The intestinal tract affects the brain through metabolites produced by gut-inhabiting bacteria. I... more The intestinal tract affects the brain through metabolites produced by gut-inhabiting bacteria. In this study, we show that the number of errors the dogs commit in a short-term memory test and also their age is linked to the gut microbiome composition. The proportion of Fusobacteria is lower in older animals. Dogs with better memory performance (i.e., fewer mistakes) have relatively fewer Actinobacteria in their fecal samples collected right after the behavior test. This result is in agreement with the high abundance of some Actinobacteria in the gastrointestinal tract of persons living with Alzheimer's disease. Links between memory performance and gut microbiota have been reported on rodents but not on dogs before. The research opens up new venues in canine aging and neurodevelopmental research.
Background and aims: Dogs have recently become an important model species for comparative social ... more Background and aims: Dogs have recently become an important model species for comparative social and cognitive neuroscience. Brain template-related label maps are essential for functional magnetic resonance imaging (fMRI) data analysis, to localize neural responses. In this study, we present a detailed, individual-based, T1-weighted MRI-based brain label map used in dog neuroimaging analysis. Methods: A typical, medium-headed dog (a 7.5-year-old male Golden Retriever) was selected from a cohort of 22 dogs, based on brain morphology (shape, size, and gyral pattern), to serve as the template for a label map. Results: Eighty-six 3-dimensional labels were created to highlight the main cortical (cerebral gyri on the lateral and medial side) and subcortical (thalamus, caudate nucleus, amygdala, and hippocampus) structures of the prosencephalon and diencephalon, and further main parts of brainstem (mesencephalon and rhombencephalon). Discussion: Importantly, this label map is (a) considerably more detailed than any available dog brain template; (b) it is easy to use with freeware and commercial neuroimaging software for MRI and fMRI analysis; and (c) it can be registered to other existing templates, including a recent average-based dog brain template. Using the coordinate system and label map proposed here can enhance precision and standard localization during future canine neuroimaging studies.
In humans, age-related changes in personality occur in a non-random fashion with respect to their... more 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.
In dogs, the social and spatial restriction associated with living in a kennel environment could ... more In dogs, the social and spatial restriction associated with living in a kennel environment could lead to chronic stress and the development of abnormal behaviors ("kennel-dog syndrome"). However, little is known about how kenneled dogs differ from their conspecifics living as pets in human families. In the current study, using a test battery exposing the dogs to novel stimuli, we compared the behavior of three groups of beagles: (1) kenneled dogs living in a restricted environment with limited human contact (N = 78), (2) family dogs living in human families as pets (N = 37), and (3) adopted dogs born in the kenneled population but raised in human families (N = 13). We found one factor comprising most of the test behaviors, labeled as Responsiveness. Family dogs and adopted dogs scored higher in Responsiveness than kenneled dogs. However, 23% of the kenneled dogs were comparable to family and adopted dogs based on a cluster analysis, indicating a similar (positive) reaction to novel stimuli, while 77% of the kenneled dogs were unresponsive (mostly immobile) in at least part of the test. To assess if the behavioral difference between the family and kenneled dogs could be due to genetic divergence of these two populations and/or to lower genetic diversity of the kenneled dogs, we analyzed their genetic structure using 11 microsatellite markers. We found no significant difference between the populations in their genetic diversity (i.e., heterozygosity, level of inbreeding), nor any evidence that the family and kenneled populations originated from different genetic pools. Thus, the behavior difference between the groups more likely reflects a G × E interaction, that is, the influence of specific genetic variants manifesting under specific environmental conditions (kennel life). Nevertheless, some kenneled individuals were (genetically) more resistant to social and environmental deprivation. Selecting for such animals could strongly improve the welfare of kenneled dog populations. Moreover, exploring the genetic background of their higher resilience could also help to better understand the genetics behind stress-and fear-related behaviors.
