Breed group differences in the unsolvable problem task: herding dogs prefer their owner, while solitary hunting dogs seek stranger proximity.

The communicating skills of dogs are well documented and especially their contact-seeking behaviours towards humans. The aim of this study was to use the unsolvable problem paradigm to investigate differences between breed groups in their contact-seeking behaviours towards their owner and a stranger. Twenty-four dogs of ancient breeds, 58 herding dogs, and 17 solitary hunting dogs were included in the study, and their behaviour when presented with an unsolvable problem task (UPT) was recorded for 3 min. All breed groups interacted with the test apparatus the same amount of time, but the herding dogs showed a longer gaze duration towards their owner compared to the other groups and they also preferred to interact with their owner instead of a stranger. Interestingly, the solitary hunting dogs were more in stranger proximity than the other groups, and they also showed a preference to make contact with a stranger instead of their owner. Hence, we found differences in contact-seeking behaviours, reflecting the dog-human relationship, between breed groups that might not only be related to their genetic similarity to wolves, but also due to the specific breeding history of the dogs.

Long-term stress in dogs is related to the human-dog relationship and personality traits.

Previously, we found that dogs belonging to the herding breed group, selected for human cooperation, synchronise their long-term stress levels with their owners. The aim of the current study was to investigate features that could influence long-term stress levels in ancient dog breeds, genetically closer to wolves, and dogs specifically selected to work independently of their owner. Twenty-four ancient breed dogs and 18 solitary hunting dogs were recruited and hair samples were obtained from both dogs and owners from which hair cortisol concentration (HCC) was analysed. Additionally, the owners completed lifestyle surveys, the Monash Dog Owner Relationship Scale (MDORS) on human-dog relationship, and both dog and owner personality questionnaires (Dog Personality questionnaire and Big Five Inventory survey). The results from the MDORS indicate that the subscale Perceived cost correlated to the dog HCC of tested breed groups: solitary hunting breeds (χ2 = 4.95, P = 0.026, ß = 0.055), ancient breeds (χ2 = 2.74, P = 0.098, ß = 0.027), and herding dogs included from a previous study (χ2 = 6.82, P = 0.009, ß = - 0.061). The HCC of the solitary hunting dogs was also related to the owner personality traits Agreeableness (χ2 = 12.30, P < 0.001, ß = - 0.060) and Openness (χ2 = 9.56, P = 0.002, ß = 0.048) suggesting a more substantial influence of the owner on the solitary hunting dog's HCC compared to the ancient breeds. No effect of owner HCC on dog HCC was found in either ancient or in solitary hunting breeds. Hence, the long-term stress synchronisation is likely to be a trait in breeds selected for human cooperation. In conclusion, dog HCC is often related to the owners' personality, but is primarily influenced by the owner-dog relationship.

Author Correction: Long-term stress levels are synchronized in dogs and their owners.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Effects of size and personality on social learning and human-directed behaviour in horses (Equus caballus).

Due to our long history of living in close association with horses, these animals are suggested to have enhanced skills in understanding and communicating with humans. Today, horses have become important to humans for sport and leisure and their understanding of human behaviour and their human-directed behaviour are therefore of great importance. In this study, we investigated 22 horses in a human contact-seeking experiment where they were presented with an unsolvable problem and a detour experiment with a human demonstrator. The unsolvable problem consisted of pieces of carrot in a closed bucket and the detour resembled the shape of V. Additionally, personality traits of the participating horses were assessed. Interestingly, the full-sized horses (N = 11) showed more human-related behaviours when presented with an unsolvable problem compared to before the carrots were made unreachable (p = 0.033), while the ponies (N = 11) did not. However, neither the full-sized horses nor the ponies were significantly more successful in the detour after human demonstrations than in control trials. When comparing the two experiments, we found the task-oriented behaviour in the detour test to positively correlate with human proximity and eye contact-seeking behaviour towards humans during the unsolvable problem in the contact-seeking test. Interestingly, again this was only true for the full-sized horses (p < 0.05) and not for the ponies. From the horse personality questionnaire results, the traits excitability and anxiousness revealed strong negative correlations with human-directed behaviour in the contact-seeking experiment (p < 0.05). Hence, size (full-sized horse/pony) and personality influenced the human-related behaviours of the horses and we suggest a future focus on these aspects to deepen our understanding of human-horse communication.

