Dogs' ability to follow temporarily invisible moving objects: the ability to track and expect is shaped by experience.

Visually tracking a moving object, even if it becomes temporarily invisible, is an important skill for animals living in complex environments. However, this ability has not been widely explored in dogs. To address this gap of knowledge and understand how experience contributes to such ability, we conducted two experiments using a violation of expectation paradigm. Dogs were shown an animation of a ball moving horizontally across a screen, passing behind an occluder, and reappearing with a timing that was faster, slower or congruent with its initial speed. In the first experiment, dogs (N = 15) were exposed to the incongruent conditions without prior experience; while in the second experiment, dogs (N = 37) were preliminarily exposed to the congruent stimulus. Dogs of the first experiment did not exhibit a surprise effect, as measured by latency to look away from the expected stimulus presentation area, in response to the incongruent conditions, suggesting they had not formed an expectation about the timing of reappearance. However, their latency to orient towards the reappearing ball depended on the condition, suggesting they were able, to some extent, to visually keep track of the stimulus' trajectory. Dogs of the second experiment were surprised when the ball stayed behind the occluder longer than expected, but showed no difference in latency to orient across conditions. This suggests they had overcome the visual tracking mechanism and had formed expectations about the timing of reappearance. In conclusion, dogs seem to use a low-level mechanism to keep visual track of a temporarily disappearing moving object, but experience is required to make expectation about its trajectory.

Meowing dogs: can dogs recognize cats in a cross-modal violation of expectancy task (Canis familiaris)?

Dogs can recognize conspecifics in cross-modal audio-video presentations. In this paper, we aimed at exploring if such capability extends to the recognition of cats, and whether it is influenced by exposure to these animals. To reach our aim, we enrolled 64 pet dogs. Half of the dogs were currently living with cats, while the rest had never been living with cats, nor were at the time of the experiment. All dogs underwent a cross-modal violation of expectancy experiment, where they were presented with either a cat or a dog vocalization, followed by a video of either species on a blank background. The result revealed that dogs did not exhibit a surprise reaction towards the incoherent stimuli of a cat vocalization and a dog video or vice-versa, implying that they had not recognized the stimuli portraying cats. The pattern of results did not differ between dogs living or not with cats, implying that exposure to a limited number of cats, however, prolonged, is not sufficient to grant dogs with the ability to recognize them on audio-video presentations. We propose that the lack of recognition could be due to the small number of individual cats the dogs are regularly exposed to, or to the possible lack of early exposure to cats during the socialization phase.

Yes, dogs are susceptible to the Kanizsa's triangle illusion: A reply to Pepperberg.

A recent paper by Pepperberg, Learning & Behavior, 51, 5-6, (2023) enquires about the validity of the finding that dogs are susceptible to the Kanizsa's triangle illusion, reported by Lõoke and coauthors (Lõoke et al., Animal Cognition, 25, 43-51, 2022). Here we elaborate on the matter, providing both theoretical considerations and further data, supporting the soundness of our previous conclusions.

Is this love? Sex differences in dog-owner attachment behavior suggest similarities with adult human bonds.

Sex differences in the behavioral responses of Labrador Retriever dogs in the Strange Situation Test were explored. Behaviors expressed by dogs during seven 3-min episodes were analyzed through a Principal Component Analysis (PCA). The scores of factors obtained were analyzed with a Generalized Linear Mixed Model to reveal the effects of the dog's sex and age and the owner's sex. In Episode 1 (dog and owner) and 5 (dog alone), the PCA identified three and two factors, respectively, which overall explained 68.7% and 59.8% of the variance, with no effect of sex. In Episodes 2 (dog, owner, and stranger), 3 and 6 (dog and stranger), and 4 and 7 (dog and owner), the PCA identified four factors, which overall explained 51.0% of the variance. Effects of sex were found on: Factor 1 (distress), with lower scores obtained by females in Episode 2 and higher in Episode 3; Factor 2 (sociability), which was overall higher in females; Factor 3 (separation-distress), with females, but not males, obtaining higher scores when left with the stranger than when with the owner. Therefore, females were overall more social but seemed more affected than males by the owner's absence. Parallels can be traced between our results and sex differences found in adult human romantic attachment, suggesting that the dog-owner bond has characteristics that are not found in the infant-mother relationship.

