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Relative duration of drowsiness plotted as a function of attachment scores measured in the Strange Situation Test. (The association is not significant after controlling for age).
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Affective neuroscience studies have demonstrated the impact of social interactions on sleep quality. In humans, trait-like social behaviors, such as attachment, are related to sleep brain activity patterns. Our aim was to investigate associations between companion dogs’ spontaneous brain activity during sleep (in the presence of the owner) and thei...
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... In other words, the willingness of participant dogs to follow task instructions, irrespective of whether their guardian was looking at them, was associated in a trait-like manner with alpha, beta, theta, delta frequency bands power during sleep. Carreiro et al. [54] used an adapted form of the Strange Situation Task (SST) [71] to index attachment levels of canine subjects and investigated whether derived attachment scores correlated with sleep activity features. They found associations between the level of attachment and the duration of NREM sleep, as the activity in certain frequency bands. ...
... As with olfaction, questions of cognitive control not only inform us about the neural underpinnings of canine cognition but also have significant practical value in the way humans communicate with and train dogs. Pre-sleep intensive activity, time and location of sleep, owner-rated hyperactivity [31,45] Alpha frequency power Age, cooperation, attachment [50,54] Beta and delta frequency power ...
The emerging field of canine cognitive neuroscience uses neuroimaging tools such as electroencephalography (EEG) and functional magnetic resonance imaging (fMRI) to map the cognitive processes of dogs to neural substrates in their brain. Within the past decade, the non-invasive use of EEG has provided real-time, accessible, and portable neuroimaging insight into canine cognitive processes. To promote systematization and create an overview of framings, methods and findings for future work, we provide a systematic review of non-invasive canine EEG studies (N=22), dissecting their study makeup, technical setup, and analysis frameworks and highlighting emerging trends. We further propose new directions of development, such as the standardization of data structures and integrating predictive modeling with descriptive statistical approaches. Our review ends by underscoring the advances and advantages of EEG-based canine cognitive neuroscience and the potential for accessible canine neuroimaging to inform both fundamental sciences as well as practical applications for cognitive neuroscience, working dogs, and human-canine interactions.
... The Strange Situation Test (SST)-a validated laboratory test (e.g., 36 ) with various precursors in the examination of social bonds across species 37 -is widely used to investigate attachment towards human individuals both intraspecifically (i.e. infant-mother relationships, e.g., 38 ) and interspecifically (e.g., dog-owner relationships [39][40][41] ). The SST has not yet been used, however, to test pigs. ...
Dogs exhibit human-analogue attachment to their owners, with similar function and mechanisms to that of infant-mother bond, but its origin is unclear. Comparative studies on socialised wolves and dogs emphasise genetic influence in dogs' preparedness for attachment to humans. We aimed to reveal if this genetic effect stems from general domestication or artificial selection that increased dogs' dependence on humans. We assessed and compared behavioural patterns of young companion pigs and dogs using a Strange Situation Test. Dogs but not pigs exhibited distinct behaviours towards their owner and a stranger along attachment-specific variables, so only dogs’ relevant behaviours fulfilled attachment criteria. From the observed behaviours, three factors were formed: Attachment (to the owner), Anxiety (in a strange situation), and Acceptance (of a stranger). Results indicate (1) higher Attachment scores in dogs than pigs, (2) greater Acceptance scores in pigs, (3) positive correlation of Attachment and Anxiety in both, (4) similar time tendency of pigs' Attachment and Acceptance scores. These suggest that in pigs, domestication and early exposure to human social stimuli did not trigger attachment to humans. Thus, along with species predispositions, the unique dog-owner attachment can be facilitated by artificial selection that increased dogs' dependence on humans.
... Carreiro and colleagues [25] studied the correlation between SST attachment and sleep patterns in mature dogs. They found that secure attachment in dogs was associated with more time spent in NREM sleep. ...
