Tailored haemodynamic response function increases detection power of fMRI in awake dogs (Canis familiaris).

Functional magnetic resonance imaging (fMRI) of awake and unrestrained dogs (Canis familiaris) has been established as a novel opportunity for comparative neuroimaging, promising important insights into the evolutionary roots of human brain function and cognition. However, data processing and analysis pipelines are often derivatives of methodological standards developed for human neuroimaging, which may be problematic due to profound neurophysiological and anatomical differences between humans and dogs. Here, we explore whether dog fMRI studies would benefit from a tailored dog haemodynamic response function (HRF). In two independent experiments, dogs were presented with different visual stimuli. BOLD signal changes in the visual cortex during these experiments were used for (a) the identification and estimation of a tailored dog HRF, and (b) the independent validation of the resulting dog HRF estimate. Time course analyses revealed that the BOLD signal in the primary visual cortex peaked significantly earlier in dogs compared to humans, while being comparable in shape. Deriving a tailored dog HRF significantly improved the model fit in both experiments, compared to the canonical HRF used in human fMRI. Using the dog HRF yielded significantly increased activation during visual stimulation, extending from the occipital lobe to the caudal parietal cortex, the bilateral temporal cortex, into bilateral hippocampal and thalamic regions. In sum, our findings provide robust evidence for an earlier onset of the dog HRF in two visual stimulation paradigms, and suggest that using such an HRF will be important to increase fMRI detection power in canine neuroimaging. By providing the parameters of the tailored dog HRF and related code, we encourage and enable other researchers to validate whether our findings generalize to other sensory modalities and experimental paradigms.

Training pet dogs for eye-tracking and awake fMRI.

In recent years, two well-developed methods of studying mental processes in humans have been successively applied to dogs. First, eye-tracking has been used to study visual cognition without distraction in unrestrained dogs. Second, noninvasive functional magnetic resonance imaging (fMRI) has been used for assessing the brain functions of dogs in vivo. Both methods, however, require dogs to sit, stand, or lie motionless while yet remaining attentive for several minutes, during which time their brain activity and eye movements are measured. Whereas eye-tracking in dogs is performed in a quiet and, apart from the experimental stimuli, nonstimulating and highly controlled environment, MRI scanning can only be performed in a very noisy and spatially restraining MRI scanner, in which dogs need to feel relaxed and stay motionless in order to study their brain and cognition with high precision. Here we describe in detail a training regime that is perfectly suited to train dogs in the required skills, with a high success probability and while keeping to the highest ethical standards of animal welfare-that is, without using aversive training methods or any other compromises to the dog's well-being for both methods. By reporting data from 41 dogs that successfully participated in eye-tracking training and 24 dogs IN fMRI training, we provide robust qualitative and quantitative evidence for the quality and efficiency of our training methods. By documenting and validating our training approach here, we aim to inspire others to use our methods to apply eye-tracking or fMRI for their investigations of canine behavior and cognition.

Domestic dogs as a comparative model for social neuroscience: Advances and challenges.

Dogs and humans have lived together for thousands of years and share many analogous socio-cognitive skills. Dog neuroimaging now provides insight into the neural bases of these shared social abilities. Here, we summarize key findings from dog fMRI identifying neocortical brain areas implicated in visual social cognition, such as face, body, and emotion perception, as well as action observation in dogs. These findings provide converging evidence that the temporal cortex plays a significant role in visual social cognition in dogs. We further briefly review investigations into the neural base of the dog-human relationship, mainly involving limbic brain regions. We then discuss current challenges in the field, such as statistical power and lack of common template spaces, and how to overcome them. Finally, we argue that the foundation has now been built to investigate and compare the neural bases of more complex socio-cognitive phenomena shared by dogs and humans. This will strengthen and expand the role of the domestic dog as a powerful comparative model species and provide novel insights into the evolutionary roots of social cognition.

Functional mapping of the somatosensory cortex using noninvasive fMRI and touch in awake dogs.