Long-term stress levels are synchronized in dogs and their owners.

This study reveals, for the first time, an interspecific synchronization in long-term stress levels. Previously, acute stress, has been shown to be highly contagious both among humans and between individuals of other species. Here, long-term stress synchronization in dogs and their owners was investigated. We studied 58 dog-human dyads and analyzed their hair cortisol concentrations (HCC) at two separate occasions, reflecting levels during previous summer and winter months. The personality traits of both dogs and their owners were determined through owner-completed Dog Personality Questionnaire (DPQ) and human Big Five Inventory (BFI) surveys. In addition, the dogs' activity levels were continuously monitored with a remote cloud-based activity collar for one week. Shetland sheepdogs (N = 33) and border collies (N = 25), balanced for sex, participated, and both pet dogs and actively competing dogs (agility and obedience) were included to represent different lifestyles. The results showed significant interspecies correlations in long-term stress where human HCC from both summer and winter samplings correlated strongly with dog HCC (summer: N = 57, χ2 = 23.697, P < 0.001, ß = 0.235; winter: N = 55, χ2 = 13.796, P < 0.001, ß = 0.027). Interestingly, the dogs' activity levels did not affect HCC, nor did the amount of training sessions per week, showing that the HCC levels were not related to general physical activity. Additionally, there was a seasonal effect in HCC. However, although dogs' personalities had little effects on their HCC, the human personality traits neuroticism, conscientiousness, and openness significantly affected dog HCC. Hence, we suggest that dogs, to a great extent, mirror the stress level of their owners.

High visual acuity revealed in dogs.

Humans have selectively bred and used dogs over a period of thousands of years, and more recently the dog has become an important model animal for studies in ethology, cognition and genetics. These broad interests warrant careful descriptions of the senses of dogs. Still there is little known about dog vision, especially what dogs can discriminate in different light conditions. We trained and tested whippets, pugs, and a Shetland sheepdog in a two-choice discrimination set-up and show that dogs can discriminate patterns with spatial frequencies between 5.5 and 19.5 cycle per degree (cpd) in the bright light condition (43 cd m-2). This is a higher spatial resolution than has been previously reported although the individual variation in our tests was large. Humans tested in the same set-up reached acuities corresponding to earlier studies, ranging between 32.1 and 44.2 cpd. In the dim light condition (0.0087 cd m-2) the acuity of dogs ranged between 1.8 and 3.5 cpd while in humans, between 5.9 and 9.9 cpd. Thus, humans make visual discrimination of objects from roughly a threefold distance compared to dogs in both bright and dim light.

Intranasal oxytocin and a polymorphism in the oxytocin receptor gene are associated with human-directed social behavior in golden retriever dogs.

The oxytocin system may play an important role in dog domestication from the wolf. Dogs have evolved unique human analogue social skills enabling them to communicate and cooperate efficiently with people. Genomic differences in the region surrounding the oxytocin receptor (OXTR) gene have previously been associated with variation in dogs' communicative skills. Here we have utilized the unsolvable problem paradigm to investigate the effects of oxytocin and OXTR polymorphisms on human-directed contact seeking behavior in 60 golden retriever dogs. Human-oriented behavior was quantified employing a previously defined unsolvable problem paradigm. Behaviors were tested twice in a repeated, counterbalanced design, where dogs received a nasal dose of either oxytocin or saline 45min before each test occasion. Buccal DNA was analysed for genotype on three previously identified SNP-markers associated with OXTR. The same polymorphisms were also genotyped in 21 wolf blood samples to explore potential genomic differences between the species. Results showed that oxytocin treatment decreased physical contact seeking with the experimenter and one of the three polymorphisms was associated with degree of physical contact seeking with the owner. Dogs with the AA-genotype at this locus increased owner physical contact seeking in response to oxytocin while the opposite effect was found in GG-genotype individuals. Hence, intranasal oxytocin treatment, an OXTR polymorphism and their interaction are associated with dogs' human-directed social skills, which can explain previously described breed differences in oxytocin response. Genotypic variation at the studied locus was also found in wolves indicating that it was present even at the start of dog domestication.

