Perception, action, and the body model.

In 1992, Goodale and Milner proposed to study the visual system based on function, thus dissociating vision for perception (ventral stream) and vision for action (dorsal stream). This became known as the Perception and Action model (PAM). Following the PAM in the visual system, a somatosensory PAM was proposed including a body representation for perception and a separate for action. This review explores the body model of the hand and how it relates to the PAM. The body model refers to the internal representation of the body that is responsible for position sense. Previous research has shown that the representation of the hand features systematic distortions: an overestimation of hand width and an underestimation of finger length. These distortions have been reported using different paradigms, different body parts, and in various settings. Thus, body model distortions appear to be a characteristic of human body representation. If the body model of the hand is distorted, how can actions like reaching and grasping be accurate? We review evidence that body model distortions may in fact provide a functional benefit to our actions, that cortical maps in the somatosensory and motor cortices reflect these distortions, and that actions rely on a distorted body model. We argue that the body model is a product of both the ventral and dorsal somatosensory streams. Further, we suggest that the body model is an example of the inextricable link between the two streams.

The distorted hand: systematic but 'independent' distortions in both explicit and implicit hand representations in young female adults.

It has long been assumed that an accurate representation of the size and shape of one's body is necessary to successfully interact with the environment. Previous research has shown accurate representations when healthy participants make overt judgments (i.e. explicit) about the size of their bodies. However, when body size is judged implicitly, studies have shown systematic distortions. One suggestion for these differences, is that explicit and implicit representations are informed by different sensory modalities. Explicit representations rely on vision whereas implicit representations are informed by haptics. We designed an experiment to investigate if explicit representations that are informed by haptics are more like implicit representation featuring systematic distortions. We asked female participants to estimate the size of their fingers and hands in three different tasks: an explicit-haptic, an implicit, and an explicit-vision task. The results showed that all three representations were distorted and furthermore, the distortions for each representation were different from one another. These results suggest that inaccurate finger and hand length are a stereotypical feature of body representation that is present in both visual and haptic domains. We discuss the results in relation to theories of body representation.

Differential effects of aging on spatial abilities.

Visuospatial functions are particularly vulnerable to the aging process. Decline of these processes can seriously affect an individual's functional independence and quality of life. Effectively assessing the spatial abilities of older adults is, therefore, crucial for identifying strategies to maintain cognitive functioning. The purpose of the present study was to use ecological tasks more comparable to activities of daily living to assess spatial ability in older adults. Three hands-on tasks (a visual search task, a low- and a high-mental rotation demand tasks) and a version of the well-known paper-based mental rotation of figures test (Shepard and Metzler, Science 171(3972):701-703, 1971) were given to 60-79-year-old female and male participants. The hands-on tasks required participants to locate, manipulate, and arrange real objects (i.e., toy bricks) in space. Age had a negative impact on visual search but not on mental rotation ability. Male participants outperformed females in the mental rotation tasks, but a trend for the opposite (better performance by females) was found for the visual search task. The results suggest that spatial abilities are not a monolithic construct and that sub-categories of this construct are affected by age and by sex differently. While visual search function is susceptible to decline during old age, mental rotation ability is not. In addition, unlike the paper-based test, the hands-on tasks were found to be age-appropriate with a feasible level of difficulty for all participants. The hands-on tasks may be more appealing as a tool to evaluate, maintain, and/or enhance spatial function in older adults.

Growing into your hand: the developmental trajectory of the body model.

We rely on accurate body representations to successfully interact with the environment. As adults, we rely on many years of experience with a body that has stayed relatively the same size. Children, however, go through periods of rapid growth and whether or not their body representation matches this physical growth is unknown. To address this question, we examined the developmental trajectory of the body model of the hand. The body model is the representation of our bodies that underlies position sense. We recruited a group of children (8-16 years) and a control group of young adults (18-26 years) and asked them to complete the body model task. In this task, participants estimated the location of ten different landmarks (the tips and metacarpophalangeal joints of each of their five fingers). The position (XY location) of each estimate was tracked using an Optotrak camera. From the XY locations we derived hand width and finger length. Not surprisingly, children's physical hand width and finger length were smaller than adults but remarkably, the body model, was similar for both groups. This result indicates that children overestimate hand size and suggests that the body model is ahead of physical growth. This result contradicts the notion that body representation lags physical growth during puberty, accounting for the clumsy motor behaviour characteristic of teens. We discuss the results in relation to the different taxonomies of body representation and how an enlarged representation of the hand during childhood may influence action.

