Andrew J. Bremner

Effects of action observation on corticospinal excitability: Muscle specificity, direction, and timing of the mirror response.

Many human behaviours and pathologies have been attributed to the putative mirror neuron system, a neural system that is active during both the observation and execution of actions. While there are now a very large number of papers on the mirror neuron system, variations in the methods and analyses employed by researchers mean that the basic characteristics of the mirror response are not clear. This review focuses on three important aspects of the mirror response, as measured by modulations in corticospinal excitability: (1) muscle specificity; (2) direction; and (3) timing of modulation. We focus mainly on electromyographic (EMG) data gathered following single-pulse transcranial magnetic stimulation (TMS), because this method provides precise information regarding these three aspects of the response. Data from paired-pulse TMS paradigms and peripheral nerve stimulation (PNS) are also considered when we discuss the possible mechanisms underlying the mirror response. In this systematic review of the literature, we examine the findings of 85 TMS and PNS studies of the human mirror response, and consider the limitations and advantages of the different methodological approaches these have adopted in relation to discrepancies between their findings. We conclude by proposing a testable model of how action observation modulates corticospinal excitability in humans. Specifically, we propose that action observation elicits an early, non-specific facilitation of corticospinal excitability (at around 90ms from action onset), followed by a later modulation of activity specific to the muscles involved in the observed action (from around 200ms). Testing this model will greatly advance our understanding of the mirror mechanism and provide a more stable grounding on which to base inferences about its role in human behaviour.

Infants lost in (peripersonal) space

A significant challenge in developing spatial representations for the control of action is one of multisensory integration. Specifically, we require an ability to efficiently integrate sensory information arriving from multiple modalities pertaining to the relationships between the acting limbs and the nearby external world (i.e. peripersonal space), across changes in body posture and limb position. Evidence concerning the early development of such spatial representations points towards the independent emergence of two distinct mechanisms of multisensory integration. The earlier-developing mechanism achieves spatial correspondence by representing body parts in their typical or default locations, and the later-developing mechanism does so by dynamically remapping the representation of the position of the limbs with respect to external space in response to changes in postural information arriving from proprioception and vision.

Exposure to an urban environment alters the local bias of a remote culture

There is substantial evidence that populations in the Western world exhibit a local bias compared to East Asian populations that is widely ascribed to a difference between individualistic and collectivist societies. However, we report that traditional Himba - a remote interdependent society - exhibit a strong local bias compared to both Japanese and British participants in the Ebbinghaus illusion and in a similarity-matching task with hierarchical figures. Critically, we measured the effect of exposure to an urban environment on local bias in the Himba. Even a brief exposure to an urban environment caused a shift in processing style: the local bias was reduced in traditional Himba who had visited a local town and even more reduced in urbanised Himba who had moved to that town on a permanent basis. We therefore propose that exposure to an urban environment contributes to the global bias found in Western and Japanese populations.

To eat or not to eat? Kinematics and muscle activity of reach-to-grasp movements are influenced by the action goal, but observers do not detect these differences

Recent evidence suggests that the mirror neuron system responds to the goals of actions, even when the end of the movement is hidden from view. To investigate whether this predictive ability might be based on the detection of early differences between actions with different outcomes, we used electromyography (EMG) and motion tracking to assess whether two actions with different goals (grasp to eat and grasp to place) differed from each other in their initial reaching phases. In a second experiment, we then tested whether observers could detect early differences and predict the outcome of these movements, based on seeing only part of the actions. Experiment 1 revealed early kinematic differences between the two movements, with grasp-to-eat movements characterised by an earlier peak acceleration, and different grasp position, compared to grasp-to-place movements. There were also significant differences in forearm muscle activity in the reaching phase of the two actions. The behavioural data arising from Experiments 2a and 2b indicated that observers are not able to predict whether an object is going to be brought to the mouth or placed until after the grasp has been completed. This suggests that the early kinematic differences are either not visible to observers, or that they are not used to predict the end-goals of actions. These data are discussed in the context of the mirror neuron system.

