Assessing acute effects of two motor-cognitive training modalities on cognitive functions, postural control, and gait stability in older adults: a randomized crossover study.

Background: The process of aging often accompanies a decline in cognitive function, postural control, and gait stability, consequently increasing the susceptibility to falls among older individuals. In response to these challenges, motor-cognitive training has emerged as a potential intervention to mitigate age-related declines. Objective: This study aims to assess the acute effects of two distinct motor-cognitive training modalities, treadmill dual-task training (TMDT) and interactive motor-cognitive training (IMCT), on cognitive function, postural control, walking ability, and dual-task performance in the elderly population. Method: In this randomized crossover study, 35 healthy elderly individuals (aged 60-75) participated in three acute training sessions involving TMDT, IMCT, and a control reading condition. Assessments of executive function, postural control, gait performance, and cognitive accuracy were conducted both before and after each session. Results: Both TMDT and IMCT improved executive functions. Notably, IMCT resulted in a significant enhancement in correct response rates and a reduction in reaction times in the Stroop task (p < 0.05) compared to TMDT and the control condition. IMCT also led to an increase in dual-task gait speed (p < 0.001) and showed a trend towards improved cognitive accuracy (p = 0.07). Conversely, TMDT increased postural sway with eyes open (p = 0.013), indicating a potential detriment to postural control. Conclusion: The findings suggest that IMCT holds greater immediate efficacy in enhancing cognitive function and gait stability among older adults compared to TMDT, with a lesser adverse impact on postural control. This underscores the potential of IMCT as a preferred approach for mitigating fall risk and enhancing both cognitive and physical functions in the elderly population.

Motor cognition in plants: from thought to real experiments.

Motor cognition involves the process of planning and executing goal-directed movements and recognizing, anticipating, and interpreting others' actions. Motor cognitive functions are generally associated with the presence of a brain and are ascribed only to humans and other animal species. A growing body of evidence suggests that aneural organisms, like climbing plants, exhibit behaviors driven by the intention to achieve goals, challenging our understanding of cognition. Here, we propose an inclusive perspective under motor cognition to explain climbing plants' behavior. We will first review our empirical research based on kinematical analysis to understand movement in pea plants. Then, we situate this empirical research within the current theoretical debate aimed at extending the principles of cognition to aneural organisms. A novel comparative perspective that considers the perception-action cycle, involving transforming perceived environmental elements into intended movement patterns, is provided.

Life-long music and dance relationships inform impressions of music- and dance-based movement therapies in individuals with and without mild cognitive impairment.

Background: No effective therapies exist to prevent degeneration from Mild Cognitive Impairment (MCI) to Alzheimer's disease. Therapies integrating music and/or dance are promising as effective, non-pharmacological options to mitigate cognitive decline. Objective: To deepen our understanding of individuals' relationships (i.e., histories, experiences and attitudes) with music and dance that are not often incorporated into music- and dance-based therapeutic design, yet may affect therapeutic outcomes. Methods: Eleven older adults with MCI and five of their care partners/ spouses participated (4M/12F; Black: n=4, White: n=10, Hispanic/ Latino: n=2; Age: 71.4±9.6). We conducted focus groups and administered questionnaires that captured aspects of participants' music and dance relationships. We extracted emergent themes from four major topics, including: (1) experience and history, (2) enjoyment and preferences, (3) confidence and barriers, and (4) impressions of music and dance as therapeutic tools. Results: Thematic analysis revealed participants' positive impressions of music and dance as potential therapeutic tools, citing perceived neuropsychological, emotional, and physical benefits. Participants viewed music and dance as integral to their lives, histories, and identities within a culture, family, and/ or community. Participants also identified lifelong engagement barriers that, in conjunction with negative feedback, instilled persistent low self-efficacy regarding dancing and active music engagement. Questionnaires verified individuals' moderately-strong music and dance relationships, strongest in passive forms of music engagement (e.g., listening). Conclusions: Our findings support that individuals' music and dance relationships and the associated perceptions toward music and dance therapy may be valuable considerations in enhancing therapy efficacy, participant engagement and satisfaction for individuals with MCI.

Object-oriented hand dexterity and grasping abilities, from the animal quarters to the neurosurgical OR: a systematic review of the underlying neural correlates in non-human, human primate and recent findings in awake brain surgery.

