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The shape of the cranium is one of the most notable physical changes induced in domestic dogs through selective breeding and is measured using the cephalic index (CI). High CI (a ratio of skull width to skull length > 60) is characterized by a short muzzle and flat face and is referred to as brachycephaly. Brachycephalic dogs display some potentially harmful changes in neuroanatomy, and there are implications for differences in behavior, as well. The path from anatomy to cognition, however, has not been charted in its entirety. Here, we report that sleep-physiological markers of white-matter loss (high delta power, low frontal spindle frequency, i.e., spindle waves/s), along with a spectral profile for REM (low beta, high delta) associated with low intelligence in humans, are each linked to higher CI values in the dog. Additionally, brachycephalic subjects spent more time sleeping, suggesting that the sleep apnea these breeds usually suffer from increases daytime sleepiness. Within sleep, more time was spent in the REM sleep stage than in non-REM, while REM duration was correlated positively with the number of REM episodes across dogs. It is currently not clear if the patterns of sleep and sleep-stage duration are mainly caused by sleep-impairing troubles in breathing and thermoregulation, present a juvenile-like sleeping profile, or are caused by neuro-psychological conditions secondary to the effects of brachycephaly, e.g., frequent REM episodes are known to appear in human patients with depression. While future studies should more directly address the interplay of anatomy, physiology, and behavior within a single experiment, this represents the first description of how the dynamics of the canine brain covary with CI, as measured in resting companion dogs using a non-invasive sleep EEG methodology. The observations suggest that the neuroanatomical changes accompanying brachycephaly alter neural systems in a way that can be captured in the sleep EEG, thus supporting the utility of the latter in the study of canine brain health and function., (© 2023. The Author(s).)

Dogs live in a complex social environment where they regularly interact with conspecific and heterospecific partners. Awake dogs are able to process a variety of information based on vocalisations emitted by dogs and humans. Whether dogs are also able to process such information while asleep, is unknown. In the current explorative study, we investigated in N = 13 family dogs, neural response to conspecific and human emotional vocalisations. Data were recorded while dogs were asleep, using a fully non-invasive event-related potential (ERP) paradigm. A species (between 250-450 and 600-800 ms after stimulus onset) and a species by valence interaction (between 550 to 650 ms after stimulus onset) effect was observed during drowsiness. A valence (750-850 ms after stimulus onset) and a species x valence interaction (between 200 to 300 ms and 450 to 650 ms after stimulus onset) effect was also observed during non-REM specific at the Cz electrode. Although further research is needed, these results not only suggest that dogs neurally differentiate between differently valenced con- and heterospecific vocalisations, but they also provide the first evidence of complex vocal processing during sleep in dogs. Assessment and detection of ERPs during sleep in dogs appear feasible., (© 2023. Springer Nature Limited.)

Recently, increased interest and efforts were observed in describing the possible interaction between sleep and emotions. Human and animal model studies addressed the implication of both sleep patterns and emotional processing in neurophysiology and neuropathology in suggesting a bidirectional interaction intimately modulated by complex mechanisms and factors. In this context, we aimed to discuss recent evidence and possible mechanisms implicated in this interaction, as provided by both human and animal models in studies. In addition, considering the affective component of brain physiological patterns, we aimed to find reasonable evidence in describing the two-way association between comorbid sleep impairments and psychiatric disorders. The main scientific literature databases (PubMed/Medline, Web of Science) were screened with keyword combinations for relevant content taking into consideration only English written papers and the inclusion and exclusion criteria, according to PRISMA guidelines. We found that a strong modulatory interaction between sleep processes and emotional states resides on the activity of several key brain structures, such as the amygdala, prefrontal cortex, hippocampus, and brainstem nuclei. In addition, evidence suggested that physiologically and behaviorally related mechanisms of sleep are intimately interacting with emotional perception and processing which could advise the key role of sleep in the unconscious character of emotional processes. However, further studies are needed to explain and correlate the functional analysis with causative and protective factors of sleep impairments and negative emotional modulation on neurophysiologic processing, mental health, and clinical contexts.