Dogs (Canis lupus familiaris) are susceptible to the Kanizsa's triangle illusion.

The ability to complete partially missing contours is widespread across the animal kingdom, but whether this extends to dogs is still unknown. To address this gap in knowledge, we assessed dogs' susceptibility to one of the most common contour illusions, the Kanizsa's triangle. Six dogs were trained to discriminate a triangle from other geometrical figures using a two-alternative conditioned discrimination task. Once the learning criterion was reached, dogs were presented with the Kanizsa's triangle and a control stimulus, where inducers were rotated around their centre, so as to disrupt what would be perceived as a triangle by a human observer. As a group, dogs chose the illusory triangle significantly more often than control stimuli. At the individual level, susceptibility to the illusion was shown by five out of six dogs. This is the first study where dogs as a group show susceptibility to a visual illusion in the same manner as humans. Moreover, the analyses revealed a negative effect of age on susceptibility, an effect that was also found in humans. Altogether, this suggests that the underling perceptual mechanisms are similar between dogs and humans, and in sharp contrast with other categories of visual illusions to which the susceptibility of dogs has been previously assessed.

I know a dog when I see one: dogs (Canis familiaris) recognize dogs from videos.

Several aspects of dogs' visual and social cognition have been explored using bi-dimensional representations of other dogs. It remains unclear, however, if dogs do recognize as dogs the stimuli depicted in such representations, especially with regard to videos. To test this, 32 pet dogs took part in a cross-modal violation of expectancy experiment, during which dogs were shown videos of either a dog and that of an unfamiliar animal, paired with either the sound of a dog barking or of an unfamiliar vocalization. While stimuli were being presented, dogs paid higher attention to the exit region of the presentation area, when the visual stimulus represented a dog than when it represented an unfamiliar species. After exposure to the stimuli, dogs' attention to different parts of the presentation area depended on the specific combination of visual and auditory stimuli. Of relevance, dogs paid less attention to the central part of the presentation area and more to the entrance area after being exposed to the barking and dog video pair, than when either was paired with an unfamiliar stimulus. These results indicate dogs were surprised by the latter pairings, not by the former, and were interested in where the barking and dog pair came from, implying recognition of the two stimuli as belonging to a conspecific. The study represents the first demonstration that dogs can recognize other conspecifics in videos.

Dogs (Canis familiaris) recognise our faces in photographs: implications for existing and future research.

Dogs are an ideal species to investigate phylogenetic and ontogenetic factors contributing to face recognition. Previous research has found that dogs can recognise their owner using visual information about the person's face, presented live. However, a thorough investigation of face processing mechanisms requires the use of graphical representations and it currently remains unclear whether dogs are able to spontaneously recognise human faces in photographs. To test this, pet dogs (N = 60) were briefly separated from their owners and, to achieve reunion, they needed to select the location indicated by a photograph of their owner's face, rather than that of an unfamiliar person concurrently presented. Photographs were taken under optimal and suboptimal (non-frontally oriented and unevenly illuminated faces) conditions. Results revealed that dogs approached their owner significantly above chance level when presented with photos taken under optimal conditions. Further analysis revealed no difference in the probability of choosing the owner between the optimal and suboptimal conditions. Dogs were more likely to choose the owner if they directed a higher percentage of looking time towards the owner's photograph compared to the stranger's one. In addition, the longer the total viewing time of both photos, the higher the probability that dogs chose the stranger. A main effect of dogs' sex was also obtained, with a higher probability of male dogs choosing the owner's photograph. This study provides direct evidence that dogs are able to recognise their owner's face from photographs. The results imply that motion and three-dimensional information is not necessary for recognition. The findings also support the ecological valence of such stimuli and increase the validity of previous investigations into dog cognition that used two-dimensional representations of faces. The effects of attention may reflect differences at the individual level in attraction towards novel faces or in the recruitment of different face processing mechanisms.

The effect of experience and of dots' density and duration on the detection of coherent motion in dogs.