Introduction: The sensorimotor integrations subserving object-oriented manipulative actions have been extensively investigated in non-human primates via direct approaches, as intracortical micro-stimulation (ICMS), cytoarchitectonic analysis and anatomical tracers. However, the understanding of the mechanisms underlying complex motor behaviors is yet to be fully integrated in brain mapping paradigms and the consistency of these findings with intraoperative data obtained during awake neurosurgical procedures for brain tumor removal is still largely unexplored. Accordingly, there is a paucity of systematic studies reviewing the cross-species analogies in neural activities during object-oriented hand motor tasks in primates and investigating the concordance with intraoperative findings during brain mapping. The current systematic review was designed to summarize the cortical and subcortical neural correlates of object-oriented fine hand actions, as revealed by fMRI and PET studies, in non-human and human primates and how those were translated into neurosurgical studies testing dexterous hand-movements during intraoperative brain mapping. Methods: A systematic literature review was conducted following the PRISMA guidelines. PubMed, EMBASE and Web of Science databases were searched. Original articles were included if they: (1) investigated cortical activation sites on fMRI and/or PET during grasping task; (2) included humans or non-human primates. A second query was designed on the databases above to collect studies reporting motor, hand manipulation and dexterity tasks for intraoperative brain mapping in patients undergoing awake brain surgery for any condition. Due to the heterogeneity in neurosurgical applications, a qualitative synthesis was deemed more appropriate. Results: We provided an updated overview of the current state of the art in translational neuroscience about the extended frontoparietal grasping-praxis network with a specific focus on the comparative functioning in non-human primates, healthy humans and how the latter knowledge has been implemented in the neurosurgical operating room during brain tumor resection. Discussion: The anatomical and functional correlates we reviewed confirmed the evolutionary continuum from monkeys to humans, allowing a cautious but practical adoption of such evidence in intraoperative brain mapping protocols. Integrating the previous results in the surgical practice helps preserve complex motor abilities, prevent long-term disability and poor quality of life and allow the maximal safe resection of intrinsic brain tumors.

The parietal architecture binding cognition to sensorimotor integration: a multimodal causal study.

Despite human's praxis abilities are unique among primates, comparative observations suggest that these cognitive motor skills could have emerged from exploitation and adaptation of phylogenetically older building blocks, namely the parieto-frontal networks subserving prehension and manipulation. Within this framework, investigating to which extent praxis and prehension-manipulation overlap and diverge within parieto-frontal circuits could help in understanding how human cognition shapes hand actions. This issue has never been investigated by combining lesion mapping and direct electrophysiological approaches in neurosurgical patients. To this purpose, 79 right-handed left-brain tumour patient candidates for awake neurosurgery were selected based on inclusion criteria. First, a lesion mapping was performed in the early postoperative phase to localize the regions associated with an impairment in praxis (imitation of meaningless and meaningful intransitive gestures) and visuo-guided prehension (reaching-to-grasping) abilities. Then, lesion results were anatomically matched with intraoperatively identified cortical and white matter regions, whose direct electrical stimulation impaired the Hand Manipulation Task. The lesion mapping analysis showed that prehension and praxis impairments occurring in the early postoperative phase were associated with specific parietal sectors. Dorso-mesial parietal resections, including the superior parietal lobe and precuneus, affected prehension performance, while resections involving rostral intraparietal and inferior parietal areas affected praxis abilities (covariate clusters, 5000 permutations, cluster-level family-wise error correction P < 0.05). The dorsal bank of the rostral intraparietal sulcus was associated with both prehension and praxis (overlap of non-covariate clusters). Within praxis results, while resection involving inferior parietal areas affected mainly the imitation of meaningful gestures, resection involving intraparietal areas affected both meaningless and meaningful gesture imitation. In parallel, the intraoperative electrical stimulation of the rostral intraparietal and the adjacent inferior parietal lobe with their surrounding white matter during the hand manipulation task evoked different motor impairments, i.e. the arrest and clumsy patterns, respectively. When integrating lesion mapping and intraoperative stimulation results, it emerges that imitation of praxis gestures first depends on the integrity of parietal areas within the dorso-ventral stream. Among these areas, the rostral intraparietal and the inferior parietal area play distinct roles in praxis and sensorimotor process controlling manipulation. Due to its visuo-motor 'attitude', the rostral intraparietal sulcus, putative human homologue of monkey anterior intraparietal, might enable the visuo-motor conversion of the observed gesture (direct pathway). Moreover, its functional interaction with the adjacent, phylogenetic more recent, inferior parietal areas might contribute to integrate the semantic-conceptual knowledge (indirect pathway) within the sensorimotor workflow, contributing to the cognitive upgrade of hand actions.