Children’s Responses to the Rubber-Hand Illusion Reveal Dissociable Pathways in Body Representation

The bodily self is constructed from multisensory information. However, little is known of the relation between multisensory development and the emerging sense of self. We investigated this question by measuring the strength of the rubber-hand illusion in young children (4 to 9 years old) and adults. Intermanual pointing showed that children were as sensitive as adults to visual-tactile synchrony cues for hand position, which indicates that a visual-tactile pathway to the bodily self matures by at least 4 years of age. However, regardless of synchrony cues, children’s perceived hand position was closer to the rubber hand than adults’ perceived hand position was. This indicates a second, later-maturing process based on visual-proprioceptive information. Furthermore, explicit feelings of embodiment were related only to the visual-tactile process. These findings demonstrate two dissociable processes underlying body representation in early life, and they call into question current models of body representation and ownership in adulthood.

Release of inattentional blindness by high working memory load: Elucidating the relationship between working memory and selective attention

An unexpected stimulus often remains unnoticed if attention is focused elsewhere. This inattentional blindness has been shown to be increased under conditions of high memory load. Here we show that increasing working memory load can also have the opposite effect of reducing inattentional blindness (i.e., improving stimulus detection) if stimulus detection is competing for attention with a concurrent visual task. Participants were required to judge which of two lines was the longer while holding in working memory either one digit (low load) or six digits (high load). An unexpected visual stimulus was presented once alongside the line judgment task. Detection of the unexpected stimulus was significantly improved under conditions of higher working memory load. This improvement in performance prompts the striking conclusion that an effect of cognitive load is to increase attentional spread, thereby enhancing our ability to detect perceptual stimuli to which we would normally be inattentionally blind under less taxing cognitive conditions. We discuss the implications of these findings for our understanding of the relationship between working memory and selective attention.

Soft skills in higher education: importance and improvement ratings as a function of individual differences and academic performance

Three UK studies on the relationship between a purpose‐built instrument to assess the importance and development of 15 ‘soft skills’ are reported. Study 1 (N = 444) identified strong latent components underlying these soft skills, such that differences between‐skills were over‐shadowed by differences between‐students. Importance and improving ratings on these skills predicted academic performance and accounted for the effects of personality on academic performance. Study 2 replicated the structure of the soft skills inventory and associations with academic performance in a larger sample (N = 1309). Examination of mean differences across faculties (humanities, life sciences, hard sciences) revealed higher soft skills ratings in ‘softer’ courses. Study 3 (N = 87) incorporated an IQ measure, which was found to be negatively related to importance ratings on soft skills. Results highlight the cohesive structure of beliefs concerning various non‐academic skills and their significant links to educationally relevant individual differences. Theoretical, methodological and applied implications are considered.

Bodily illusions in young children: developmental change in visual and proprioceptive contributions to perceived hand position.

We examined the visual capture of perceived hand position in forty-five 5- to 7-year-olds and in fifteen young adults, using a mirror illusion task. In this task, participants see their left hand on both the left and right (by virtue of a mirror placed at the midline facing the left arm, and obscuring the right). The accuracy of participants’ reaching was measured when proprioceptive and visual cues to the location of the right arm were put into conflict (by placing the arms at different distances from the mirror), and also when only proprioceptive information was available (i.e., when the mirror was covered). Children in all age-groups (and adults) made reaching errors in the mirror condition in accordance with the visually-specified illusory starting position of their hand indicating a visual capture of perceived hand position. Data analysis indicated that visual capture increased substantially up until 6 years of age. These findings are interpreted with respect to the development of the visual guidance of action in early childhood.

Skin and Touch

The skin contains a wide variety of receptors that give rise to our experience of touch. Tactile experience depends on attention, and is typically the result of multisensory integration. Many of our tactile experiences are the result of the integration of inputs from different classes of receptors – known as touch blends. The sense of touch is an important albeit understudied sense. While the bandwidth of the sense of touch is relatively low, stimulating the skin can nevertheless convey emotional benefits.

Do Local and Global Perceptual Biases Tell Us Anything About Local and Global Selective Attention

Local, as opposed to global, perceptual bias has been linked to a lesser ability to attend globally. We examined this proposed link in Himba observers, members of a remote Namibian population who have demonstrated a strong local bias compared with British observers. If local perceptual bias is related to a lesser ability to attend globally, Himba observers, relative to British observers, should be less distracted by global information when performing a local-selection task but more distracted by local information when performing a global-selection task. However, Himba observers performed better than British observers did on both a local-selection task and a global-selection task (both of which used local/global hierarchical figures as stimuli), which suggests that they possessed greater control over attentional selection in response to task demands. We conclude that local and global perceptual biases must be distinguished from local and global selective attention.