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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.

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 comparison of human related communication skills of socialized canids may help to understand the evolution and the epigenesis of gesture comprehension in humans. To reconcile previously contradicting views on the origin of dogs' outstanding performance in utilizing human gestures, we suggest that dog-wolf differences should be studied in a more complex way. We present data both on the performance and the behaviour of dogs and wolves of different ages in a two-way object choice test. Characteristic behavioural differences showed that for wolves it took longer to establish eye contact with the pointing experimenter, they struggled more with the handler, and pups also bit her more before focusing on the human's signal. The performance of similarly hand-reared 8-week-old dogs and wolves did not differ in utilizing the simpler proximal momentary pointing. However, when tested with the distal momentary pointing, 4-month-old pet dogs outperformed the same aged hand reared wolves. Thus early and intensive socialisation does not diminish differences between young dogs and wolves in behaviour and performance. Socialised adult wolves performed similarly well as dogs in this task without pretraining. The success of adult wolves was accompanied with increased willingness to cooperate. Thus, we provide evidence for the first time that socialised adult wolves are as successful in relying on distal momentary pointing as adult pet dogs. However, the delayed emergence of utilising human distal momentary pointing in wolves shows that these wild canines react to a lesser degree to intensive socialisation in contrast to dogs, which are able to control agonistic behaviours and inhibition of actions in a food related task early in development. We suggest a \"synergistic\" hypothesis, claiming that positive feedback processes (both evolutionary and epigenetic) have increased the readiness of dogs to attend to humans, providing the basis for dog-human communication.

Forming eye contact is important in dog–human communication. In this study we measured what factors affect dogs’ propensity for forming eye contact with an experimenter. We investigated the effect of [1] cephalic index (head shape’s metric, indicator of higher visual acuity at the centre of the visual field), [2] breed function (visual cooperativeness), [3] age and [4] playfulness with strangers in 125 companion dogs. Cephalic index was measured individually and analysed as a continuous variable. Results showed that [1] dogs with a higher cephalic index (shorter head) established eye contact faster. Since cephalic index is highly variable even within a breed, using artificial head shape groups or breed average cephalic index values is not recommended. [2] Breed function also affected dogs’ performance: cooperative breeds and mongrels established eye contact faster than dogs from non-cooperative breeds. [3] Younger dogs formed eye contact faster than older ones. [4] More playful dogs formed eye contact faster. Our results suggest that several factors affect dogs’ interspecific attention, and therefore their visual communication ability.

Modern dog breeds show a trend towards shorter muzzles in the Western world. Using an international online questionnaire ( N  = 780), we examined how muzzle length, as the sole varying feature, influenced visual preference for dogs, using digitally morphed images of the same individuals. We investigated how preferences for shorter muzzles were influenced by demographics, personality, pre-existing attitudes towards brachycephalic dogs, awareness of health issues, and desired dog traits. Shorter muzzles were generally preferred. Associating more health problems with brachycephaly was negatively associated with preference for shorter muzzles, except among those with positive attitudes towards brachycephalic dogs. Preference was positively associated with positive attitudes towards brachycephalic dogs, older age, conscientiousness, extraversion, and appreciation of aesthetics; and negatively associated with dog-related professional expertise, openness, and a liking for control in dog keeping. Among respondents without prior attitudes, preference was positively linked to viewing the ideal dog as childlike, and negatively to valuing a humorous personality in the ideal dog. Our results show how age, dog-related profession, personality, health awareness, and attitudes towards brachycephalic dogs and the ideal dog intersect to shape preferences for shorter muzzle length. Our findings support the notion that individuals preferring shorter muzzles may prioritise aesthetic traits over health concerns.