Brain plasticity underlying sleep-dependent motor consolidation after motor imagery.

Motor imagery can, similarly to physical practice, improve motor performance through experience-based plasticity. Using magnetoencephalography, we investigated changes in brain activity associated with offline consolidation of motor sequence learning through physical practice or motor imagery. After an initial training session with either physical practice or motor imagery, participants underwent overnight consolidation. As control condition, participants underwent wake-related consolidation after training with motor imagery. Behavioral analyses revealed that overnight consolidation of motor learning through motor imagery outperformed wake-related consolidation (95% CI [0.02, 0.07], P < 0.001, RP2 = 0.05). As regions of interest, we selected the generators of event-related synchronization/desynchronization of alpha (8-12 Hz) and beta (15-30 Hz) oscillations, which predicted the level of performance on the motor sequence. This yielded a primary sensorimotor-premotor network for alpha oscillations and a cortico-cerebellar network for beta oscillations. The alpha network exhibited increased neural desynchronization after overnight consolidation compared to wake-related consolidation. By contrast, the beta network exhibited an increase in neural synchronization after wake-related consolidation compared to overnight consolidation. We provide the first evidence of parallel brain plasticity underlying behavioral changes associated with sleep-dependent consolidation of motor skill learning through motor imagery and physical practice.

Visuo-motor interference is modulated by task interactivity: A kinematic study.

Extensive evidence shows that action observation can influence action execution, a phenomenon often referred to as visuo-motor interference. Little is known about whether this effect can be modulated by the type of interaction agents are involved in, as different studies show conflicting results. In the present study, we aimed at shedding light on this question by recording and analyzing the kinematic unfolding of reach-to-grasp movements performed in interactive and noninteractive settings. Using a machine learning approach, we investigated whether the extent of visuo-motor interference would be enhanced or reduced in two different joint action settings compared with a noninteractive one. Our results reveal that the detrimental effect of visuo-motor interference is reduced when the action performed by the partner is relevant to achieve a common goal, regardless of whether this goal requires to produce a concrete sensory outcome in the environment (joint outcome condition) or only a joint movement configuration (joint movement condition). These findings support the idea that during joint actions we form dyadic motor plans, in which both our own and our partner's actions are represented in predictive terms and in light of the common goal to be achieved. The formation of a dyadic motor plan might allow agents to shift from the automatic simulation of an observed action to the active prediction of the consequences of a partner's action. Overall, our results demonstrate the unavoidable impact of others' action on our motor behavior in social contexts, and how strongly this effect can be modulated by task interactivity.

Does aging amplify the rule-based efficiency effect in action selection?

When it comes to the selection of adequate movements, people may apply varying strategies. Explicit if-then rules, compared to implicit prospective action planning, can facilitate action selection in young healthy adults. But aging alters cognitive processes. It is unknown whether older adults may similarly, profit from a rule-based approach to action selection. To investigate the potential effects of aging, the Rule/Plan Motor Cognition (RPMC) paradigm was applied to three different age groups between 31 and 90 years of age. Participants selected grips either instructed by a rule or by prospective planning. As a function of age, we found a general increase in a strategy-specific advantage as quantified by the difference in reaction time between plan- and rule-based action selection. However, in older age groups, these differences went in both directions: some participants initiated rule-based action selection faster, while for others, plan-based action selection seemed more efficient. The decomposition of reaction times into speed of the decision process, action encoding, and response caution components suggests that rule-based action selection may reduce action encoding demands in all age groups. There appears a tendency for the younger and middle age groups to have a speed advantage in the rule task when it comes to information accumulation for action selection. Thus, one influential factor determining the robustness of the rule-based efficiency effect across the lifespan may be presented by the reduced speed of information uptake. Future studies need to further specify the role of these parameters for efficient action selection.

Establishing conversational engagement and being effective: The role of body movement in mediated communication.

A model for investigating the effects of body movement on conversational effectiveness in computer-mediated communication (CMC) is developed based on theories of motor cognition and embodiment. Movement is relevant to a wide range of CMC settings, including remote interviews, court testimonials, instructing, medical consultation, and socializing. The present work allows for a consideration of different forms of motoric activation, including gesturing and full-body motion, in mediated conversational settings and the derivation of a range of testable hypothesis. Motor cognition and embodiment provide an account of how speaker and listener become subject to the consequences of the muscular activation patterns that come with body movement. While movement supports internal elaboration, thus helping the speaker in formulating messages, it also has direct effects on the listener through behavioral synchrony and motor contagion. The effects of movement in CMC environments depend on two general characteristics: the level of visibility of movement and the extent to which the technology facilitates or inhibits movement. Available channels, set-up of technology, and further customization therefore determine whether movement can fulfil its internal functions (relevant to cognitive-affective elaboration of what is being said by the speaker) and its external functions (relevant to what is being perceived by and activated within the listener). Several indicators of conversational effectiveness are identified that serve as outcome variables. This MCEE model is intended to help users, developers and service provides to make CMC more engaging and more meaningful.