Introduction: The aim of this study was to explore spontaneous social interactions between dyads ... more Introduction: The aim of this study was to explore spontaneous social interactions between dyads of unfamiliar adult dogs. Although intraspecific encounters are frequent events in the life of pet dogs, the factors that might influence encounters, such as sex, dyad composition, reproductive status, age, and state of cohabitation (keeping the dogs singly or in groups), remained unexplored. Methods: In this study, we assigned unfamiliar, non-aggressive dogs to three types of dyads defined by sex and size. We observed their unrestrained, spontaneous behaviors in an unfamiliar dog park, where only the two dogs, the owners, and experimenter were present. Results: We found that the dogs, on average, spent only 17% of the time (less than 1 min) in proximity. Sex, dyad composition, reproductive status, and age influenced different aspects of the interactions in dyads. Female dogs were more likely to initiate the first contact in their dyad but later approached the partner less frequently, were less likely to move apart, and displayed less scent marking. Following and moving apart were more frequent in male-male interactions. Neutered dogs spent more time following the other dog and sniffed other dogs more frequently. The time companion dogs spent in proximity and number of approaches decreased with age. Conclusion: The study provides guidance for dog owners about the outcomes of intraspecific encounters based on the dog's age, sex, and reproductive status, as well as the sex of the interacting partner.
Dominance is well defined in ethology, debated in psychology, and is often unclear among the dog ... more Dominance is well defined in ethology, debated in psychology, and is often unclear among the dog owning public and in the press. However, to date, no study has examined how owners perceive dominance in dogs, and what different behaviours and personality types are used to describe dominant and subordinate individuals. A questionnaire study was launched to investigate the external validity of owner-derived estimates of dominance in dog dyads sharing the same household (N = 1,151). According to the owners, dogs rated as dominant (87%) have priority access to resources (resting place, food, and rewards), undertake certain tasks (defend and lead the group, bark more), display dominance (win fights, lick the other's mouth less, and mark over the other's urine), share certain personality traits (smarter, more aggressive and impulsive), and are older than their partner dog (all p < 0.0001). An age-related hypothesis has been suggested to explain dominance in dogs; but we found that dog age did not explain the occurrence of dominance related behaviours over the owners' estimate of dominance status. Results suggest that owner-derived reports of dominance ranks of dogs living in multi-dog households correspond to ethologically valid behavioural markers of dominance. Size and physical condition were unrelated to the perceived dominance. Surprisingly, in mixed-sex dyads, females were more frequently rated as dominant than males, which might correspond to a higher proportion of neutered females in this subgroup. For future studies that wish to allocate dominance status using owner report, we offer a novel survey.
Most common methods that directly show macro-or microscopic anatomy of the brain usually require ... more Most common methods that directly show macro-or microscopic anatomy of the brain usually require the removal of the organ from the neurocranium. However, the brain can be revealed in situ by using proper sectioning techniques. Our aim was to both improve the cryosectioning method, test its limits and create a high-resolution macro-anatomical image series of a Beagle brain, which at the time of the study did not exist. A two-year-old female Beagle has been scanned with CT and MRI ante and post mortem, then the arteries of the head were filled with red resin. After freezing to -80˚C, a neurocranium block was created and was embedded into a water-gelatin mix. Using a special milling device and a DSLR camera, 1112 consecutive RGB-color cryosections were made with a 100 μm layer thickness and captured in high resolution (300 dpi, 24-bit color, and pixel size was 19.5 x 19.5 μm). Image post-processing was done with Adobe Photoshop CS3 and Thermo Scientific Amira 6.0 softwares, and as a result of the proper alignment and coregistration, visualization and comparing was possible with all the applied imaging modalities (CT, MRI, cryosectioning) in any arbitrary plane. Surface models from the arteries, veins, brain and skull were also generated after segmentation in the same coordinate system, giving a unique opportunity for comparing the two-dimensional and three-dimensional anatomy. This is the first study which focuses directly to this high-definition multimodal visualization of the canine brain, and it provides the most accurate results compared to previous cryosectioning studies, as using an improved method, higher image quality, more detailed image, proper color fidelity and lower artefact formation were achieved. Based on the methodology we described, it can serve as a base for future multimodal (CT, MR, augmented-or virtual reality) imaging atlases for medical, educational and scientific purposes.