Genomic Regions Associated With Interspecies Communication in Dogs Contain Genes Related to Human Social Disorders.

Unlike their wolf ancestors, dogs have unique social skills for communicating and cooperating with humans. Previously, significant heritabilities for human-directed social behaviors have been found in laboratory beagles. Here, a Genome-Wide Association Study identified two genomic regions associated with dog's human-directed social behaviors. We recorded the propensity of laboratory beagles, bred, kept and handled under standardized conditions, to initiate physical interactions with a human during an unsolvable problem-task, and 190 individuals were genotyped with an HD Canine SNP-chip. One genetic marker on chromosome 26 within the SEZ6L gene was significantly associated with time spent close to, and in physical contact with, the human. Two suggestive markers on chromosome 26, located within the ARVCF gene, were also associated with human contact seeking. Strikingly, four additional genes present in the same linkage blocks affect social abilities in humans, e.g., SEZ6L has been associated with autism and COMT affects aggression in adolescents with ADHD. This is, to our knowledge, the first genome-wide study presenting candidate genomic regions for dog sociability and inter-species communication. These results advance our understanding of dog domestication and raise the use of the dog as a novel model system for human social disorders.

Hair cortisol varies with season and lifestyle and relates to human interactions in German shepherd dogs.

It is challenging to measure long-term endocrine stress responses in animals. We investigated whether cortisol extracted from dog hair reflected the levels of activity and stress long-term, during weeks and months. Hair samples from in total 59 German shepherds were analysed. Samples for measuring cortisol concentrations were collected at three occasions and we complemented the data with individual scores from the Canine Behavioural Assessment and Research Questionnaire (C-BARQ). Generalised linear mixed model (GLMM) results showed that hair cortisol varied with season and lifestyle: competition dogs had higher levels than companion, and professional working dogs, and levels were higher in January than in May and September. In addition, a positive correlation was found between the cortisol levels and the C-BARQ score for stranger-directed aggression (r = 0.31, P = 0.036). Interestingly, the factor "playing often with the dog" (r = -0.34, P = 0.019) and "reward with a treat/toy when the dog behaves correctly" (r = -0.37, P = 0.010) correlated negatively with cortisol levels, suggesting that positive human interactions reduce stress. In conclusion, hair cortisol is a promising method for revealing the activity of the HPA-axis over a longer period of time, and human interactions influence the cortisol level in dogs.

The impact of domestication on the chicken optical apparatus.

Domestication processes tend to release animals from natural selection and favour traits desired by humans, such as food-production and co-operative behaviour. A side effect of such selective breeding is the alteration of unintended traits. In this paper, we investigate how active selection for egg production in chickens has affected the visual system, in particular the optical sensitivity that relates to the ability of chickens to see in dim light. We measured eye dimensions as well as the pupil diameter at different light intensities (the steady state pupil dynamics), in adult male and female White Leghorns and the closest relatives to their ancestor, the Red Junglefowls. With this information, we calculated the focal length and optical sensitivity (f-number) of the eyes. Males have larger eyes than females in both breeds and White Leghorn eyes are larger than those of Red Junglefowls in both sexes. The steady state pupil dynamics is less variable, however, the combination of pupil dynamics and eye size gives a higher optical sensitivity in Red Junglefowl eyes than in White Leghorns at light intensities below approximately 10 cd/m(2). While eye size and focal length match the larger body size in White Leghorns compared to Red Junglefowls, the steady state pupil dynamics do not. The reason for this is likely to be that eye morphology and the neuro-muscular control of the pupil have been affected differently by the strong selection for egg production and the simultaneous release of the selection pressure for high performing vision. This study is the first description of how optical sensitivity has changed in a domesticated species and our results demonstrate important considerations regarding domestication processes and sensory ability.

The pupils and optical systems of gecko eyes.