Knowledge about the mechanisms underlying canine vision is far from being exhaustive, especially that concerning post-retinal elaboration. One aspect that has received little attention is motion perception, and in spite of the common belief that dogs are extremely apt at detecting moving stimuli, there is no scientific support for such an assumption. In fact, we recently showed that dogs have higher thresholds than humans for coherent motion detection (Kanizsar et al. in Sci Rep UK 7:11259, 2017). This term refers to the ability of the visual system to perceive several units moving in the same direction, as one coherently moving global unit. Coherent motion perception is commonly investigated using random dot displays, containing variable proportions of coherently moving dots. Here, we investigated the relative contribution of local and global integration mechanisms for coherent motion perception, and changes in detection thresholds as a result of repeated exposure to the experimental stimuli. Dogs who had been involved in the previous study were given a conditioned discrimination task, in which we systematically manipulated dot density and duration and, eventually, re-assessed our subjects' threshold after extensive exposure to the stimuli. Decreasing dot duration impacted on dogs' accuracy in detecting coherent motion only at very low duration values, revealing the efficacy of local integration mechanisms. Density impacted on dogs' accuracy in a linear fashion, indicating less efficient global integration. There was limited evidence of improvement in the re-assessment but, with an average threshold at re-assessment of 29%, dogs' ability to detect coherent motion remains much poorer than that of humans.

Sex differences in dogs' social learning of spatial information.

We used a modified version of the Do as I Do paradigm to investigate dogs' preference and flexibility in the acquisition of different types of spatial information in social learning situations. When required to match the location of the demonstration, dogs (N = 16) preferentially relied on allocentric information, i.e., the relationship between the location of the demonstration and the various objects surrounding it. However, when allocentric cues were inadequate to solve the task, dogs learned to rely on egocentric information, i.e., the direction-left/right-taken by the human demonstrator. The ease of resorting to the non-preferred egocentric strategy was sex-dependent with males acquiring the egocentric strategy in fewer trials than females. This study shows that dogs rely preferentially on allocentric cues when recalling socially acquired spatial information. However, they are impressively flexible in switching to egocentric strategies according to the task requirements. Whether preference for the allocentric strategy in processing spatial information is embedded in the nature of social learning or restricted to our paradigm is an open question. This study also supports the idea that sex differences in cognitive domains are widespread among mammals and calls for further investigations aimed at shedding light on the evolution, function and mechanisms of such differences.

Recognition of human faces by dogs (Canis familiaris) requires visibility of head contour.

Researchers have suggested that dogs are able to recognise human faces, but conclusive evidence has yet to be found. Experiment 1 of this study investigated whether dogs can recognise humans using visual information from the face/head region, and whether this also occurs in conditions of suboptimal visibility of the face. Dogs were presented with their owner's and a stranger's heads, protruding through openings of an apparatus in opposite parts of the experimental setting. Presentations occurred in conditions of either optimal or suboptimal visibility; the latter featured non-frontal orientation, uneven illumination and invisibility of outer contours of the heads. Instances where dogs approached their owners with a higher frequency than predicted by chance were considered evidence of recognition. This occurred only in the optimal condition. With a similar paradigm, Experiment 2 investigated which of the alterations in visibility that characterised the suboptimal condition accounted for dogs' inability to recognise owners. Dogs approached their owners more frequently than predicted by chance if outer head contours were visible, but not if heads were either frontally oriented or evenly illuminated. Moreover, male dogs were slightly better at recognition than females. These findings represent the first clear demonstration that dogs can recognise human faces and that outer face elements are crucial for such a task, complementing previous research on human face processing in dogs. Parallels with face recognition abilities observed in other animal species, as well as with human infants, point to the relevance of these results from a comparative standpoint.

Global bias reliability in dogs (Canis familiaris).

Dogs enrolled in a previous study were assessed two years later for reliability of their local/global preference in a discrimination test with the same hierarchical stimuli used in the previous study (Experiment 1) and with a novel stimulus (Experiment 2). In Experiment 1, dogs easily re-learned to discriminate the positive stimulus; their individual global/local choices were stable compared to the previous study; and an overall clear global bias was found. In Experiment 2, dogs were slower in acquiring the initial discrimination task; the overall global bias disappeared; and, individually, dogs tended to make inverse choices compared to the original study. Spontaneous attention toward the test stimulus resembling the global features of the probe stimulus was the main factor affecting the likeliness of a global choice of our dogs, regardless of the type of experiment. However, attention to task-irrelevant elements increased at the expense of attention to the stimuli in the test phase of Experiment 2. Overall, the results suggest that the stability of global bias in dogs depends on the characteristics of the assessment contingencies, likely including the learning requirements of the tasks. Our results also clearly indicate that attention processes have a prominent role on dogs' global bias, in agreement with previous findings in humans and other species.