The close bond that can exist between humans and their dogs is an important aspect of the evolutionary, economic, and social connections between the two species. There is a need for a better understanding of the place of the dog within the human–dog bond and on ways the human–dog bond affects dog welfare. We conducted a scoping review to investigate to what extent and in what ways dog welfare is addressed in the research literature on the human–dog bond. We identified 706 publications on the human–dog bond from across the globe that were published from 2012 to 2023. We found that 246 of these 706 publications had a focus on dog welfare. Our review showed that the interplay of characteristics and backgrounds of owners/handlers and their dogs was linked to dog welfare in multiple, both positive and negative, ways. Our review is limited by the fact that most of the research that we reviewed involved pet dogs and in majority came from Western, Educated, Industrialized, Rich, Democratic (WEIRD) societies. There is a need for a better understanding of how the human–dog bond affects the welfare of working, assistance, and service dogs.
... sleep/wake states and derived parameters) and the power spectral density of dogs in a variety of different experiments investigating, e.g., the effects of physical activity (Kis et al. 2014;Bunford et al. 2018), learning (Kis et al. 2017a), emotional experiences (Kis et al. 2017b), location of sleep (Bunford et al. 2018), developmental changes (Reicher et al. 2021a), or first-nights ) on different sleep characteristics, as well as for describing the physiological changes associated with different sleep stages (Bálint et al. 2019). It has also been revealed that dogs show interhemispheric asymmetry during deep sleep (Reicher et al. 2021b)-certain EEG parameters are associated with the attachment of dogs toward their owners (Carreiro et al. 2022) and that sleep spindle characteristics are associated with learning, age, and sex in dogs (Iotchev et al. 2019. Recently, this methodology has also been successfully applied in studying the closest wild relative of dogs-the wolf-providing the first descriptive analysis of its natural sleep . ...
... This result is especially interesting, since although we expected that cats will be generally more anxious in a novel environment with unfamiliar people, cats participating in the present study represent a special subset of family cats in that they all have successfully passed the "habituation test" (Uccheddu et al. 2022) prior to the sleep measurement, showing that they could conform to the lab environment. This result may indicate that while owners provide a "safe haven" for dogs in unfamiliar or even potentially dangerous situations and places (Gácsi et al. 2013;Carreiro et al. 2022), the same may not hold true for cats. Other reasons may include a more pronounced manifestation of the so-called first-night effect in cats than in dogs caused by, e.g., the unfamiliar surroundings, discomfort (e.g. ...
We have successfully measured the sleep electroencephalogram (EEG) of 12 family cats during an afternoon nap using a completely noninvasive methodology originally developed and validated for family dogs. Extracting both macrostructural and spectral sleep variables from the acquired data, we: (1) provided a descriptive analysis of sleep structure in cats and the power spectral density (PSD) distribution considering 3 sleep stages—drowsiness, non-rapid eye movement (NREM), and rapid eye movement (REM) sleep; and (2) compared the results to those obtained in family dogs measured under the same conditions and using the same methodology. Importantly, our description of sleep structure and PSD distribution in cats proved to be comparable to those of earlier invasive studies, highlighting that appropriate noninvasive methodologies may provide a viable alternative to those that are invasive in some cases. While no macrostructural differences were found between the sleep of cats and dogs, and the characteristic PSDs were mostly similar across sleep stages within the 2 species, the high-frequency resolution comparison of PSD distributions revealed differences between the 2 species in all sleep stages (concerning the delta, theta, alpha, sigma, and beta bands in drowsiness and NREM sleep; and the delta, alpha, and sigma bands in REM sleep). Potential factors underlying these differences are discussed, including differences in circadian rhythms, sleep homeostatic regulation, experienced stress, or even differential attitudes toward owners—highlighting important links between sleep characteristics and often more complex neural and behavioral features.
