Dogs' social susceptibility is differentially affected by various dog-Human interactions. A study on family dogs, former shelter dogs and therapy dogs.

When pre-treated with social stimuli prior to testing, dogs are more susceptible to human influence in a food preference task. This means, after a positive social interaction they are more willing to choose the smaller amount of food indicated by the human, as opposed to their baseline preference for the bigger amount. In the current study we investigate if and how various forms of social interaction modulate choices in the same social susceptibility task, testing dogs with varying early life history (pet dogs, therapy dogs, former shelter dogs). In line with previous studies, dogs in general were found to be susceptible to human influence as reflected in the reduced number of "bigger" choices in the human influence, compared to baseline, trials. This was true not only for pet dogs with a normal life history, but also for dogs adopted from a shelter. Therapy dogs, however, did not uniformly change their preference for the bigger quantity of food in the human influence trials; they only did so if prior to testing they had been pre-treated with social stimuli by their owner (but not by a stranger). Pet dogs were also more influenced after pre-treatment with social stimuli by their owner compared to ignoring and separation; however after pre-treatment by a stranger their behaviour did not differ from ignoring and separation. Former shelter dogs on the other hand were equally influenced regardless of pre-treatment by owner versus stranger. In summary these results show that dogs' social susceptibility is modulated by both interactions immediately preceding the test as well as by long term social experiences.

Methodological comparison of cancellation versus two-way choice spatial attention tests in humans and dogs.

Introduction: Behavioural problems in family dogs are amongst the leading reasons for relinquishment to shelters which adversely affects animal welfare. Recent research suggests that certain problematic behavioural patterns might be analogous to human psychiatric disorders. Veterinary diagnosis of such conditions, however, is scarce, probably due to the lack of appropriate measurement tools. The current study focuses on dog behaviour resembling the human hemispatial neglect condition, which manifests itself as a deficit in attention to and awareness of one side of the space. Methods: Healthy human subjects (N = 21) and adult family dogs (N = 23) were tested with tools aimed to measure spatial attention. Tests administered to humans included validated paper and pencil neuropsychological tools to assess hemispatial neglect (cancellation tasks), as well as the canine version of that task (visuo-spatial search task). Dogs were tested with the same visuo-spatial search task as well as a two-way choice task. Results: Results show that both in case of dogs and humans the visuo-spatial search task detects individual variation in subjects' side preferences. However, subjects' performance in the different tasks were not related.

The influence of voice familiarity and linguistic content on dogs' ability to follow human voice direction.

Domestic dogs are well-known for their abilities to utilize human referential cues for problem solving, including following the direction of human voice. This study investigated whether dogs can locate hidden food relying only on the direction of human voice and whether familiarity with the speaker (owner/stranger) and the relevance of auditory signal features (ostensive addressing indicating the intent for communication to the receiver; linguistic content) affect performance. N = 35 dogs and their owners participated in four conditions in a two-way object choice task. Dogs were presented with referential auditory cues representing different combinations of three contextual parameters: the (I) 'familiarity with the human informant' (owner vs. stranger), the (II) communicative function of attention getter (ostensive addressing vs. non-ostensive cueing) and the (III) 'tone and content of the auditory cue' (high-pitched/potentially relevant vs. low-pitched/potentially irrelevant). Dogs also participated in a 'standard' pointing condition where a visual cue was provided. Significant differences were observed between conditions regarding correct choices and response latencies, suggesting that dogs' response to auditory signals are influenced by the combination of content and intonation of the message and the identity of the speaker. Dogs made correct choices the most frequently when context-relevant auditory information was provided by their owners and showed less success when auditory signals were coming from the experimenter. Correct choices in the 'Pointing' condition were similar to the experimenter auditory conditions, but less frequent compared to the owner condition with potentially relevant auditory information. This was paralleled by shorter response latencies in the owner condition compared to the experimenter conditions, although the two measures were not related. Subjects' performance in response to the owner- and experimenter-given auditory cues were interrelated, but unrelated to responses to pointing gestures, suggesting that dogs' ability to understand the referential nature of auditory cues and visual gestures partly arise from different socio-cognitive skills.

Sleep-physiological correlates of brachycephaly in dogs.

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.

Event-related potentials indicate differential neural reactivity to species and valence information in vocal stimuli in sleeping dogs.

