2). Because different types of fear-related behavioral disorders commonly co-occur, the SNPs in exon 2 were also investigated for possible association to noise reactivity in 5 breeds: Havanese (n = 121), collie (n = 94), Irish soft-coated wheaten terrier (n = 44), Nova Scotia duck tolling retriever (n = 33), and standard poodle (n = 29). Significant associations were detected between SNPs in exon 2 of the DRD2 gene and noise reactivity in the Irish soft-coated wheaten terrier (respective allelic odds ratio: 2.64, 2.88) and collie (allelic odds ratio: 3.03). The same SNP alleles were associated with the beneficial phenotypes in the 3 breeds.
Article
Accurate phenotyping is a critical component of any behavioral research. We compared owner responses to the Canine Behavior and Research Questionnaire section on separation-related behaviors with video footage of their dogs (n = 31). A focal animal continuous recording method was used to describe the behavior of dogs during the first 25 minutes after owners' departure. Nine dogs were recorded in the presence of 1 or 2 conspecifics, whereas 23 other dogs were alone. There was a significant moderate positive correlation between the separation-related behavior questionnaire score and the percentage of time exhibiting behaviors suggestive of anxiety (rs = 0.438; P = 0.003). There was a similar moderate positive correlation between the questionnaire score and the first principal component score of recorded behaviors (rs = 0.385; P = 0.008). Correlations between the questionnaire responses for individual behaviors and the occurrence of those behaviors on video were higher for the more apparent behaviors such as destruction. Although the significant positive correlation between owner responses and the dog's behavior in the initial 25 minutes of being left alone is suggestive that owners may have an indicative knowledge of their dog's behavior during their absence, further work with more participants and repeated recordings is required.
Article
The objective of our research was to identify pet-, owner-, and environment-related risk factors for euthanasia or rehoming of dogs presenting to a veterinary behavioral service. In this retrospective clinical study, we examined 302 patient records for dog and owner demographics, presenting complaints, clinical diagnoses, home environment, behavioral and medical history, and disposition of the patient at 3 and 6 months after visit. Data were analyzed for the association of pet-, owner-, and environment-related factors with (A) owners having considered euthanasia before the behavioral consultation; (B) owners having considered either rehoming or euthanasia before the consultation; (C) owners having rehomed or euthanized the dog up to 3 months after the consultation; (D) owners having rehomed or euthanized the dog 6 months after the consultation. In order to individuate the risk factors for rehoming and euthanasia, logistic regression analysis was conducted on pet-, owner-, and environment-related factors significantly associated (P < 0.05) with A, B, C, and D. Canine variables associated with a greater risk of rehoming or euthanasia and/or the owners considering rehoming or euthanasia included heavier weight; mixed breed; aggression to familiar people over resources, resting places, or when groomed/medicated; aggression to unfamiliar people during interactions; a history of biting; and living in a family with children aged 13-17 years. Associated owner variables included the use of punishment-based training and previous consultation with a nonveterinary behaviorist or trainer. Several pet-, owner-, and environment-related factors, other than the behavioral problem itself, can increase the likelihood of rehoming and euthanasia. These findings can provide an indication to closely follow-up with owners after the behavioral consultation to minimize the likelihood of actual rejection of the dog.
Article
The purpose of this paper is to review a series of studies (N= 17) regarding the effects of using various methods when training dogs. The reviewed studies examined the differences between training methods (e.g., methods based on positive reinforcement, positive punishment, escape/avoidance, etc.) on a dog’s physiology, welfare, and behavior towards humans and other dogs. The reviewed studies included surveys, observational studies, and interventions. The results show that using aversive training methods (e.g., positive punishment and negative reinforcement) can jeopardize both the physical and mental health of dogs. In addition, while positive punishment can be effective, there is no evidence that it is more effective than positive reinforcement-based training. In fact, there is some evidence that the opposite is true. A few methodological concerns arose from the reviewed studies. Among them are small sample sizes, missing data on effect size, possible bias when coding behavior in observational studies, and the need to publish case reports of bodily damage caused by aversive training methods. In conclusion, those working with or handling dogs should rely on positive reinforcement methods, and avoid using positive punishment and negative reinforcement as much as possible.
Article
Noise reactivity is a common problem for dogs and may progress to true phobia. Survey studies report that some type of noise reaction occurs in up to half of all pet dogs throughout their lifetimes, indicating that noise reactivity and/or phobia is a welfare issue. Familial aggregations of affected dogs have been reported, and increased prevalence in certain breeds has been suggested. Reactivity to noise can severely compromise function in both pet and working dogs. Noise reactivity may be comorbid with many anxiety disorders for both canines and humans and is postulated to effect information processing in associated human, rodent, and dog conditions. Any putative effect of noise on information processing becomes a concern for problem solving and other aspects of cognition that are important to working dogs. Accordingly, we sought to phenotype 3 breeds of herding dogs commonly used for work as detection dogs, police and/or patrol dogs, search and rescue dogs, and/or service dogs: Australian shepherds (AUS), border collies (BOC), and German shepherds (GSD). We analyzed demographic information and behavioral responses to noises (guns, storms, and fireworks) known to provoke fearful or phobic responses for 59 AUS, 81 BOC, and 58 GSD, who were also included in a genetic analysis. Behaviors were compared using a metric constructed from information on type, frequency, and intensity of response, and the Anxiety Intensity Rank (AIR) score. Reactivity to noise was found to segregate in some family lines for the dogs in this study, although individuals expressed considerable variation in noise response. Such variation may be time and exposure dependent and presents a phenotyping challenge. In this study, the presence and intensity of reactivity as represented by AIR scores varied by breed but only slightly with age. The BOC studied were older, and BOC and AUS were more severely affected (higher AIR scores) than were GSD. Source and/or purpose of dog may also affect severity of affliction. Determination of crisp and accurate phenotypes is essential for understanding underlying genetic contributions. For noise reactivity and/or phobia, accurate phenotypes include age of onset and specific behavioral characterization. Standardized and objective assessments are essential for assessment of progression and comorbidity. Our data imply that accurate phenotypic assessment is possible at a relatively early age, providing for both humane treatment and accurate phenotyping that facilitates good genotyping.