... The stress related explanation was also supported by the results of Mäses and Wascher 47 who reported no decline in the choices of cats in their familiar environment when they used food distraction instead of restraint to keep shelter cats at the starting point. Besides the direct biases, the testability of dogs could have been enhanced by the presence of their O, providing a secure base/safe haven in a strange novel environment 73,74 , while the lack of such a role of the O may have contributed to the cats feeling uncomfortable in the laboratory 41 . Cats' more difficult habituation to the unfamiliar environment may originate from two factors. ...
The dog is a so far unique species to study interspecific communication and a promising evolutionary model for preverbal human communication. Recently cats were reported to show some similar skills to dogs. Here we directly compared both the testability and the success of companion dogs and cats in relying on human distal pointing gestures. Due to differences in their domestication, social and ecological background, and developmental processes, we expected better performance from dogs compared to cats. Using an object-choice task, cats made considerably fewer choices than dogs in the laboratory environment, and their tendency to make a choice declined during trials. They were slightly more testable at home, where their willingness to choose did not decrease over time. Dogs made more successful choices than cats, both at the group and individual level, irrespective of the type of the pointing gesture. Older cats were more successful. Despite the two species’ rather similar role nowadays as human companions, our results support previous findings suggesting that, compared to the dog, the cat is a less ideal model to study some human communicative abilities in a laboratory environment.
... Dogs provide further evidence for possessing a functional attachment system. Earlier research from multiple laboratories, using the same methodology (SST) and large number of adult dogs has established that the attachment system of dogs and humans share most significant features; the owner's role as a secure base (Palmer & Custance, 2008) and as safe haven (Gácsi et al., 2013), and that dogs even develop new attachment relationship in adulthood (shelter dogs: Carreiro et al., 2022;Gácsi et al., 2001, and guide dogs for the blind: Valsecchi et al., 2010). Dogs' attachment scores, calculated from the SST, were reported to be associated with their brain activity during sleep at an unfamiliar place with the owner (Carreiro et al., 2022) and with their neural reward responses to verbal praise in an fMRI study (Gábor et al., 2021). ...
... Earlier research from multiple laboratories, using the same methodology (SST) and large number of adult dogs has established that the attachment system of dogs and humans share most significant features; the owner's role as a secure base (Palmer & Custance, 2008) and as safe haven (Gácsi et al., 2013), and that dogs even develop new attachment relationship in adulthood (shelter dogs: Carreiro et al., 2022;Gácsi et al., 2001, and guide dogs for the blind: Valsecchi et al., 2010). Dogs' attachment scores, calculated from the SST, were reported to be associated with their brain activity during sleep at an unfamiliar place with the owner (Carreiro et al., 2022) and with their neural reward responses to verbal praise in an fMRI study (Gábor et al., 2021). Thus, we can conclude that, unless more systematic empirical research on wolves indicates otherwise, the dog-human bond has specific characteristics that have not been observed so far in wolf pups, and especially not in adult wolves. ...
In their recent paper, Hansen Wheat et al. ( Ecology and Evolution , 2022, 12 , e9299) claimed that hand raised 23‐week‐old wolves showed the same attachment behaviour towards their handler in the Strange Situation Test (SST) ( Determinants of infant behavior , 1969, 4 , 111) as dogs. At first glance, their results seem to contradict previous findings that domestication caused a unique change in social‐affiliative behaviours in dogs ( Animal Behaviour , 2005, 70 , 1367). We argue that no persuading evidence was presented to claim that “wolves can show attachment behaviours towards humans comparable to those of dogs”. When dealing with a behaviour system ( Child Development , 1977, 48 , 1184), the subjects' behaviour must meet consistent criteria ( Behavioural and Brain Science , 1978, 3 , 417), and a few behavioural preferences should not be used to claim the presence of an attachment system, especially, if the experiment violates basic assumptions of the original test. We believe the intriguing scientific question is whether the dog‐owner relationship is qualitatively different from what could be observed in the wolf‐hand raiser relation. Assessing all available data, our answer is still yes; dogs are unique in this respect.