Dogs living on Bali Island have been free-ranging for thousands of years. A large group of expatr... more Dogs living on Bali Island have been free-ranging for thousands of years. A large group of expatriates sometimes adopt Bali dogs and keep them restricted to their houses and backyards, as is typical in modern western cultures. This provides us with the unique opportunity to compare the personality traits of dogs to their lifestyle either living as human companions or as free-ranging animals, exploring at the same time the impact of demographic variables (such as age, sex, and neutered status) on personality. After controlling for internal consistency of the scales and between-observer variation, we found that free-ranging Bali dogs were rated as less active, less excitable, less aggressive towards animals, and less inclined to chase animals or humans than Bali dogs living as human companions. Among free-ranging dogs, females were found to be more excitable. Females in the whole sample were also more fearful of people. The results of this preliminary study suggest that a change in lifestyle, i.e. being adopted, and living in a confined environment has negative consequences on some canine personality traits, such as activity/excitability, aggression towards animals, and prey drive.
Visual and auditory impairments can have a large impact on performance in cognitive tests. It is ... more Visual and auditory impairments can have a large impact on performance in cognitive tests. It is important to evaluate the sensory capacities of dogs before enrolling them in cognitive tests, in order to exclude sensory impairment as confounding effect. Therefore we designed multiple non-invasive testing paradigms to detect subjects with potential auditory and visual impairment, without requiring extensive training for the dog. Multiple testing was a means to add internal reliability, and to reduce the risk of false positives due to habituation and previous learning or false negatives due to random errors. Our sensory test battery consisted of four subtests: (1) 'Clapping' auditory test, (2) 'Recorded sound' auditory test, (3) 'Distance' visual test, and (4) 'Darkness' visual test. The 'Clapping test' was similar to the clapping test used by veterinarians, with the addition that the clapping was performed at various distances from the dog. In the 'Recorded sound test', the dogs' reaction to various sounds played back at different volumes. In the 'Distance test' we placed a small piece of food on one of four plates placed on the grounds at varying distances from the dogs. In the 'Darkness test', we measured the dogs' performance in walking through an S-shaped route during artificial dusk and daylight-like conditions.
Aging is the largest risk factor in many diseases and mortality alike. As the elderly population ... more Aging is the largest risk factor in many diseases and mortality alike. As the elderly population is expected to increase at an accelerating rate in the future, these phenomena will pose a growing socio-economic burden on societies. To successfully cope with this challenge, a deeper understanding of aging is crucial. In many aspects, the companion dog is an increasingly popular model organism to study aging, with the promise of producing results that are more applicable to humans than the findings that come from the studies of classical model organisms. In this preliminary study we used the whole-genome sequence of two extremely old dogs -age: 22 and 27 years (or 90-135% more, than the average lifespan of dogs) -in order to make the first steps to understand the genetic background of extreme longevity in dogs. We identified more than ∼80 1000 novel SNPs in the two dogs (7500 of which overlapped between them) when compared to three publicly available canine SNP databases, which included SNP information from850 dogs. Most novel mutations (∼52000 SNPs) were identified at non-coding regions, while 4.6% of the remaining SNPs (n∼1600) were at exons, including 670 missense variants -76 of which overlapped between the two animalsacross 472 genes. Based on their gene ontologies, these genes were related -among others -to gene transcription/translation and its regulation, to immune response and the nervous system in general. We also detected 12 loss-of-function mutations, although their actual effect is unclear. Several genetic pathways were also identified, which pathways may be tempting candidates to be investigated in large sample sizes in order to confirm their relevance in extreme longevity in dogs (and possibly, in humans). We hypothesize a possible link between extreme longevity and the regulation of gene transcription/translation, which hypothesis should be further investigated in the future. This phenomenon could define an interesting direction for future research aiming to better understand longevity. The presented preliminary results highlight the utility of the companion dog in the study of the genetic background of longevity and aging.