Can they touch? A novel mental motor imagery task for the assessment of back pain.

Introduction: As motor imagery is informed by the anticipated sensory consequences of action, including pain, we reasoned that motor imagery could provide a useful indicator of chronic back pain. We tested the hypothesis that mental motor imagery regarding body movements can provide a reliable assessment of low back pain. Methods: Eighty-five subjects with back pain and forty-five age-matched controls were shown two names of body parts and asked to indicate if they could imagine moving so that the named body parts touched. Three types of imagined movements were interrogated: movements of arms, movements of legs and movements requiring flexion and/or rotation of the low back. Results: Accuracy and reaction times were measured. Subjects with back pain were less likely to indicate that they could touch body parts than age-matched controls. The effect was observed only for those movements that required movement of the low back or legs, suggesting that the effect was not attributable to task difficulty or non-specific effects. There was an effect of pain severity. Compared to subjects with mild pain, subjects with severe pain were significantly less likely to indicate that they could move so that named body parts touched. There was a correlation between pain ratings and impaired performance for stimuli that involved the lower but not upper body. Discussion: As the Can They Touch task is quick, easy to administer and does not require an explicit judgment of pain severity, it may provide useful information to supplement the assessment of subjects with chronic pain.

Mapping relational links between motor imagery, action observation, action-related language, and action execution.

Actions can be physically executed, observed, imagined, or simply thought about. Unifying mental processes, such as simulation, emulation, or predictive processing, are thought to underlie different action types, whether they are mental states, as in the case of motor imagery and action observation, or involve physical execution. While overlapping brain activity is typically observed across different actions which indicates commonalities, research interest is also concerned with investigating the distinct functional components of these action types. Unfortunately, untangling subtleties associated with the neurocognitive bases of different action types is a complex endeavour due to the high dimensional nature of their neural substrate (e.g., any action process is likely to activate multiple brain regions thereby having multiple dimensions to consider when comparing across them). This has impeded progress in action-related theorising and application. The present study addresses this challenge by using the novel approach of multidimensional modeling to reduce the high-dimensional neural substrate of four action-related behaviours (motor imagery, action observation, action-related language, and action execution), find the least number of dimensions that distinguish or relate these action types, and characterise their neurocognitive relational links. Data for the model comprised brain activations for action types from whole-brain analyses reported in 53 published articles. Eighty-two dimensions (i.e., 82 brain regions) for the action types were reduced to a three-dimensional model, that mapped action types in ordination space where the greater the distance between the action types, the more dissimilar they are. A series of one-way ANOVAs and post-hoc comparisons performed on the mean coordinates for each action type in the model showed that across all action types, action execution and concurrent action observation (AO)-motor imagery (MI) were most neurocognitively similar, while action execution and AO were most dissimilar. Most action types were similar on at least one neurocognitive dimension, the exception to this being action-related language. The import of the findings are discussed in terms of future research and implications for application.

The pantomime of mental rotation: Left-handers are less lateralized.

INTRODUCTION: The conceptualization of skilled hand movements (praxis) may be grounded in hemispherically specialized functions. However, a left-hemispherical advantage of (tool-use) pantomime gestures and a right-hemispherical advantage of spatial gestures may be more prominent in right-handed than left-handed individuals. We therefore investigated the hypothesis that right-handed but not left-handed individuals show a superiority of the left hemisphere (/right-hand preference) for the execution of pantomime (rotation of an object) gestures as well as a right-hemispherical superiority (/left-hand preference) for gestures that depict spatial information (/positioning of an object). METHODS: 20 right- and 20 left-handed participants were asked in two experiments to demonstrate with their two hands how to move tachistoscopically (in the left (LVF) or right visual hemifields (RVF)) presented geometric objects of different rotations into an identical final position. Two independent blind raters evaluated the videotaped hand gestures employing the Neuropsychological Gesture (NEUROGES) Coding System. RESULTS: In contrast to left-handed individuals, right-handed individuals present increased pantomime - rotation gestures with the right hand and pantomime - position gestures with the left hand during stimuli presentation in either visual field. Left-handers showed significantly increased left-hand pantomime - rotation gestures during stimulus presentation within the LVF (only). DISCUSSION: Right-handed individuals increase their pantomime - rotation gestures with the right hand to depict motion but use their left hand for pantomime - position gestures to describe spatial relations of the objects. Left-handers do not show a clear lateralization of the right and left hand with regards to either handedness or hemispherically lateralized motor functions. The hemispherical lateralization of praxis functions is therefore more pronounced in right-handed than left-handed individuals.