One brick at a time: Building a developmental profile of spatial abilities.

Spatial abilities are not only fundamental for activities of daily living, but they are also markers of academic and professional success. It has remained a challenge, however, to understand their development across childhood and adolescence, partly because of the lack of spatial tasks that are appropriate across age groups. For example, the well-established paper-based mental rotation test (MRT) has been shown to be too difficult for children. In the current study, we tested girls and boys in three age groups: younger children (5-8-years-old), older children (9-12), and adolescents (13-17) on a hands-on visuospatial task using toy bricks: the brick-building task (BBT). Children completed a low- and a high-mental rotation demand (LMR and HMR) version of the BBT and the paper-based MRT. Correlations were found between all tasks. Children, especially females, found the HMR more challenging than the LMR condition, but all children successfully completed the BBT. In contrast, the MRT was too difficult for the younger children to solve performing at chance. Given this result and that the BBT is a game-like, 3D, interactive task, the BBT could be used to examine the biological and/or environmental factors that affect the early development of visuospatial abilities.

Which hand knows the "right" word? What hand selection reveals about vocabulary in pre-and school-aged children.

Research has shown that infants with increased right-hand selection for their first gestures perform better at an array of language tasks when they are tested later as toddlers. There is a smaller body of literature which focuses on preschoolers and how their right-handed movements relate to their speech and vocabulary development. Some research has established a connection between right-hand preference for grasping and speech production ability in preschool children, but the link to gestures is relatively unexplored in this age group. We investigated if lateralized gestures (pointing) are related to measures of language development (vocabulary) in a preschool-age sample. Specifically, typically developing children (aged 3-6) completed the Peabody Picture Vocabulary Test (PPVT) to assess receptive language. We recorded their hand preference for pointing during the PPVT and the incidence of mistakes (pointing to the wrong picture). Despite the length of the test, children were more likely to select a correct response with their right hand. This result suggests a relationship between lateralized communicative gestures (pointing) and receptive language. This study provides evidence for an intimate relationship between right-handed manual movement and language development. Implications of this finding include developing simple fine-motor tasks to detect and/or ameliorate delayed language development.

Promoting Executive Function Skills in Preschoolers Using a Play-Based Program.

In recent years, play has been shown to be a powerful means to enhance learning and brain development. It is also known that through play children enhance their executive function (EF) skills. Furthermore, well-developed EF in preschoolers has been shown to be an important predictor for later academic and life success. Armed with this information a program, Building Brains and Futures (BBF), for developing EF through play was designed for 3-5-year-old. The program consisted of 10 simple, fun, and interactive games selected to enhance various facets of EF. The 10 games included were: dimensional change card sort, lips and ears, block building, musical freeze, opposites, pretend play, red light/green light, shared project, Simon says, and wait for it. The program was implemented with a group of children shown to have challenges with respect to kindergarten readiness. The approach was first, to build adult capability by sharing knowledge of brain development, EF, and the importance of play with educators, caregivers, and parents. Second, to build skills in delivering the program in the school setting. Children engaged with the program of games for a minimum of 6 weeks. Their performance on a battery of direct measures of EF, language, and motor skills, were recorded before and after the program. The results showed improvement in all three domains. In addition, adopters of the BBF program reported it was easily and successfully integrated into their existing preschool curricula. The importance of intentional adult directed play in building developmental learning, including EF, is discussed.

An evaluation of visuospatial skills using hands-on tasks.

Several tests of mental rotation ability have been used to investigate its development and the origins of sex differences. One of the most used tests is the mental rotation test (MRT) by Vandenberg and Kuse. A limitation of the MRT is that it is a pen-and-paper test with 2D images of 3D objects. This is a challenge to the ecological validity of the MRT because mental rotation typically involves physical 3D objects that are also physically manipulated. The purpose of the present study was to compare mental rotation ability as evaluated by the MRT to three new tasks with physical objects (toy bricks) that were physically manipulated. The different tasks allowed us to vary the processing demands on mental rotation while standardizing other aspects of the tasks. Fifty-nine females and twenty-eight males completed the LMR and HMR conditions (low- and high-mental rotation demands, respectively) of the brick building task (BBT), a visual search task, and the MRT. As demands on mental rotation for the BBT increased, performance decreased and a sex difference, with males outperforming females, increased. There were correlations between all tasks, but they were larger between the versions of the BBT with the MRT. The results suggest that spatial skill is an assembly of interrelated subskills and that the sex difference is sensitive to the demands on mental rotation and dimensionality crossing. The benefits of the BBT are that it is ecologically valid, avoids dimensionality crossing, and the demands on mental rotation can be manipulated.