Several studies on age-related cognitive decline in dogs involve laboratory dogs and prolonged tr... more Several studies on age-related cognitive decline in dogs involve laboratory dogs and prolonged training. We developed two spatial tasks that required a single 1-h session. We tested 107 medium-large sized dogs: Byoung^(N=41, aged 2.5-6.5 years) and Bold^(N=66, aged 8-14.5 years). Our results indicated that, in a discrimination learning task and in a reversal learning task, young dogs learned significantly faster than the old dogs, indicating that these two tasks could successfully be used to investigate differences in spatial learning between young and old dogs. We also provide two novel findings. First, in the reversal learning, the dogs trained based on the location of stimuli learned faster than the dogs trained based on stimulus characteristics. Most old dogs did not learn the task within our cut-off of 50 trials. Training based on an object's location is therefore more appropriate for reversal learning tasks. Second, the contrast between the response to the positive and negative stimuli was narrower in old dogs, compared to young dogs, during the reversal learning task, as well as the cognitive bias test. This measure favors comparability between tasks and between studies. Following the cognitive bias test, we could not find any indication of differences in the positive and negative expectations between young and old dogs. Taken together, these findings do not support the hypothesis that old dogs have more negative expectations than young dogs and the use of the cognitive bias test in older dogs requires further investigation.
To describe the extent of age-related cognitive decline in dogs, information regarding the baseli... more To describe the extent of age-related cognitive decline in dogs, information regarding the baseline occurrence of associated behaviours in the general population is necessary. With a seven-item, data driven Age-Related Changes scale, we evaluated the relationship between sensory functions, training, sex, and the occurrence of behavioural signs associated with cognitive decline across the whole adult lifespan. The twofold difference in lifespan between small and large dogs presents challenges for ageing studies, with no widely accepted method to control for body size as it relates to chronological age and longevity, when comparing behavioural signs of cognitive decline. To address this issue, we utilized relative age, calculated using the estimated expected lifespan of the individuals in our questionnaire study. Signs of cognitive decline were already detectable in' Mature' dogs (at 50-75% of the expected lifespan). Visual, auditory and olfactory impairments all resulted in significantly higher scores on the Age-Related Changes scale. Participating in dog training activities was revealed to be protective against behavioural signs of cognitive decline in aged dogs as perceived by the owners. These results revealed possible beneficial effects of training on cognitive ageing and emphasize the importance of routinely screening the sensory capacities of ageing dogs. (D. Szabó).
The intestinal tract affects the brain through metabolites produced by gut-inhabiting bacteria. I... more The intestinal tract affects the brain through metabolites produced by gut-inhabiting bacteria. In this study, we show that the number of errors the dogs commit in a short-term memory test and also their age is linked to the gut microbiome composition. The proportion of Fusobacteria is lower in older animals. Dogs with better memory performance (i.e., fewer mistakes) have relatively fewer Actinobacteria in their fecal samples collected right after the behavior test. This result is in agreement with the high abundance of some Actinobacteria in the gastrointestinal tract of persons living with Alzheimer's disease. Links between memory performance and gut microbiota have been reported on rodents but not on dogs before. The research opens up new venues in canine aging and neurodevelopmental research.
Background and aims: Dogs have recently become an important model species for comparative social ... more Background and aims: Dogs have recently become an important model species for comparative social and cognitive neuroscience. Brain template-related label maps are essential for functional magnetic resonance imaging (fMRI) data analysis, to localize neural responses. In this study, we present a detailed, individual-based, T1-weighted MRI-based brain label map used in dog neuroimaging analysis. Methods: A typical, medium-headed dog (a 7.5-year-old male Golden Retriever) was selected from a cohort of 22 dogs, based on brain morphology (shape, size, and gyral pattern), to serve as the template for a label map. Results: Eighty-six 3-dimensional labels were created to highlight the main cortical (cerebral gyri on the lateral and medial side) and subcortical (thalamus, caudate nucleus, amygdala, and hippocampus) structures of the prosencephalon and diencephalon, and further main parts of brainstem (mesencephalon and rhombencephalon). Discussion: Importantly, this label map is (a) considerably more detailed than any available dog brain template; (b) it is easy to use with freeware and commercial neuroimaging software for MRI and fMRI analysis; and (c) it can be registered to other existing templates, including a recent average-based dog brain template. Using the coordinate system and label map proposed here can enhance precision and standard localization during future canine neuroimaging studies.
In humans, age-related changes in personality occur in a non-random fashion with respect to their... more 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.