Do graspable objects always leave a motor signature? A study on memory traces.

Several studies have reported the existence of reciprocal interactions between the type of motor activity physically performed on objects and the conceptual knowledge that is retained of them. Whether covert motor activity plays a similar effect is less clear. Certainly, objects are strong triggers for actions, and motor components can make the associated concepts more memorable. However, addition of an action-related memory trace may not always be automatic and could rather depend on 'how' objects are encountered. To test this hypothesis, we compared memory for objects that passive observers experienced as verbal labels (the word describing them), visual images (color photographs) and actions (pantomimes of object use). We predicted that the more direct the involvement of action-related representations the more effective would be the addition of a motor code to the experience and the more accurate would be the recall. Results showed that memory for objects presented as words i.e., a format that might only indirectly prime the sensorimotor system, was generally less accurate compared to memory for objects presented as photographs or pantomimes, which are more likely to directly elicit motor simulation processes. In addition, free recall of objects experienced as pantomimes was more accurate when these items afforded actions performed towards one's body than actions directed away from the body. We propose that covert motor activity can contribute to objects' memory, but the beneficial addition of a motor code to the experience is not necessarily automatic. An advantage is more likely to emerge when the observer is induced to take a first-person stance during the encoding phase, as may happen for objects affording actions directed towards the body, which obviously carry more relevance for the actor.

The unexplored link between aesthetic perception and creativity: A theory-driven meta-analysis of fMRI studies in the visual domain.

Creative production (related to art-making) and aesthetic appreciation (related to art-viewing) are inherently linked in visual arts, but their relationship has never been explored explicitly in cognitive neuroscience, nor the nature of such connection. The available literature suggests two cognitive processes as possible foundations of these two experiences: motor simulation or inhibitory control. In a meta-analysis of fMRI studies, we addressed this issue: we investigated whether there are shared neurofunctional underpinnings behind aesthetic and creative experiences in the visual domain; further, we examined whether any shared brain activation may reflect either motor simulation or inhibitory processes. A conjunction analysis revealed a common involvement of the pre-SMA in both classes of studies, a brain region, if anything, more concerned with top-down inhibitory motor and volitional cognitive control rather than bottom-up motor simulation. In the art-viewing domain, this finding was primarily driven by figurative rather than abstract art. The methodological limitations in the available literature are discussed together with possible new ways to expand the existing findings.

Action goals and the praxis network: an fMRI study.

The praxis representation network (PRN) of the left cerebral hemisphere is typically linked to the control of functional interactions with familiar tools. Surprisingly, little is known about the PRN engagement in planning and execution of tool-directed actions motivated by non-functional but purposeful action goals. Here we used functional neuroimaging to perform both univariate and multi-voxel pattern analyses (MVPA) in 20 right-handed participants who planned and later executed, with their dominant and non-dominant hands, disparate grasps of tools for different goals, including: (1) planning simple vs. demanding functional grasps of conveniently vs. inconveniently oriented tools with an intention to immediately use them, (2) planning simple-but non-functional-grasps of inconveniently oriented tools with a goal to pass them to a different person, (3) planning reaching movements directed at such tools with an intention to move/push them with the back of the hand, and (4) pantomimed execution of the earlier planned tasks. While PRN contributed to the studied interactions with tools, the engagement of its critical nodes, and/or complementary right hemisphere processing, was differently modulated by task type. E.g., planning non-functional/structural grasp-to-pass movements of inconveniently oriented tools, regardless of the hand, invoked the left parietal and prefrontal nodes significantly more than simple, non-demanding functional grasps. MVPA corroborated decoding capabilities of critical PRN areas and some of their right hemisphere counterparts. Our findings shed new lights on how performance of disparate action goals influences the extraction of object affordances, and how or to what extent it modulates the neural activity within the parieto-frontal brain networks.

Social Touch Somatotopically Affects Mental Body Representations.