Hierarchical stimulus processing by dogs (Canis familiaris).

The purpose of this study was to assess the visual processing of global and local levels of hierarchical stimuli in domestic dogs. Fourteen dogs were trained to recognise a compound stimulus in a simultaneous conditioned discrimination procedure and were then tested for their local/global preference in a discrimination test. As a group, dogs showed a non-significant trend for global precedence, although large inter-individual variability was observed. Choices in the test were not affected by either dogs' sex or the type of stimulus used for training. However, the less time a dog took to complete the discrimination training phase, the higher the probability that it chose the global level of test stimulus. Moreover, dogs that showed a clear preference for the global level in the test were significantly less likely to show positional responses during discrimination training. These differences in the speed of acquisition and response patterns may reflect individual differences in the cognitive requirements during discrimination training. The individual variability in global/local precedence suggests that experience in using visual information may be more important than predisposition in determining global/local processing in dogs.

Determining Hearing Thresholds in Dogs Using the Staircase Method.

There is a growing interest in performing playback experiments to understand which acoustical cues trigger specific behavioral/emotional responses in dogs. However, very limited studies have focused their attention on more basic aspects of hearing such as sensitivity, i.e., the identification of minimal intensity thresholds across different frequencies. Most previous studies relied on electrophysiological methods for audiograms for dogs, but these methods are considered less accurate than assessments based on behavioral responses. To our knowledge, only one study has established hearing thresholds using a behavioral assessment on four dogs but using a method that did not allow potential improvement throughout the sessions. In the present study, we devised an assessment procedure based on a staircase method. Implying the adaptation of the assessed intensity on the dogs' performance, this approach grants several assessments around the actual hearing threshold of the animal, thereby increasing the reliability of the result. We used such a method to determine hearing thresholds at three frequencies (0.5, 4.0, and 20.0 kHz). Five dogs were tested in each frequency. The hearing thresholds were found to be 19.5 ± 2.8 dB SPL at 0.5 kHz, 14.0 ± 4.5 dB SPL at 4.0 kHz, and 8.5 ± 12.8 dB SPL at 20.0 kHz. No improvement in performance was visible across the procedure. While the thresholds at 0.5 and 4.0 kHz were in line with the previous literature, the threshold at 20 kHz was remarkably lower than expected. Dogs' ability to produce vocalization beyond 20 kHz, potentially used in short-range communication, and the selective pressure linked to intraspecific communication in social canids are discussed as potential explanations for the sensitivity to higher frequencies.

Canine Behavioral Assessment and Research Questionnaire (C-BARQ): Validation of the Italian Translation.

The aim of this study is to develop an Italian translation of the 100-item Canine Behavioral Assessment and Research Questionnaire (C-BARQ) version and to validate its psychometric properties, in order to facilitate systematic, large-scale studies on dog behavior for Italian-speaking dog owners. A total number of 803 responses by dog owners were collected online. Using the Principal Axis Method and Common Factor Analysis with Quartimin oblique rotation (p < 0.05), a factorial structure was found including 13 factors composed of 62 items and explaining 53.5% of the total variance. Eight factors showed high reliability (Cronbach's alpha > 0.70), namely: stranger-directed aggression/fear, dog-directed fear, owner-directed aggression, separation-related behavior, chasing, dog-directed aggression, attachment/attention seeking, and elimination problems. Three factors were slightly under the threshold and two had only modest reliability (non-social fear, energy level, touch sensitivity, excitability and trainability). A potential explanation for factors with low reliability is that the composing items do not describe behaviors resulting from homogeneous stimuli or situations. Although our factorial structure resembled in most respects that of the most recently published Canadian version, some important exceptions are present regarding dog rivalry, intraspecific aggression, fear/aggression towards strangers, touch sensitivity and chewing inappropriate objects. Such differences may be due to demographic and/or cultural differences between the sampled populations. Overall, the results suggest that a 62-item Italian C-BARQ can be reliably used in studies on dog behavior.