In dogs, the social and spatial restriction associated with living in a kennel environment could ... more In dogs, the social and spatial restriction associated with living in a kennel environment could lead to chronic stress and the development of abnormal behaviors ("kennel-dog syndrome"). However, little is known about how kenneled dogs differ from their conspecifics living as pets in human families. In the current study, using a test battery exposing the dogs to novel stimuli, we compared the behavior of three groups of beagles: (1) kenneled dogs living in a restricted environment with limited human contact (N = 78), (2) family dogs living in human families as pets (N = 37), and (3) adopted dogs born in the kenneled population but raised in human families (N = 13). We found one factor comprising most of the test behaviors, labeled as Responsiveness. Family dogs and adopted dogs scored higher in Responsiveness than kenneled dogs. However, 23% of the kenneled dogs were comparable to family and adopted dogs based on a cluster analysis, indicating a similar (positive) reaction to novel stimuli, while 77% of the kenneled dogs were unresponsive (mostly immobile) in at least part of the test. To assess if the behavioral difference between the family and kenneled dogs could be due to genetic divergence of these two populations and/or to lower genetic diversity of the kenneled dogs, we analyzed their genetic structure using 11 microsatellite markers. We found no significant difference between the populations in their genetic diversity (i.e., heterozygosity, level of inbreeding), nor any evidence that the family and kenneled populations originated from different genetic pools. Thus, the behavior difference between the groups more likely reflects a G × E interaction, that is, the influence of specific genetic variants manifesting under specific environmental conditions (kennel life). Nevertheless, some kenneled individuals were (genetically) more resistant to social and environmental deprivation. Selecting for such animals could strongly improve the welfare of kenneled dog populations. Moreover, exploring the genetic background of their higher resilience could also help to better understand the genetics behind stress-and fear-related behaviors.
Introduction: The aim of this study was to explore spontaneous social interactions between dyads ... more Introduction: The aim of this study was to explore spontaneous social interactions between dyads of unfamiliar adult dogs. Although intraspecific encounters are frequent events in the life of pet dogs, the factors that might influence encounters, such as sex, dyad composition, reproductive status, age, and state of cohabitation (keeping the dogs singly or in groups), remained unexplored. Methods: In this study, we assigned unfamiliar, non-aggressive dogs to three types of dyads defined by sex and size. We observed their unrestrained, spontaneous behaviors in an unfamiliar dog park, where only the two dogs, the owners, and experimenter were present. Results: We found that the dogs, on average, spent only 17% of the time (less than 1 min) in proximity. Sex, dyad composition, reproductive status, and age influenced different aspects of the interactions in dyads. Female dogs were more likely to initiate the first contact in their dyad but later approached the partner less frequently, were less likely to move apart, and displayed less scent marking. Following and moving apart were more frequent in male-male interactions. Neutered dogs spent more time following the other dog and sniffed other dogs more frequently. The time companion dogs spent in proximity and number of approaches decreased with age. Conclusion: The study provides guidance for dog owners about the outcomes of intraspecific encounters based on the dog's age, sex, and reproductive status, as well as the sex of the interacting partner.
Dominance is well defined in ethology, debated in psychology, and is often unclear among the dog ... more Dominance is well defined in ethology, debated in psychology, and is often unclear among the dog owning public and in the press. However, to date, no study has examined how owners perceive dominance in dogs, and what different behaviours and personality types are used to describe dominant and subordinate individuals. A questionnaire study was launched to investigate the external validity of owner-derived estimates of dominance in dog dyads sharing the same household (N = 1,151). According to the owners, dogs rated as dominant (87%) have priority access to resources (resting place, food, and rewards), undertake certain tasks (defend and lead the group, bark more), display dominance (win fights, lick the other's mouth less, and mark over the other's urine), share certain personality traits (smarter, more aggressive and impulsive), and are older than their partner dog (all p < 0.0001). An age-related hypothesis has been suggested to explain dominance in dogs; but we found that dog age did not explain the occurrence of dominance related behaviours over the owners' estimate of dominance status. Results suggest that owner-derived reports of dominance ranks of dogs living in multi-dog households correspond to ethologically valid behavioural markers of dominance. Size and physical condition were unrelated to the perceived dominance. Surprisingly, in mixed-sex dyads, females were more frequently rated as dominant than males, which might correspond to a higher proportion of neutered females in this subgroup. For future studies that wish to allocate dominance status using owner report, we offer a novel survey.