In pre-Covid days, many daily actions such as hand shaking or cheek kissing implied physical contact between our body and that of other people. With respect to touching an inanimate object (objectual touch), touching a person (social touch) concerns not only touching a human body, but also that this body belongs to a living person. This fundamental difference also may affect the way we figure our own movements and perceptions or, in other words, how we mentally represent our own body. To test this hypothesis, we asked 30 neurotypical participants to perform mental rotation of images representing hands, full bodies, and feet (an active cognitive task able to activate body representations without need of moving) in two tactile conditions: holding (one in each hand) either the thumbs of another person (social touch) or two plastic cylinders (objectual touch) of about the same circumference and size. Results showed that only mental rotation of hand images was affected by varying the tactile conditions, in that participants were faster during social than objectual touch. This suggests that the nature of hand-related tactile input (social or objectual touch) influences local (hand) and not global (body) mental representations of the body, and in a very somatotopic manner (hands but not feet). We interpret these findings with reference to the differentiation between sensorimotor (body schema) and visuospatial (body image) dynamics in the mental representation of our body. The present study shows that external social factors can affect the internal mental representations of one's own body.

Translation, Cross-Cultural Adaptation, and Validation of a Dutch Version of the Actions and Feelings Questionnaire in Autistic and Neurotypical Adults.

The actions and feelings questionnaire (AFQ) provides a short, self-report measure of how well someone uses and understands visual communicative signals such as gestures. The objective of this study was to translate and cross-culturally adapt the AFQ into Dutch (AFQ-NL) and validate this new version in neurotypical and autistic populations. Translation and adaptation of the AFQ consisted of forward translation, synthesis, back translation, and expert review. In order to validate the AFQ-NL, we assessed convergent and divergent validity. We additionally assessed internal consistency using Cronbach's alpha. Validation and reliability outcomes were all satisfactory. The AFQ-NL is a valid adaptation that can be used for both autistic and neurotypical populations in the Netherlands.

Qualitative Evaluation Informs Understanding of Motor Cognition and Therapies in Older Adults with Mild Cognitive Impairment.

BACKGROUND: 10%to 20%of Americans aged 65 and older have mild cognitive impairment (MCI) with 10%progressing to Alzheimer's disease (AD) each year. Underserved groups, including African Americans (AAs), are among the most vulnerable to MCI and AD. Although evidence continues to amass, the benefits of exercise and movement for AD is still understudied in AD. OBJECTIVE: Understanding the attitudes, perceptions, and beliefs about motor-cognitive integration and examining the physical activity of a sample of predominantly Black women community members with self-reported memory problems will allow improved recruitment and refinement of multimodal interventions designed to improve motor-cognitive and cognitive function. METHODS: We conducted focus groups with older adults who reported subjective memory complaints (n = 15; Black: n = 12, White: n = 3, mean age 71.7±5.8). RESULTS: Findings from thematic analysis showed most participants knew of benefits of exercise. However, most participants reported not getting adequate exercise due to factors such as pain, increased responsibilities, and fear of injury. Despite barriers, participants expressed enthusiasm for multimodal interventions designed to target body and brain health and provided several suggestions to improve or enhance the proposed interventions. CONCLUSION: Results provide useful insights regarding improving participation among historically under-represented groups in clinical movement-based research. Participants' discussion focused primarily on the way motor-cognitive integration prevents falls, maintains memory, and provides a social benefit. The reported perceived benefits and limitations of exercise, as this population understands it, can help researchers and physicians better engage the community for lifestyle changes that will support greater motor-cognitive health.

What modulates the Mirror Neuron System during action observation?: Multiple factors involving the action, the actor, the observer, the relationship between actor and observer, and the context.

Seeing an agent perform an action typically triggers a motor simulation of that action in the observer's Mirror Neuron System (MNS). Over the past few years, it has become increasingly clear that during action observation the patterns and strengths of responses in the MNS are modulated by multiple factors. The first aim of this paper is therefore to provide the most comprehensive survey to date of these factors. To that end, 22 distinct factors are described, broken down into the following sets: six involving the action; two involving the actor; nine involving the observer; four involving the relationship between actor and observer; and one involving the context. The second aim is to consider the implications of these findings for four prominent theoretical models of the MNS: the Direct Matching Model; the Predictive Coding Model; the Value-Driven Model; and the Associative Model. These assessments suggest that although each model is supported by a wide range of findings, each one is also challenged by other findings and relatively unaffected by still others. Hence, there is now a pressing need for a richer, more inclusive model that is better able to account for all of the modulatory factors that have been identified so far.