Age-related differences in dog sleep and the age at which dogs reach adulthood as indexed by sleep electrophysiology are unknown. We assessed, in (1) a Juvenile sample (n = 60) of 2-14-month-old dogs (weight range: 4-68 kg), associations between age, sleep macrostructure, and non-rapid eye movement (NREM) EEG power spectrum, whether weight moderates associations, and (2) an extended sample (n = 91) of 2-30-months-old dogs, when sleep parameters stabilise. In Juvenile dogs, age was positively associated with time in drowsiness between 2 and 8 months, and negatively with time in rapid eye movement (REM) sleep between 2 and 6 months. Age was negatively associated with delta and positively with theta and alpha power activity, between 8 and 14 months. Older dogs exhibited greater sigma and beta power activity. Larger, > 8-month-old dogs had less delta and more alpha and beta activity. In extended sample, descriptive data suggest age-related power spectrum differences do not stabilise by 14 months. Drowsiness, REM, and delta power findings are consistent with prior results. Sleep electrophysiology is a promising index of dog neurodevelopment; some parameters stabilise in adolescence and some later than one year. Determination of the effect of weight and timing of power spectrum stabilisation needs further inquiry. The dog central nervous system is not fully mature by 12 months of age., (© 2021. The Author(s).)

Although a positive link between sleep spindle occurrence and measures of post-sleep recall (learning success) is often reported for humans and replicated across species, the test-retest reliability of the effect is sometimes questioned. The largest to date study could not confirm the association, however methods for automatic spindle detection diverge in their estimates and vary between studies. Here we report that in dogs using the same detection method across different learning tasks is associated with observing a positive association between sleep spindle density (spindles/minute) and learning success. Our results suggest that reducing measurement error by averaging across measurements of density and learning can increase the visibility of this effect, implying that trait density (estimated through averaged occurrence) is a more reliable predictor of cognitive performance than estimates based on single measures.

The importance of dogs (Canis familiaris) in sleep research is primarily based on their comparability with humans. In spite of numerous differences, dogs' comparable sleep pattern, as well as several phenotypic similarities on both the behavioural and neural levels, make this species a most feasible model in many respects. Our aim was to investigate whether the so-called first-night effect, which in humans manifests as a marked macrostructure difference between the first and second sleep occasions, can be observed in family dogs. We used a non-invasive polysomnographic method to monitor and compare the characteristics of dogs' (N = 24) 3-hr-long afternoon naps on three occasions at the same location. We analysed how sleep macrostructure variables differed between the first, second and third occasions, considering also the effects of potential confounding variables such as the dogs' age and sleeping habits. Our findings indicate that first-night effect is present in dogs' sleep architecture, although its specifics somewhat deviate from the pattern observed in humans. Sleep macrostructure differences were mostly found between occasions 1 and 3; dogs slept more, had less wake after the first drowsiness episode, and reached drowsiness sleep earlier on occasion 3. Dogs, which had been reported to sleep rarely not at home, had an earlier non-rapid eye movement sleep, a shorter rapid eye movement sleep latency, and spent more time in rapid eye movement sleep on occasion 3, compared with occasion 1. Extending prior dog sleep data, these results help increase the validity of further sleep electroencephalography investigations in dogs., (© 2020 The Authors. Journal of Sleep Research published by John Wiley & Sons Ltd on behalf of European Sleep Research Society.)

Dogs have outstanding capabilities to read human emotional expressions, both vocal and facial. It has also been shown that positively versus negatively valenced dog-human social interactions substantially affect dogs' subsequent sleep. In the present study, we manipulated dogs' (N = 15, in a within subject design) sleep structure by specifically disrupting REM versus Non-REM sleep, while maintaining equal sleep efficiency (monitored via non-invasive polysomnography). We found that both the number of awakenings as well as relative Non-REM (but not relative REM) duration influenced dogs' viewing patterns in a task where sad and happy human faces were simultaneously projected with sad or happy human voice playbacks. In accordance with the emotion laterality hypothesis, the interaction between sound valence and Non-REM sleep duration was specific to images projected to the left (regardless of image-sound congruency). These results reveal the first evidence of a causal link between sleep structure and inter-specific emotion-processing in the family dog.