Most common methods that directly show macro-or microscopic anatomy of the brain usually require ... more Most common methods that directly show macro-or microscopic anatomy of the brain usually require the removal of the organ from the neurocranium. However, the brain can be revealed in situ by using proper sectioning techniques. Our aim was to both improve the cryosectioning method, test its limits and create a high-resolution macro-anatomical image series of a Beagle brain, which at the time of the study did not exist. A two-year-old female Beagle has been scanned with CT and MRI ante and post mortem, then the arteries of the head were filled with red resin. After freezing to -80˚C, a neurocranium block was created and was embedded into a water-gelatin mix. Using a special milling device and a DSLR camera, 1112 consecutive RGB-color cryosections were made with a 100 μm layer thickness and captured in high resolution (300 dpi, 24-bit color, and pixel size was 19.5 x 19.5 μm). Image post-processing was done with Adobe Photoshop CS3 and Thermo Scientific Amira 6.0 softwares, and as a result of the proper alignment and coregistration, visualization and comparing was possible with all the applied imaging modalities (CT, MRI, cryosectioning) in any arbitrary plane. Surface models from the arteries, veins, brain and skull were also generated after segmentation in the same coordinate system, giving a unique opportunity for comparing the two-dimensional and three-dimensional anatomy. This is the first study which focuses directly to this high-definition multimodal visualization of the canine brain, and it provides the most accurate results compared to previous cryosectioning studies, as using an improved method, higher image quality, more detailed image, proper color fidelity and lower artefact formation were achieved. Based on the methodology we described, it can serve as a base for future multimodal (CT, MR, augmented-or virtual reality) imaging atlases for medical, educational and scientific purposes.
Dogs living on Bali Island have been free-ranging for thousands of years. A large group of expatr... more Dogs living on Bali Island have been free-ranging for thousands of years. A large group of expatriates sometimes adopt Bali dogs and keep them restricted to their houses and backyards, as is typical in modern western cultures. This provides us with the unique opportunity to compare the personality traits of dogs to their lifestyle either living as human companions or as free-ranging animals, exploring at the same time the impact of demographic variables (such as age, sex, and neutered status) on personality. After controlling for internal consistency of the scales and between-observer variation, we found that free-ranging Bali dogs were rated as less active, less excitable, less aggressive towards animals, and less inclined to chase animals or humans than Bali dogs living as human companions. Among free-ranging dogs, females were found to be more excitable. Females in the whole sample were also more fearful of people. The results of this preliminary study suggest that a change in lifestyle, i.e. being adopted, and living in a confined environment has negative consequences on some canine personality traits, such as activity/excitability, aggression towards animals, and prey drive.
Visual and auditory impairments can have a large impact on performance in cognitive tests. It is ... more Visual and auditory impairments can have a large impact on performance in cognitive tests. It is important to evaluate the sensory capacities of dogs before enrolling them in cognitive tests, in order to exclude sensory impairment as confounding effect. Therefore we designed multiple non-invasive testing paradigms to detect subjects with potential auditory and visual impairment, without requiring extensive training for the dog. Multiple testing was a means to add internal reliability, and to reduce the risk of false positives due to habituation and previous learning or false negatives due to random errors. Our sensory test battery consisted of four subtests: (1) 'Clapping' auditory test, (2) 'Recorded sound' auditory test, (3) 'Distance' visual test, and (4) 'Darkness' visual test. The 'Clapping test' was similar to the clapping test used by veterinarians, with the addition that the clapping was performed at various distances from the dog. In the 'Recorded sound test', the dogs' reaction to various sounds played back at different volumes. In the 'Distance test' we placed a small piece of food on one of four plates placed on the grounds at varying distances from the dogs. In the 'Darkness test', we measured the dogs' performance in walking through an S-shaped route during artificial dusk and daylight-like conditions.