The nocturnal helmet gecko, Tarentola chazaliae, discriminates colors in dim moonlight when humans are color blind. The sensitivity of the helmet gecko eye has been calculated to be 350 times higher than human cone vision at the color vision threshold. The optics and the large cones of the gecko are important reasons why they can use color vision at low light intensities. Using photorefractometry and an adapted laboratory Hartmann-Shack wavefront sensor of high resolution, we also show that the optical system of the helmet gecko has distinct concentric zones of different refractive powers, a so-called multifocal optical system. The intraspecific variation is large but in most of the individuals studied the zones differed by 15 diopters. This is of the same magnitude as needed to focus light of the wavelength range to which gecko photoreceptors are most sensitive. We compare the optical system of the helmet gecko to that of the diurnal day gecko, Phelsuma madagascariensis grandis. The optical system of the day gecko shows no signs of distinct concentric zones and is thereby monofocal.

The absolute threshold of colour vision in the horse.

Arrhythmic mammals are active both during day and night if they are allowed. The arrhythmic horses are in possession of one of the largest terrestrial animal eyes and the purpose of this study is to reveal whether their eye is sensitive enough to see colours at night. During the day horses are known to have dichromatic colour vision. To disclose whether they can discriminate colours in dim light a behavioural dual choice experiment was performed. We started the training and testing at daylight intensities and the horses continued to choose correctly at a high frequency down to light intensities corresponding to moonlight. One Shetland pony mare, was able to discriminate colours at 0.08 cd/m(2), while a half blood gelding, still discriminated colours at 0.02 cd/m(2). For comparison, the colour vision limit for several human subjects tested in the very same experiment was also 0.02 cd/m(2). Hence, the threshold of colour vision for the horse that performed best was similar to that of the humans. The behavioural results are in line with calculations of the sensitivity of cone vision where the horse eye and human eye again are similar. The advantage of the large eye of the horse lies not in colour vision at night, but probably instead in achromatic tasks where presumably signal summation enhances sensitivity.

Colour perception in a dichromat.

Most mammals have dichromatic colour vision based on two different types of cones: a short-wavelength-sensitive cone and a long-wavelength-sensitive cone. Comparing the signal from two cone types gives rise to a one-dimensional chromatic space when brightness is excluded. The so-called ;neutral point' refers to the wavelength that the animal cannot distinguish from achromatic light such as white or grey because it stimulates both cone types equally. The question is: how do dichromats perceive their chromatic space? Do they experience a continuous scale of colours or does the neutral point divide their chromatic space into two colour categories, i.e. into colours of either short or long wavelengths? We trained horses to different colour combinations in a two-choice behavioural experiment and tested their responses to the training and test colours. The horses chose colours according to their similarity/relationship to rewarded and unrewarded training colours. There was no evidence for a categorical boundary at the neutral point or elsewhere. This study suggests that dichromats perceive their chromatic space as a continuous scale of colours, treating the colour at the neutral point as any other colour they can distinguish.

Nocturnal colour vision--not as rare as we might think.

The dual retina of humans and most vertebrates consists of multiple types of cone for colour vision in bright light and one single type of rod, leaving these animals colour-blind at night. Instead of comparing the signals from different spectral types of photoreceptors, they use one highly sensitive receptor, thus improving the signal-to-noise ratio. However, nocturnal moths and geckos can discriminate colours at extremely dim light intensities when humans are colour-blind, by sacrificing spatial and temporal rather than spectral resolution. The advantages of colour vision are just as obvious at night as they are during the day. Colour vision is much more reliable than achromatic contrast, not only under changing light intensities, but also under the colour changes occurring during dusk and dawn. It can be expected that nocturnal animals other than moths and geckos make use of the highly reliable colour signals in dim light.

Nocturnal colour vision in geckos.

Nocturnal animals are said to sacrifice colour vision in favour of increased absolute sensitivity. This is true for most vertebrates that possess a dual retina with a single type of rod for colour-blind night vision and multiple types of cone for diurnal colour vision. However, among the nocturnal vertebrates, geckos are unusual because they have no rods but three cone types. Here, we show that geckos use their cones for colour vision in dim light. Two specimens of the nocturnal helmet gecko Tarentola (formerly Geckonia) chazaliae were able to discriminate blue from grey patterns by colour alone. Experiments were performed at 0.002 cd m(-2), a light intensity similar to dim moonlight. We conclude that nocturnal geckos can use cone-based colour vision at very dim light levels when humans rely on colour-blind rod vision.