Temperature and humidity index (THI) affects salivary cortisol (HC) and dehydroepiandrosterone (DHEA) concentrations in growing bulls following stress generated by performance test procedures.

The hypothalamus-pituitary-adrenal axis response to a challenge was proposed for genetic selection of robust and resilient animals. As ACTH (adrenocorticotropic hormone) test and hormone measurements in blood may result impractical, it may be useful to measure salivary hormones in response to natural stressors, after an accurate biological validation, to control factors that could contribute to the response. We evaluated whether animal handling during performance test affects salivary HC and DHEA secretion and could be used for selection. We tested the effects of habituation to repeated handling and THI as putative bias. Bull calves (N = 273) undergoing performance test were sampled at 8-9 and 11-13 months (N = 101), 8-9 months (N = 131), or 11-13 months (N = 41). On each test day (D0), calves were isolated, conducted to a squeeze chute and immobilized for 6 min. Saliva samples were collected in the morning after feed administration (T0), and after 6 min immobilization in the squeeze chute (T1) for HC and DHEA measurement. Environmental temperature and relative humidity were recorded every hour from 1:00 h to 24:00 h during the 6 days before the performance test and on D0. Salivary HC and DHEA concentrations were higher in T1 (p < 0.01), although a clear individual positive response to handling could be observed in less than 10% of subjects. The mixed model revealed: (i) HC and HC/DHEA were higher in Young bulls (p < 0.05). (ii) The time of T0 sample collection significantly affected DHEA (p < 0.01) and HC/DHEA (p < 0.05). (iii) THI affected both steroids (p < 0.001) but not HC/DHEA. Spearman correlations suggested that THI weakly affected salivary HC at T0 only (ρ = 0.150, p < 0.01), while moderate statistically significant correlations were found between DHEA and THI at T0 (ρ = 0.316, p < 0.001), and T1 (ρ = 0.353, p < 0.001). Salivary HC and DHEA in response to handling procedures might identify subpopulations of subjects with sensitive HPA axis. Habituation to repeated handling played a role, as the hormone response was lower in older animals. Chronic exposure to high THI had a minor effect on salivary HC visible at T0. A more intense THI effect was observed on salivary DHEA concentrations at both T0 and T1, which should be worth of further investigations.

Sound Localization Ability in Dogs.

The minimum audible angle (MAA), defined as the smallest detectable difference between the azimuths of two identical sources of sound, is a standard measure of spatial auditory acuity in animals. Few studies have explored the MAA of dogs, using methods that do not allow potential improvement throughout the assessment, and with a very small number of dog(s) assessed. To overcome these limits, we adopted a staircase method on 10 dogs, using a two-forced choice procedure with two sound sources, testing angles of separation from 60° to 1°. The staircase method permits the level of difficulty for each dog to be continuously adapted and allows for the observation of improvement over time. The dogs' average MAA was 7.6°, although with a large interindividual variability, ranging from 1.3° to 13.2°. A global improvement was observed across the procedure, substantiated by a gradual lowering of the MAA and of choice latency across sessions. The results indicate that the staircase method is feasible and reliable in the assessment of auditory spatial localization in dogs, highlighting the importance of using an appropriate method in a sensory discrimination task, so as to allow improvement over time. The results also reveal that the MAA of dogs is more variable than previously reported, potentially reaching values lower than 2°. Although no clear patterns of association emerged between MAA and dogs' characteristics such as ear shape, head shape or age, the results suggest the value of conducting larger-scale studies to determine whether these or other factors influence sound localization abilities in dogs.

The Cat-Owner Relationship: Validation of the Italian C/DORS for Cat Owners and Correlation with the LAPS.