Dogs are increasingly used as a model for neuroscience due to their ability to undergo functional MRI fully awake and unrestrained, after extensive behavioral training. Still, we know rather little about dogs' basic functional neuroanatomy, including how basic perceptual and motor functions are localized in their brains. This is a major shortcoming in interpreting activations obtained in dog fMRI. The aim of this preregistered study was to localize areas associated with somatosensory processing. To this end, we touched N = 22 dogs undergoing fMRI scanning on their left and right flanks using a wooden rod. We identified activation in anatomically defined primary and secondary somatosensory areas (SI and SII), lateralized to the contralateral hemisphere depending on the side of touch, and importantly also activation beyond SI and SII, in the cingulate cortex, right cerebellum and vermis, and the sylvian gyri. These activations may partly relate to motor control (cerebellum, cingulate), but also potentially to higher-order cognitive processing of somatosensory stimuli (rostral sylvian gyri), and the affective aspects of the stimulation (cingulate). We also found evidence for individual side biases in a vast majority of dogs in our sample, pointing at functional lateralization of somatosensory processing. These findings not only provide further evidence that fMRI is suited to localize neuro-cognitive processing in dogs, but also expand our understanding of in vivo touch processing in mammals, beyond classically defined primary and secondary somatosensory cortices.

Functionally analogous body- and animacy-responsive areas are present in the dog (Canis familiaris) and human occipito-temporal lobe.

Comparing the neural correlates of socio-cognitive skills across species provides insights into the evolution of the social brain and has revealed face- and body-sensitive regions in the primate temporal lobe. Although from a different lineage, dogs share convergent visuo-cognitive skills with humans and a temporal lobe which evolved independently in carnivorans. We investigated the neural correlates of face and body perception in dogs (N = 15) and humans (N = 40) using functional MRI. Combining univariate and multivariate analysis approaches, we found functionally analogous occipito-temporal regions involved in the perception of animate entities and bodies in both species and face-sensitive regions in humans. Though unpredicted, we also observed neural representations of faces compared to inanimate objects, and dog compared to human bodies in dog olfactory regions. These findings shed light on the evolutionary foundations of human and dog social cognition and the predominant role of the temporal lobe.

Validation of a New Coil Array Tailored for Dog Functional Magnetic Resonance Imaging Studies.

Comparative neuroimaging allows for the identification of similarities and differences between species. It provides an important and promising avenue, to answer questions about the evolutionary origins of the brain´s organization, in terms of both structure and function. Dog functional magnetic resonance imaging (fMRI) has recently become one particularly promising and increasingly used approach to study brain function and coevolution. In dog neuroimaging, image acquisition has so far been mostly performed with coils originally developed for use in human MRI. Since such coils have been tailored to human anatomy, their sensitivity and data quality is likely not optimal for dog MRI. Therefore, we developed a multichannel receive coil (K9 coil, read "canine") tailored for high-resolution functional imaging in canines, optimized for dog cranial anatomy. In this paper we report structural (n = 9) as well as functional imaging data (resting-state, n = 6; simple visual paradigm, n = 9) collected with the K9 coil in comparison to reference data collected with a human knee coil. Our results show that the K9 coil significantly outperforms the human knee coil, improving the signal-to-noise ratio (SNR) across the imaging modalities. We noted increases of roughly 45% signal-to-noise in the structural and functional domain. In terms of translation to fMRI data collected in a visual flickering checkerboard paradigm, group-level analyses show that the K9 coil performs better than the knee coil as well. These findings demonstrate how hardware improvements may be instrumental in driving data quality, and thus, quality of imaging results, for dog-human comparative neuroimaging.

Socio-cognitive training impacts emotional and perceptual self-salience but not self-other distinction.