... In other words, the willingness of participant dogs to follow task instructions, irrespective of whether their guardian was looking at them, was associated in a trait-like manner with alpha, beta, theta, delta frequency bands power during sleep. Carreiro et al. [18] used an adapted form of the Strange Situation Task (SST) [2] . CC-BY 4.0 International license available under a (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. ...
A bstract
The emerging field of canine cognitive neuroscience uses neuroimaging tools such as electroencephalography (EEG) and functional magnetic resonance imaging (fMRI) to map the cognitive processes of dogs to neural substrates in their brain. Within the past decade, the non-invasive use of EEG has provided real-time, accessible, and portable neuroimaging insight into canine cognitive processes. To promote systematization and create an overview of framings, methods and findings for future work, we provide a systematic review of non-invasive canine EEG studies (N=22), dissecting their study makeup, technical setup, and analysis frameworks and highlighting emerging trends. We further propose new directions of development, such as the standardization of data structures and integrating predictive modeling with descriptive statistical approaches. Our review ends by underscoring the advances and advantages of EEG-based canine cognitive neuroscience and the potential for accessible canine neuroimaging to inform both fundamental sciences as well as practical applications for cognitive neuroscience, working dogs, and human-canine interactions.
... Thus, ADHD has been seen as a socio-emotional dysfunction. These would be important to consider in future canine research as, e.g., the dog-owner social bond is in association with altered SWA dynamics during sleep (alpha-delta anticorrelation 28 ). ...
Subjective sleep disturbances are reported by humans with attention-deficit/hyperactivity disorder (ADHD). However, no consistent objective findings related to sleep disturbances led to the removal of sleep problems from ADHD diagnostic criteria. Dogs have been used as a model for human ADHD with questionnaires validated for this purpose. Also, their sleep physiology can be measured by non-invasive methods similarly to humans. In the current study, we recorded spontaneous sleep EEG in family dogs during a laboratory session. We analyzed the association of sleep macrostructure and deep sleep (NREM) slow-wave activity (SWA) with a validated owner-rated ADHD questionnaire, assessing inattention (IA), hyperactivity/impulsivity (H/I) and total (T) scores. Higher H/I and T were associated with lower sleep efficiency and longer time awake after initial drowsiness and NREM. IA showed no associations with sleep variables. Further, no association was found between ADHD scores and SWA. Our results are in line with human studies in which poor sleep quality reported by ADHD subjects is associated with some objective EEG macrostructural parameters. This suggests that natural variation in dogs’ H/I is useful to gain a deeper insight of ADHD neural mechanisms.
... Thus, ADHD has been seen as a socio-emotional dysfunction. These would be important to consider in future canine research as, e.g., the dog-owner social bond is in association with altered SWA dynamics during sleep (alpha-delta anticorrelation 28 ). ...
Subjective sleep disturbances are reported by humans with attention-deficit/hyperactivity disorder (ADHD). However, no consistent objective findings related to sleep disturbances led to sleep problems removal from ADHD diagnostic criteria. As a model for human ADHD with questionnaires validated for this purpose, dogs have been used also because their sleep physiology can be measured by non-invasive methods similarly to humans. We recorded spontaneous sleep EEG in family dogs during a laboratory session. We analyzed the association of sleep macrostructure and deep sleep (NREM) slow-wave activity (SWA) with a validated owner-rated ADHD questionnaire, assessing inattention (IA), hyperactivity/impulsivity (H/I) and total (T) scores. Higher H/I and T were associated with lower sleep efficiency and longer time awake after initial drowsiness and NREM. IA showed no associations with sleep variables. Further, no association was found between ADHD scores and SWA. Our results are in line with human studies in which poor sleep quality reported by ADHD subjects is associated with some objective EEG macrostructural parameters. This suggests that dogs’ H/I natural variation is useful to gain a deeper insight of ADHD neural mechanisms.