In humans, age-related changes in personality occur in a non-random fashion with respect to their direction, timing, and magnitude. In dogs, there are still gaps in our knowledge about the detailed dynamics of age-related personality changes. We analysed the personality of 217 Border collies aged from 0.5 to 15 years both cross-sectionally and longitudinally using a test battery, to specify age periods when changes most prominently occur, assess the magnitude of changes, and analyse individual differences in personality change. We found that similar to humans, changes in personality occur unevenly during the dogs’ life course, however, their dynamics seems to be specific for each trait. Activity-independence decreased mostly from puppyhood (0.5–1 years) to adolescence (> 1–2 years), then continued to decrease in a slowing rate. Novelty seeking did not change markedly until middle age (> 3–6 years), then showed a steady linear decrease. Problem orientation increased strongly until middle age then showed no marked changes in later age periods. We also revealed individual differences in personality change over time, and showed that a few individuals with potential age-related impairments significantly affected the general age trajectory of some traits. These results raise caution against the over-generalisation of global age trends in dogs.

Non-REM bursts of activity in the sigma range (9–16 Hz) typical of sleep spindles predict learning in dogs, similar to humans and rats. Little is known, however, about the age-related changes in amplitude, density (spindles/minute) and frequency (waves/second) of canine spindles. We investigated a large sample (N = 155) of intact and neutered pet dogs of both sexes, varying in breed and age, searching for spindles in segments of non-REM sleep. We recorded EEG from both a frontal midline electrode (Fz) and a central midline electrode (Cz) in 55.5% of the dogs, in the remaining animals only the Fz electrode was active (bipolar derivation). A similar topography was observed for fast (≥13 Hz) spindle occurrence as in humans (fast spindle number, density on Cz > Fz). For fast spindles, density was higher in females, and increased with age. These effects were more pronounced among intact animals and on Fz. Slow spindle density declined and fast spindle frequency increased with age on Cz, while on Fz age-related amplitude decline was observed. The frequency of fast spindles on Fz and slow spindles on Cz was linked to both sex and neutering, suggesting modulation by sexual hormones. Intact females displayed higher frequencies than males and neutered females. Our findings support the argument that sigma bursts in the canine non-REM sleep are analogous to human sleep spindles, and suggest that slow and fast spindles display different trajectories related to age, of which an increase in frontal fast spindles is unique to dogs.

During social interactions, acoustic parameters of tetrapods’ vocalisations reflect the emotional state of the caller. Higher levels of spectral noise and the occurrence of irregularities (non-linear phenomena NLP) might be negative arousal indicators in alarm calls, although less is known about other distress vocalisations. Family dogs experience different levels of stress during separation from their owner and may vocalise extensively. Analysing their whines can provide evidence for the relationship between arousal and NLP. We recorded 167 family dogs’ separation behaviour including vocalisations, assessed their stress level based on behaviour and tested how these, their individual features, and owner reported separation-related problems (SRP) relate to their whines’ (N = 4086) spectral noise and NLP. Dogs with SRP produced NLP whines more likely. More active dogs and dogs that tried to escape produced noisier whines. Older dogs’ whines were more harmonic than younger ones’, but they also showed a higher NLP ratio. Our results show that vocal harshness and NLP are associated with arousal in contact calls, and thus might function as stress indicators. The higher occurrence of NLP in older dogs irrespective to separation stress suggests loss in precise neural control of the larynx, and hence can be a potential ageing indicator.

Body-awareness is one of the fundamental modules of self-representation. We investigated how body-awareness could contribute to dogs' decision making in a novel spatial problem where multiple solutions are possible. Family dogs (N = 68) had to obtain a treat from behind a transparent fence. They had two options: either detour around the fence (7 m), or take a shortcut through a doorway (2 m). We had three conditions: small door open, large door open, and doors closed. Our results indicated that dogs assess the size of the doorway, and if they find it too small, they decide to detour instead, while in the case of the open large door, they rather opted for the shortcut without hesitation. Shorter headed dogs tended to choose open doors more often, while longer headed dogs rather chose detours, probably because of their better peripheral vision. While body size awareness did not manifest differently in dogs with short or long heads, we showed for the first time a connection between head shape and physical cognition in dogs. We showed that dogs rely on their body-awareness in a naturalistic setting where multiple solutions exist simultaneously. Dogs make decisions without lengthy trial-and-error learning and choose between options based on their body-awareness.