Training to inhibit imitative tendencies has been shown to reduce self-other interferences in both automatic imitation and perspective taking, suggesting that an enhancement of self-other distinction is transferrable from the motor to the cognitive domain. This study examined whether socio-cognitive training specifically enhances self-other distinction, or rather modulates self-salience, that is, the relative attentional priority of information pertaining to the self-perspective over information pertaining to the other person's perspective. Across two experiments, participants trained on one day to either imitate, inhibit imitation, inhibit control stimuli, or they were imitated. On the following day they completed a visuo-tactile affective perspective-taking paradigm measuring both self-other distinction and emotional self-salience, and a shape matching paradigm measuring perceptual self-salience. Results indicate no significant or consistent impact of training on self-other distinction performance, but reveal an increased emotional and perceptual self-salience following training to inhibit imitative tendencies. Together, these findings raise the question whether socio-cognitive training improves performance via enhanced self-other distinction, and invite to consider self-salience as a complementary angle to explain the past, present, and future findings on self-other distinction.

Exploring the dog-human relationship by combining fMRI, eye-tracking and behavioural measures.

Behavioural studies revealed that the dog-human relationship resembles the human mother-child bond, but the underlying mechanisms remain unclear. Here, we report the results of a multi-method approach combining fMRI (N = 17), eye-tracking (N = 15), and behavioural preference tests (N = 24) to explore the engagement of an attachment-like system in dogs seeing human faces. We presented morph videos of the caregiver, a familiar person, and a stranger showing either happy or angry facial expressions. Regardless of emotion, viewing the caregiver activated brain regions associated with emotion and attachment processing in humans. In contrast, the stranger elicited activation mainly in brain regions related to visual and motor processing, and the familiar person relatively weak activations overall. While the majority of happy stimuli led to increased activation of the caudate nucleus associated with reward processing, angry stimuli led to activations in limbic regions. Both the eye-tracking and preference test data supported the superior role of the caregiver's face and were in line with the findings from the fMRI experiment. While preliminary, these findings indicate that cutting across different levels, from brain to behaviour, can provide novel and converging insights into the engagement of the putative attachment system when dogs interact with humans.

Does Really One in Ten Believe Capital Punishment Exists in a Contemporary European Community Country? An Endorsed, Prereviewed, Preregistered Replication Study and Meta-Analysis.

BACKGROUND: Till et al. (2016) reported that in an Austrian sample approximately one in ten respondents incorrectly believed that Austria still practices, or recently practiced, the death penalty, and that there is a positive association between the amount of weekly television viewing and this gross misperception of the Austrian justice system. METHODS: An endorsed, prereviewed, preregistered close (N = 597) served to test the veracity of these reported effects. This was coupled with the conceptual extension part, which (a) investigated the potential influence of watching American crime series, (b) accounted for further possible confounds, and (c) tested the generalizability of the effect of television viewing to online streaming. RESULTS: Online survey data (N = 597) replicated the one-in-ten prevalence of incorrect answers with the 5-item death penalty questionnaire used in the original study, but not, when asking directly about Austria's death penalty practices (prevalence: 0.3%). Younger age, but not the amount of television viewing or online streaming, suggestibility, or preferred TV genre consistently predicted incorrect answers in the death penalty questionnaire. Incorrect answers were Mokken-scalable (i.e., formed a common scale, complying with a non-parametric item response model) and were highly consistent. In contrast to the replication study results, a small meta-analysis of all available evidence (three studies, including the present replication) suggested that the aggregate effect of television viewing nominally was significant, albeit small. CONCLUSION: The replication study yielded mixed results, which indicate the perception of a high prevalence of beliefs that there is capital punishment in a country without death penalty probably is due to a faultily designed questionnaire and thus a research artifact. Also, positive associations of television viewing with such beliefs likely are only small at best.

The multiple facets of empathy.

We discuss the definition of empathy provided by Kujala (2017) and argue that research in this field, in assigning the cognitive component of empathy only a secondary role, misses crucial information. Further knowledge about dogs' ability for higher cognitive processes helps (a) in interpreting results such as potential prosocial behavior in dogs and (b) sheds light on the question of whether abilities like perspective-taking and self-other distinction are uniquely human.