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.

Dog brains are sensitive to infant- and dog-directed prosody.

When addressing preverbal infants and family dogs, people tend to use specific speech styles. While recent studies suggest acoustic parallels between infant- and dog-directed speech, it is unclear whether dogs, like infants, show enhanced neural sensitivity to prosodic aspects of speech directed to them. Using functional magnetic resonance imaging on awake unrestrained dogs we identify two non-primary auditory regions, one that involve the ventralmost part of the left caudal Sylvian gyrus and the temporal pole and the other at the transition of the left caudal and rostral Sylvian gyrus, which respond more to naturalistic dog- and/or infant-directed speech than to adult-directed speech, especially when speak by female speakers. This activity increase is driven by sensitivity to fundamental frequency mean and variance resulting in positive modulatory effects of these acoustic parameters in both aforementioned non-primary auditory regions. These findings show that the dog auditory cortex, similarly to that of human infants, is sensitive to the acoustic properties of speech directed to non-speaking partners. This increased neuronal responsiveness to exaggerated prosody may be one reason why dogs outperform other animals when processing speech.

Sleep and cognition in aging dogs. A polysomnographic study.

Introduction: Sleep is fundamental for cognitive homeostasis, especially in senior populations since clearance of amyloid beta (key in the pathophysiology of Alzheimer's disease) occurs during sleep. Some electroencephalographic characteristics of sleep and wakefulness have been considered a hallmark of dementia. Owners of dogs with canine cognitive dysfunction syndrome (a canine analog to Alzheimer's disease) report that their dogs suffer from difficulty sleeping. The aim of this study was to quantify age-related changes in the sleep-wakefulness cycle macrostructure and electroencephalographic features in senior dogs and to correlate them with their cognitive performance. Methods: We performed polysomnographic recordings in 28 senior dogs during a 2 h afternoon nap. Percentage of time spent in wakefulness, drowsiness, NREM, and REM sleep, as well as latency to the three sleep states were calculated. Spectral power, coherence, and Lempel Ziv Complexity of the brain oscillations were estimated. Finally, cognitive performance was evaluated by means of the Canine Dementia Scale Questionnaire and a battery of cognitive tests. Correlations between age, cognitive performance and sleep-wakefulness cycle macrostructure and electroencephalographic features were calculated. Results: Dogs with higher dementia scores and with worse performance in a problem-solving task spent less time in NREM and REM sleep. Additionally, quantitative electroencephalographic analyses showed differences in dogs associated with age or cognitive performance, some of them reflecting shallower sleep in more affected dogs. Discussion: Polysomnographic recordings in dogs can detect sleep-wakefulness cycle changes associated with dementia. Further studies should evaluate polysomnography's potential clinical use to monitor the progression of canine cognitive dysfunction syndrome.

Six facial prosodic expressions caregivers similarly display to infants and dogs.

Parents tend to use a specific communication style, including specific facial expressions, when speaking to their preverbal infants which has important implications for children's healthy development. In the present study, we investigated these facial prosodic features of caregivers with a novel method that compares infant-, dog- and adult-directed communication. We identified three novel facial displays in addition to the already described three facial expressions (i.e. the 'prosodic faces') that mothers and fathers are typically displaying when interacting with their 1-18 month-old infants and family dogs, but not when interacting with another adult. The so-called Special Happy expression proved to be the most frequent face type during infant- and dog-directed communication which always includes a Duchenne marker to convey an honest and intense happy emotion of the speaker. These results suggest that the 'prosodic faces' play an important role in both adult-infant and human-dog interactions and fulfil specific functions: to call and maintain the partner's attention, to foster emotionally positive interactions, and to strengthen social bonds. Our study highlights the relevance of future comparative studies on facial prosody and its potential contribution to healthy emotional and cognitive development of infants.

Stereoselective Synthesis and Antiproliferative Activities of Tetrafunctional Diterpene Steviol Derivatives.