In both humans and dogs sleep spindle occurrence between acquisition and recall of a specific memory correlate with learning performance. However, it is not known whether sleep spindle characteristics are also linked to performance beyond the span of a day, except in regard to general mental ability in humans. Such a relationship is likely, as both memory and spindle expression decline with age in both species (in dogs specifically the density and amplitude of slow spindles). We investigated if spindle amplitude, density (spindles/minute) and/or frequency (waves/second) correlate with performance on a short-term memory and a reversal-learning task in old dogs (> 7 years), when measurements of behavior and EEG were on average a month apart. Higher frequencies of fast (≥ 13 Hz) spindles on the frontal and central midline electrodes, and of slow spindles (≤ 13 Hz) on the central midline electrode were linked to worse performance on a reversal-learning task. The present findings suggest a role for spindle frequency as a biomarker of cognitive aging across species: Changes in spindle frequency are associated with dementia risk and onset in humans and declining learning performance in the dog.

The domestic dog (Canis familiaris) is a promising animal model. Yet, the canine neuroscience literature is predominantly comprised of studies wherein (semi-)invasive methods and intensive training are used to study awake dog behavior. Given prior findings with humans and/or dogs, our goal was to assess, in 16 family dogs (1.5-7 years old; 10 males; 10 different breeds) the effects of pre-sleep activity and timing and location of sleep on sleep electrophysiology. All three factors had a main and/or interactive effect on sleep macrostructure. Following an active day, dogs slept more, were more likely to have an earlier drowsiness and NREM, and spent less time in drowsiness and more time in NREM and REM. Activity also had location- and time of day-specific effects. Time of day had main effects; at nighttime, dogs slept more and spent less time in drowsiness and awake after first drowsiness, and more time in NREM and in REM. Location had a main effect; when not at home, REM sleep following a first NREM was less likely. Findings are consistent with and extend prior human and dog data and have implications for the dog as an animal model and for informing future comparative research on sleep.

The effects of emotionally valenced events on sleep physiology are well studied in humans and laboratory rodents. However, little is known about these effects in other species, despite the fact that several sleep characteristics differ across species and thus limit the generalizability of such findings. Here we studied the effect of positive and negative social experiences on sleep macrostructure in dogs, a species proven to be a good model of human social cognition. A non-invasive polysomnography method was used to collect data from pet dogs ( n = 16) participating in 3-hour-long sleep occasions. Before sleep, dogs were exposed to emotionally positive or negative social interactions (PSI or NSI) in a within-subject design. PSI consisted of petting and ball play, while NSI was a mixture of separation, threatening approach and still face test. Sleep macrostructure was markedly different between pre-treatment conditions, with a shorter sleep latency after NSI and a redistribution of the time spent in the different sleep stages. Dogs' behaviour during pre-treatments was related to the macrostructural difference between the two occasions, and was further modulated by individual variability in personality. This result provides the first direct evidence that emotional stimuli affect subsequent sleep physiology in dogs., (© 2017 The Author(s).)

Recently dogs (Canis familiaris) have been demonstrated to be a promising model species for studying human behavior as they have adapted to the human niche and developed human-like socio-cognitive skills. Research on dog behavior, however, has so far almost exclusively focused on awake functioning. Here we present a self-developed non-invasive canine polysomnography method that can easily be applied to naive pet dogs. N=22 adult pet dogs (with their owners present) and N=12 adult humans participated in Study I. From these subjects, N=7 dogs returned on two more occasions for Study II. In Study I, we give a descriptive analysis of the sleep electroencephalogram of the dog and compare it to human data. In order to validate our canine polysomnography method in Study II, we compare the sleep macrostructure and the EEG spectrum of dogs after a behaviorally active day without sleep versus passive day with sleep. In Study I, we found that dogs' sleep EEG resembled that of human subjects and was generally in accordance with previous literature using invasive technology. In Study II, we show that similarly to previous results on humans daytime load of novel experiences and sleep deprivation affects the macrostructural and spectral aspects of subsequent sleep. Our results validate the family dog as a model species for studying the effects of pre-sleep activities on the EEG pattern under natural conditions and, thus, broaden the perspectives of the rapidly growing fields of canine cognition and sleep research., (Copyright © 2014 Elsevier Inc. All rights reserved.)