Aging is the largest risk factor in many diseases and mortality alike. As the elderly population ... more Aging is the largest risk factor in many diseases and mortality alike. As the elderly population is expected to increase at an accelerating rate in the future, these phenomena will pose a growing socio-economic burden on societies. To successfully cope with this challenge, a deeper understanding of aging is crucial. In many aspects, the companion dog is an increasingly popular model organism to study aging, with the promise of producing results that are more applicable to humans than the findings that come from the studies of classical model organisms. In this preliminary study we used the whole-genome sequence of two extremely old dogs -age: 22 and 27 years (or 90-135% more, than the average lifespan of dogs) -in order to make the first steps to understand the genetic background of extreme longevity in dogs. We identified more than ∼80 1000 novel SNPs in the two dogs (7500 of which overlapped between them) when compared to three publicly available canine SNP databases, which included SNP information from850 dogs. Most novel mutations (∼52000 SNPs) were identified at non-coding regions, while 4.6% of the remaining SNPs (n∼1600) were at exons, including 670 missense variants -76 of which overlapped between the two animalsacross 472 genes. Based on their gene ontologies, these genes were related -among others -to gene transcription/translation and its regulation, to immune response and the nervous system in general. We also detected 12 loss-of-function mutations, although their actual effect is unclear. Several genetic pathways were also identified, which pathways may be tempting candidates to be investigated in large sample sizes in order to confirm their relevance in extreme longevity in dogs (and possibly, in humans). We hypothesize a possible link between extreme longevity and the regulation of gene transcription/translation, which hypothesis should be further investigated in the future. This phenomenon could define an interesting direction for future research aiming to better understand longevity. The presented preliminary results highlight the utility of the companion dog in the study of the genetic background of longevity and aging.
Several studies on age-related cognitive decline in dogs involve laboratory dogs and prolonged tr... more Several studies on age-related cognitive decline in dogs involve laboratory dogs and prolonged training. We developed two spatial tasks that required a single 1-h session. We tested 107 medium-large sized dogs: Byoung^(N=41, aged 2.5-6.5 years) and Bold^(N=66, aged 8-14.5 years). Our results indicated that, in a discrimination learning task and in a reversal learning task, young dogs learned significantly faster than the old dogs, indicating that these two tasks could successfully be used to investigate differences in spatial learning between young and old dogs. We also provide two novel findings. First, in the reversal learning, the dogs trained based on the location of stimuli learned faster than the dogs trained based on stimulus characteristics. Most old dogs did not learn the task within our cut-off of 50 trials. Training based on an object's location is therefore more appropriate for reversal learning tasks. Second, the contrast between the response to the positive and negative stimuli was narrower in old dogs, compared to young dogs, during the reversal learning task, as well as the cognitive bias test. This measure favors comparability between tasks and between studies. Following the cognitive bias test, we could not find any indication of differences in the positive and negative expectations between young and old dogs. Taken together, these findings do not support the hypothesis that old dogs have more negative expectations than young dogs and the use of the cognitive bias test in older dogs requires further investigation.
To describe the extent of age-related cognitive decline in dogs, information regarding the baseli... more To describe the extent of age-related cognitive decline in dogs, information regarding the baseline occurrence of associated behaviours in the general population is necessary. With a seven-item, data driven Age-Related Changes scale, we evaluated the relationship between sensory functions, training, sex, and the occurrence of behavioural signs associated with cognitive decline across the whole adult lifespan. The twofold difference in lifespan between small and large dogs presents challenges for ageing studies, with no widely accepted method to control for body size as it relates to chronological age and longevity, when comparing behavioural signs of cognitive decline. To address this issue, we utilized relative age, calculated using the estimated expected lifespan of the individuals in our questionnaire study. Signs of cognitive decline were already detectable in' Mature' dogs (at 50-75% of the expected lifespan). Visual, auditory and olfactory impairments all resulted in significantly higher scores on the Age-Related Changes scale. Participating in dog training activities was revealed to be protective against behavioural signs of cognitive decline in aged dogs as perceived by the owners. These results revealed possible beneficial effects of training on cognitive ageing and emphasize the importance of routinely screening the sensory capacities of ageing dogs. (D. Szabó).
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Papers by Eniko Kubinyi