A new family of diterpene-type aminotriol derivatives has been synthesised from stevioside in a stereoselective manner. The key intermediate spiro-epoxide was prepared through the methyl ester of the allilyc diol derived from steviol. The oxirane ring was opened with primary and secondary amines, providing a versatile library of aminotriols. The corresponding primary aminotriol was formed by palladium-catalysed hydrogenation, and an N,O-heterocyclic compound was synthesised in a regioselective reaction. All new compounds were characterised by 1D- and 2D-NMR techniques and HRMS measurements. In our in vitro investigations, we found that the aromatic N-substituted derivatives exhibited high inhibition of cell growth on human cancer cell lines (HeLa, SiHa, A2780, MCF-7 and MDA-MB-231). The antiproliferative activities were assayed by the MTT method. Furthermore, the introduction of an additional hydroxy group slightly increased the biological activity. The drug-likeness of the compounds was assessed by in silico and experimental physicochemical characterisations, completed by kinetic aqueous solubility and in vitro intestinal-specific parallel artificial membrane permeability assay (PAMPA-GI) measurements.

Owner-rated hyperactivity/impulsivity is associated with sleep efficiency in family dogs: a non-invasive EEG study.

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.

Attachment towards the Owner Is Associated with Spontaneous Sleep EEG Parameters in Family Dogs.

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 their relevant behavior in a task-free social context assessing their attachment towards the owner. In random order, each dog participated in a non-invasive sleep electroencephalogram (EEG) measurement and in the Strange Situation Test (SST) to assess their attachment behavior. We found that higher attachment scores were associated with more time spent in NREM sleep, lower NREM alpha power activity and lower NREM alpha-delta anticorrelation. Our results reveal that, when dogs sleep in a novel environment in the company of their owners, differences in their attachment are reflected in their sleep EEG characteristics. This could be best explained by the different degree that owners could be used as a safe haven in an unfamiliar environment and during the unusual procedure of the first EEG measurement.

Differences in dogs' event-related potentials in response to human and dog vocal stimuli; a non-invasive study.

Recent advances in the field of canine neuro-cognition allow for the non-invasive research of brain mechanisms in family dogs. Considering the striking similarities between dog's and human (infant)'s socio-cognition at the behavioural level, both similarities and differences in neural background can be of particular relevance. The current study investigates brain responses of n = 17 family dogs to human and conspecific emotional vocalizations using a fully non-invasive event-related potential (ERP) paradigm. We found that similarly to humans, dogs show a differential ERP response depending on the species of the caller, demonstrated by a more positive ERP response to human vocalizations compared to dog vocalizations in a time window between 250 and 650 ms after stimulus onset. A later time window between 800 and 900 ms also revealed a valence-sensitive ERP response in interaction with the species of the caller. Our results are, to our knowledge, the first ERP evidence to show the species sensitivity of vocal neural processing in dogs along with indications of valence sensitive processes in later post-stimulus time periods.

Social Stimulation by the Owner Increases Dogs' (Canis familiaris) Social Susceptibility in a Food Choice Task-The Possible Effect of Endogenous Oxytocin Release.

Recent evidence suggests a human-like susceptibility to social influence in dogs. For example, dogs tend to ignore their 'natural' preference for the larger amount of food after having seen a human's explicit preference for a smaller quantity. However, it is still unclear whether this tendency to conform to the partner's behaviour can be influenced by social stimuli and/or the neurohormone oxytocin as primers to prosocial predispositions. In Experiment I, eighty two dogs were tested using Prato-Previde et al.'s food quantity preference task. In Experiment I, we investigated in a 2 × 2 design how (i) a 10-minute-long social stimulation by the owner versus a socially ignoring pre-treatment as well as (ii) on-line ostensive communications versus no communication during task demonstration affect dogs' (N = 82) choices in the abovementioned food choice task. Results indicate that the owners' pre-treatment with social stimuli (eye contact, petting) increased dogs' susceptibility to the experimenter's food preference, but the salient ostensive addressing signals accompanying human demonstration masked this social priming effect. In Experiment II, N = 32 dogs from the subjects of Experiment I were retested after oxytocin (OT) or placebo (PL) pre-treatments. This experiment aimed to study whether intranasal administration of oxytocin as compared to placebo treatment would similarly increase dogs' tendency to re-enact the human demonstrator's counterproductive choice in the same task. Results showed an increased susceptibility to the human preference in the OT group, suggesting that both socially stimulating pre-treatment and the intranasal administration of oxytocin have similar priming effects on dogs' social susceptibility.

Interactions between Sleep and Emotions in Humans and Animal Models.