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. [ABSTRACT FROM AUTHOR]

Functional hemispheric asymmetry was evidenced in many species during sleep. Dogs seem to show hemispheric asymmetry during wakefulness; however, their asymmetric neural activity during sleep was not yet explored. The present study investigated interhemispheric asymmetry in family dogs using non-invasive polysomnography. EEG recordings during 3-h-long afternoon naps were carried out (N = 19) on two occasions at the same location. Hemispheric asymmetry was assessed during NREM sleep, using bilateral EEG channels. To include periods with high homeostatic sleep pressure and to reduce the variance of the time spent in NREM sleep between dogs, the first two sleep cycles were analysed. Left hemispheric predominance of slow frequency range was detected in the first sleep cycle of sleep recording 1, compared to the baseline level of zero asymmetry as well as to the first sleep cycle of sleep recording 2. Regarding the strength of hemispheric asymmetry, we found greater absolute hemispheric asymmetry in the second sleep cycle of sleep recording 1 and 2 in the frequency ranges of alpha, sigma and beta, compared to the first sleep cycle. Differences between sleep recordings and consecutive sleep cycles might be indicative of adaptation-like processes, but do not closely resemble the results described in humans. [ABSTRACT FROM AUTHOR]

The effects of emotionally valenced events on sleep physiology are well studied in humans and laboratory rodents. However, little is known about these effects in other species, despite the fact that several sleep characteristics differ across species and thus limit the generalizability of such findings. Herewe studied the effect of positive and negative social experiences on sleep macrostructure in dogs, a species proven to be a good model of human social cognition. A non-invasive polysomnography method was used to collect data from pet dogs (n = 16) participating in 3-hour-long sleep occasions. Before sleep, dogs were exposed to emotionally positive or negative social interactions (PSI or NSI) in a withinsubject design. PSI consisted of petting and ball play, while NSI was a mixture of separation, threatening approach and still face test. Sleep macrostructure was markedly different between pre-treatment conditions, with a shorter sleep latency after NSI and a redistribution of the time spent in the different sleep stages. Dogs' behaviour during pre-treatmentswas related to themacrostructural difference between the two occasions, andwas further modulated by individual variability in personality. This result provides the first direct evidence that emotional stimuli affect subsequent sleep physiology in dogs. [ABSTRACT FROM AUTHOR]

Thefirst-night effect—marked differences between the first- and the second-night sleep spent in a laboratory—is a widely known phenomenon that accounts for the common practice of excluding the first-night sleep from any polysomnographic analysis. The extent to which the first-night effect is present in a participant, as well as its duration (1 or more nights), might have diagnostic value and should account for different protocols used for distinct patient groups. This study investigated the first-night effect on nightmare sufferers (NM;N= 12) and healthy controls (N= 15) using both objective (2-night-long polysomnography) and subjective (Groningen Sleep Quality Scale for the 2 nights spent in the laboratory and 1 regular night spent at home) methods. Differences were found in both the objective (sleep efficiency, wakefulness after sleep onset, sleep latency, Stage-1 duration, Stage-2 duration, slow-wave sleep duration, and REM duration) and subjective (self-rating) variables between the 2 nights and the 2 groups, with a more pronounced first-night effect in the case of the NM group. Furthermore, subjective sleep quality was strongly related to polysomnographic variables and did not differ among 1 regular night spent at home and the second night spent in the laboratory. The importance of these results is discussed from a diagnostic point of view. [ABSTRACT FROM AUTHOR]