Globally, most people now own a pet. Scales have been developed to understand the impact of pet ownership on people's lives and to measure specific aspects of the owner-pet relationship. The Cat-/Dog-Owner Relationship Scale (C/DORS) is a tool developed to investigate this relationship in both dog and cat owners. The aim of the study was to refine and validate the C/DORS for cat owners in Italian. Exploratory Factor Analysis and Confirmatory Factor Analysis (CFA) were used to determine the most appropriate factor model. Construct validity was confirmed by correlating the C/DORS subscales with the Lexington Attachment to Pets (LAPS) subscales. Results confirmed the original structure of the English version (i.e., three factors: Pet Owner Interaction = POI, Perceived Emotional Closeness = PEC, Perceived Costs = PC) and CFA confirmed the structure of LAPS and C/DORS scales. Cronbach's alpha demonstrated the Italian version of the two scales to have good internal reliability for all domains. Owners of cats living exclusively indoors reported higher scores on POI and PEC compared to indoor/outdoor cats. Owning both cats and dogs was correlated with lower scores on POI, and fewer perceived costs (i.e., PC) of cat ownership. Finally, behaviour problems, not being neutered, and lack of previous experience with cat ownership were associated with lower scores on PC.

Dogs (canis familiaris) underestimate the quantity of connected items: first demonstration of susceptibility to the connectedness illusion in non-human animals.

In humans, numerical estimation is affected by perceptual biases, such as those originating from the spatial arrangement of elements. Different animal species can also make relative quantity judgements. This includes dogs, who have been proposed as a good model for comparative neuroscience. However, dogs do not show the same perceptual biases observed in humans. Thus, the exact perceptual/cognitive mechanisms underlying quantity estimations in dogs and their degree of similarity with humans are still a matter of debate. Here we explored whether dogs are susceptible to the connectedness illusion, an illusion based on the tendency to underestimate the quantity of interconnected items. Dogs were first trained to choose the larger of two food arrays. Then, they were presented with two arrays containing the same quantity of food, of which one had items interconnected by lines. Dogs significantly selected the array with unconnected items, suggesting that, like in humans, connectedness determines underestimation biases, possibly disrupting the perceptual system's ability to segment the display into discrete objects. The similarity in dogs' and humans' susceptibility to the connectedness, but not to other numerical illusions, suggests that different mechanisms are involved in the estimation of quantity of stimuli with different characteristics.

Dogs fail to recognize a human pointing gesture in two-dimensional depictions of motion cues.

Few studies have investigated biological motion perception in dogs and it remains unknown whether dogs recognise the biological identity of two-dimensional animations of human motion cues. To test this, we assessed the dogs' (N = 32) responses to point-light displays of a human performing a pointing gesture towards one of two pots. At the start of the experiment the demonstrator was a real-life person, but over the course of the test dogs were presented with two-dimensional figurative representations of pointing gestures in which visual information was progressively removed until only the isolated motion cues remained. Dogs' accuracy was above chance level only with real-life and black-and-white videos, but not with the silhouette or the point-light figure. Dogs' accuracy during these conditions was significantly lower than in the real-life condition. This result could not be explained by trial order since dogs' performance was still not higher than chance when only the point-light figure condition was presented after the initial demonstration. The results imply that dogs are unable to recognise humans in two-dimensional depictions of human motion cues only. In spite of extensive exposure to human movement, dogs need more perceptual cues to detect equivalence between human two-dimensional animations and the represented living entity.

Do Dehydroepiandrosterone (DHEA) and Its Sulfate (DHEAS) Play a Role in the Stress Response in Domestic Animals?

In animal husbandry, stress is often associated with poor health and welfare. Stress occurs when a physiological control system detects a state of real or presumptive threat to the animal's homeostasis or a failure to control a fitness-critical variable. The definition of stress has mostly relied on glucocorticoids measurement, even though glucocorticoids represent one stress-response system, the hypothalamus-pituitary-adrenocortical axis, which is not precise enough as it is also related to metabolic regulation and activated in non-stressful situations (pleasure, excitement, and arousal). The mammal adrenal can synthesize the androgenic steroid dehydroepiandrosterone (DHEA) and its sulfate metabolite (DHEAS), which have been associated to the stress response in several studies performed mostly in humans and laboratory animals. Although the functions of these steroids are not fully understood, available data suggest their antagonistic effects on glucocorticoids and, in humans, their secretion is affected by stress. This review explores the scientific literature on DHEA and DHEAS release in domestic animals in response to stressors of different nature (inflammatory, physical, or social) and duration, and the extra-adrenal contribution to circulating DHEA. Then, the potential use of DHEA in conjunction with cortisol to improve the definition of the stress phenotype in farmed animals is discussed. Although the focus of this review is on farmed animals, examples from other species are reported when available.