Dual-task performance of speech and motor skill: verb generation facilitates grasping behaviour.

Pronouncing nouns or verbs while grasping distinctly alters movement. Changes in hand speed and final position occur according to the meaning of the words spoken. These results are typically found when executing a single movement paired with a single word. For example, pronouncing the word 'fast' increased the speed of the hand when reaching to grasp. Our objective was to compare how verb and noun fluency tasks interact with grasping behaviour in a grasp-to-construct task. Because previous imaging research shows that verb and noun production activates distinct neural areas, we reasoned that grasping outcomes would differ according to the category of word produced by participants. Specifically, we hypothesized that verb pronunciation would distinctly affect grasping behaviour compared to producing nouns. We recruited 38 young adults who performed a grasp-to-construct task and two different verbal fluency tasks. Participants completed each task (grasp, verb fluency, and noun fluency) separately as control conditions, and the grasping and each speaking task simultaneously for dual-task conditions. We found that during the dual-task condition, when generating nouns and grasping, participants made significantly more grasping errors (inaccurate grasps) compared to the control and verb dual-task conditions. Moreover, our results revealed a relationship between the number of verbs generated and grasping performance. Participants who generated more verbs were faster and more accurate during the motor component of the dual-task condition. This relationship was not observed when nouns were produced, indicating a unique relationship between verb production and functional grasping. The result is a facilitation effect, diminishing the negative outcome on motor control associated with increased cognitive load (as observed during noun pronunciation).

The association between sedentary behavior and cognitive ability in older adults.

Executive functions (EF) are a grouping of cognitive abilities essential for daily life. Previous research has shown that physical activity (PA) may in fact preserve EF in older adults, but the link between sedentary behavior (SB) and cognitive ability has been less explored. The purpose of this study was to examine the relationship between SB and cognition (executive function and memory) in older adults. Seventy five older adults (74.6 ± 9 years) self-reported their sedentary time (ST) and PA, as well as EF ability (paper-based measure of EF). Participants also completed several performance-based measures of EF and a memory task. Older adults who were less sedentary had superior EF and memory (e.g., Stroop time was significantly faster in less sedentary adults (34.7 s ± 1.9) compared to more sedentary adults (39.6 s ± 1.8), p = .02). Regression analysis showed that total ST was associated with several measures of EF after adjusting for age, and physical activity (e.g., Stroop time ß = .005 (.002, .009). Less cognitively demanding SB (TV viewing and napping) was associated with worse performance on most EF and in the memory task. Performing a hobby was also associated with lower levels of EF and memory. For example, the building times for the Lego task were positively related to napping (r2 = .34), watching TV (r2 = .27), and performing a hobby (r2 = .46). Associations of ST with cognitive abilities were more pronounced in older adults who engaged in less PA. These results suggest that SB may play an important role in cognitive abilities of older adults. Longitudinal studies using performance-based assessments of EF are needed. Lara Coelho and Kayla Hauck contributed equally to the manuscript.

On the Neurocircuitry of Grasping: The influence of action intent on kinematic asymmetries in reach-to-grasp actions.

Evidence from electrophysiology suggests that nonhuman primates produce reach-to-grasp movements based on their functional end goal rather than on the biomechanical requirements of the movement. However, the invasiveness of direct-electrical stimulation and single-neuron recording largely precludes analogous investigations in humans. In this review, we present behavioural evidence in the form of kinematic analyses suggesting that the cortical circuits responsible for reach-to-grasp actions in humans are organized in a similar fashion. Grasp-to-eat movements are produced with significantly smaller and more precise maximum grip apertures (MGAs) than are grasp-to-place movements directed toward the same objects, despite near identical mechanical requirements of the two subsequent (i.e., grasp-to-eat and grasp-to-place) movements. Furthermore, the fact that this distinction is limited to right-handed movements suggests that the system governing reach-to-grasp movements is asymmetric. We contend that this asymmetry may be responsible, at least in part, for the preponderance of right-hand dominance among the global population.

Wait wait, don't tell me: Handedness questionnaires do not predict hand preference for grasping.