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.

Developmental features of sleep electrophysiology in family dogs.

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.

Interhemispheric asymmetry during NREM sleep in the dog.

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.

The implicit reward value of the owner's face for dogs.

It is increasingly assumed that domestication has equipped dogs with unique socio-cognitive skills, which raises the possibility of intriguing parallels between the social motivational systems of the two species. However, the positive incentive value of human facial stimuli is a largely unexplored area. Here, we investigated whether the owner's face serves as a social reinforcer. In a two-way choice task N = 39 dogs were presented with a short video about their owners' head showing the face (facing owner [FO]) vs. the back of the head (non-facing owner). Despite both locations containing equal food reward, dogs approached the container associated with FO more frequently (p < .001), and this was not affected by side, trial order, and choice latency. However, the considerable inter-individual differences in dogs' task performance suggest that the added social component required special social skills which need to be further explored.

Longitudinal Volumetric Assessment of Ventricular Enlargement in Pet Dogs Trained for Functional Magnetic Resonance Imaging (fMRI) Studies.

BACKGROUND: Recent studies suggest that clinically sound ventriculomegaly in dogs could be a preliminary form of the clinically significant hydrocephalus. We evaluated changes of ventricular volumes in awake functional magnetic resonance imaging (fMRI) trained dogs with indirectly assessed cognitive abilities over time (thus avoiding the use of anaesthetics, which can alter the pressure). Our research question was whether ventricular enlargement developing over time would have any detrimental effect on staying still while being scanned; which can be extrapolated to the ability to pay attention and to exert inhibition. METHODS: Seven healthy dogs, 2-8 years old at the baseline scan and 4 years older at rescan, participated in a rigorous and gradual training for staying motionless (<2 mm) in the magnetic resonance (MR) scanner without any sedation during 6 minute-long structural MR sequences. On T1 structural images, volumetric analyses of the lateral ventricles were completed by software guided semi-automated tissue-type segmentations performed with FMRIB Software Library (FSL, Analysis Group, Oxford, UK). RESULTS AND CONCLUSION: We report significant enlargement for both ventricles (left: 47.46 %, right: 46.07 %) over time while dogs retained high levels of attention and inhibition. The results suggest that even considerable ventricular enlargement arising during normal aging does not necessarily reflect observable pathological changes in behavior.

Behavioral and Neurophysiological Correlates of Dogs' Individual Sensitivities to Being Observed by Their Owners While Performing a Repetitive Fetching Task.

Ample evidence suggests that dogs possess enhanced skills in reading human visual attention, but it remains to be explored whether they are sensitive to the audience effect in their interactions with humans. The present study aimed to investigate how dogs' behavior is affected by their owners' visual attention while performing a repetitive task (bringing an object back to an unfamiliar experimenter while the owner waited passively). We assumed that if dogs are susceptible to the audience effect, their task persistence and task performance would vary according to their owners' attentiveness. A group of adult pet dogs (N = 27) were repeatedly presented with an object retrieval task by the experimenter (N = 20 trials) while owners either ignored their dogs (Inattentive Owner condition) or paid attention to their dogs' actions (Attentive Owner condition). Behavioral observations were complemented with the owner's reports of their relationships with their dogs (assessed by means of an owner-pet attachment questionnaire) and dogs' spectral EEG sleep profile (recorded during 3-h-long daytime sleep). Although dogs, independently of their owners' attentional state, were generally willing to comply with the fetching task, they were faster to approach the toy object and gazed significantly longer at their owners when he/she was paying attention. This finding is reminiscent of peer influence observed in humans. Further, characteristics of relationship insecurity (relationship anxiety and avoidance) were associated with dogs' task persistence and performance. Dogs of owners with higher relationship anxiety tended to approach the toy object less frequently, and dogs of owners with higher relationship avoidance and anxiety were more hesitant to approach the toy object. We also found that dogs' individual susceptibilities to the audience effect is related to EEG spectral power of both REM and non-REM sleep as well as in pre-sleep (drowsiness) in a trait-like manner. These results, in line with previous findings, support the notion that dogs have a somewhat human-like susceptibility to the audience effect, a trait which might be linked to more complex mechanisms, such as self-presentation or reputation management, helping the two species to become effective social partners.