Simple Summary: Sleep alterations are known to be severe accompanying symptoms of many human psychiatric conditions, and validated clinical protocols are in place for their diagnosis and treatment. However, sleep monitoring is not yet part of standard veterinary practice, and the possible importance of sleep-related physiological alterations for certain behavioral problems in pets is seriously understudied. Recently, a non-invasive electroencephalography (EEG) method was developed for pet dogs that is well-suited for untrained individuals. This so called polysomnography protocol could easily be implemented in veterinary diagnosis. However, in order to make the procedure more effective and standardized, methodological questions about the validity and reliability of data processing need to be answered. As a first step, the present study tests the effect of several factors on the manual scoring of the different sleep stages (a standard procedure adopted from human studies). Scoring the same recordings but varying the number of EEG channels visible to the scorer (emulating the difference between single channel versus four channel recordings) resulted in significant differences in hypnograms. This finding suggests that using more recording electrodes may provide a more complete picture of dog brain electrophysiological activity. Visual sleep staging by three different expert raters also did not provide a full agreement, but despite this, there were no significant differences between raters in important output values such as sleep structure and the spectral features of the EEG. This suggests that the non-invasive canine polysomnography method is ready to be implemented for veterinary use, but there is room for further refinement of the data processing. Here, we describe which parts of the sleep recording yield the lowest agreement and present the first form of an automated method that can reliably distinguish awake from sleep stages and could thus accelerate the time-consuming manual data processing. The translation of the findings into clinical practice will open the door to the more effective diagnosis and treatment of disorders with sleep-related implications. Non-invasive polysomnography recording on dogs has been claimed to produce data comparable to those for humans regarding sleep macrostructure, EEG spectra and sleep spindles. While functional parallels have been described relating to both affective (e.g., emotion processing) and cognitive (e.g., memory consolidation) domains, methodologically relevant questions about the reliability of sleep stage scoring still need to be addressed. In Study 1, we analyzed the effects of different coders and different numbers of visible EEG channels on the visual scoring of the same polysomnography recordings. The lowest agreement was found between independent coders with different scoring experience using full (3 h-long) recordings of the whole dataset, and the highest agreement within-coder, using only a fraction of the original dataset (randomly selected 100 epochs (i.e., 100 × 20 s long segments)). The identification of drowsiness was found to be the least reliable, while that of non-REM (rapid eye movement, NREM) was the most reliable. Disagreements resulted in no or only moderate differences in macrostructural and spectral variables. Study 2 targeted the task of automated sleep EEG time series classification. Supervised machine learning (ML) models were used to help the manual annotation process by reliably predicting if the dog was sleeping or awake. Logistic regression models (LogREG), gradient boosted trees (GBT) and convolutional neural networks (CNN) were set up and trained for sleep state prediction from already collected and manually annotated EEG data. The evaluation of the individual models suggests that their combination results in the best performance: ~0.9 AUC test scores. [ABSTRACT FROM AUTHOR]

Simple Summary: It is common knowledge that negative emotions in humans are accompanied by both impaired subjective experience as well as maladaptive changes in behavior and physiology. The present paper investigates heart rate—one of the most commonly used emotion-related physiology measures—in the family dog, with the aim of uncovering its potential relationship with emotions. Sleep recordings were conducted following a positive versus a negative social interaction, as sleep alternations are one of the most conspicuous changes in response to negative affect. We observed differences in heart rate following the positive versus negative interactions, however these were only apparent during wakefulness, but not during sleep. The domestic dog (Canis familiaris) has been shown to both excel in recognising human emotions and produce emotion-related vocalisations and postures that humans can easily recognise. However, little is known about the effect of emotional experiences on subsequent sleep physiology, a set of phenomena heavily interrelated with emotions in the case of humans. The present paper examines heart rate (HR) and heart rate variability (HRV) during dogs’ sleep, measures that are influenced by both positive and negative emotions in awake dogs. In Study I, descriptive HR and HRV data is provided on N = 12 dogs about the different sleep stages (wake, drowsiness, non-rapid eye movement (non-REM), REM; scoring based on electroencephalogram (EEG) data). We conclude that wakefulness is characterised by higher HR and lower HRV compared to all sleep stages. Furthermore, drowsiness is characterised by higher HR and lower HRV than non-REM and REM, but only if the electrocardiogram (ECG) samples are taken from the first occurrence of a given sleep stage, not when the longest periods of each sleep stage are analysed. Non-REM and REM sleep were not found to be different from each other in either HR or HRV parameters. In Study II, sleep HR and HRV measures are compared in N = 16 dogs after a positive versus negative social interaction (within-subject design). The positive social interaction consisted of petting and ball play, while the negative social interaction was a mixture of separation, threatening approach and still face test. Results are consistent with the two-dimensional emotion hypothesis in that following the intense positive interaction more elevated HR and decreased HRV is found compared to the mildly negative (lower intensity) interaction. However, although this trend can be observed in all sleep stages except for REM, the results only reach significance in the wake stage. In sum, the present findings suggest that HR and HRV are possible to measure during dogs’ sleep, and can potentially be used to study the effect of emotions not only during but also after such interactions. [ABSTRACT FROM AUTHOR]