Handedness questionnaires are a common screening tool in psychology and neuroscience, used whenever a participant's performance on a given task may conceivably be affected by their laterality. Two widely-used examples of such questionnaires are the Edinburgh Handedness Inventory and the Waterloo Handedness Questionnaire. Both instruments ask respondents to report their hand preference for performing a variety of common tasks (e.g., throwing a ball, or opening a drawer). Here we combined questions from the two instruments (E-WHQ; 22 questions total) and asked participants to report their preferred hand for each via a five-point scale. The purpose of this study was to determine whether responses on the E-WHQ are accurate, reliable, and/or predictive of hand-preference for a simple grasp-to-construct task. Regarding accuracy, handedness scores were 5% lower when participants used a scrambled response key versus a consistent one. Test-retest reliability of the questionnaire was weak, with any given inventory item eliciting a different response from 34% of respondents upon retesting. Neither was the E-WHQ predictively useful-although both left- and right-handers preferred their dominant hands, E-WHQ score did not correlate with overall percentage of dominant-hand grasps in either group. We conclude that the E-WHQ is unsuited for predicting hand preference for grasping.

Chubby hands or little fingers: sex differences in hand representation.

Disturbed body representation is a condition defined by the perception that one's body size is different from their anatomical size. While equal amounts of males and females suffer from disturbed body representation, there appear to be differences in the direction of this distortion. Females will typically overestimate, whereas males will typically underestimate body size. One part of the body that has been consistently misperceived is the hands. This misrepresentation consists of two distinct characteristics: an overestimation of hand width, and an underestimation of finger length. Many of these studies, however, have used predominately female participants, allowing for the possibility that women are driving this distortion. The aim of the present study was to examine possible sex differences in hand perception. To this end, participants estimated the location of ten landmarks on their hands when their hands were hidden from view. Our results indicate that females follow the characteristic distortion, whereas males only underestimate finger length (albeit more than females). These findings are surprising, because the hands are not an area of concern for weight gain/loss. We discuss these findings in relation to body dysmorphia literature.

Long- but not short-term tool-use changes hand representation.

Tool-use has been found to change body representation. For example, participants who briefly used a mechanical grabber to pick up objects perceived their forearms to be longer immediately after its use (e.g., Cardinali et al., Curr Biol 19(12):R478-R479, 2009; they incorporated the tool into their perceived arm size). While some studies have investigated the long-term effects of tool-use on body representation, none of these studies have used a tool that encapsulates the entire body part (e.g., a glove). Moreover, the relationship between tool-use and the body model (the representation of the body's spatial characteristics) has yet to be explored. To test this, we recruited 19 elite baseball players (EBP) and 18 age-matched controls to participate in a hand representation task. We included EBP because of their many years (8+) of training with a tool (baseball glove). The task required participants to place their hands underneath a covered glass tabletop (no vision of their hands), and to point to where they believed 10 locations (the tips and bases of each finger) were on their hands (Coelho et al., Psychol Res 81(6):1224-1231, 2017). Each point's XY coordinates was tracked using an Optotrak camera. From these coordinates, we mapped out the participants perceived hand size. The results showed that when compared to the controls, EBP underestimated hand width and finger length of both hands. This indicates that long-term tool use produces changes in the body model for both, the trained and untrained hands. We conducted a follow-up study to examine if 15 min of glove use would change perceived hand size in control participants. Novice baseball players (participants without baseball experience: NBP) were recruited and hand maps were derived before and after 15 min of active catching with a glove. Results showed no significant differences between the pre and post hand maps. When we compared between the two experiments, the EBP showed smaller hand representation for both hand width and finger length, than the NBP. We discuss these results in relation to theories of altered body ownership.

Hear speech, change your reach: changes in the left-hand grasp-to-eat action during speech processing.

Research has shown that the kinematic characteristics of right-hand movements change when executed during both speech production and processing. Despite the variety of prehension and manual actions used to examine this relationship, the literature has yet to examine potential movement effects using an action with a distinct kinematic signature: the hand-to-mouth (grasp-to-eat) action. In this study, participants performed grasp-to-eat and grasp-to-place actions in (a) a quiet environment and (b) while processing speech. Results during the quiet condition replicated the previous findings; consistently smaller grasp-to-eat (compared to grasp-to-place), maximum grip apertures appeared only when using the right hand. Interestingly, in the listen condition, smaller maximum grip apertures in the grasp-to-eat movement appeared in both the right and left hands, despite the fact that participants were right-handed. This paper addresses these results in relation with similar behaviour observed in children, and discusses implications for functional lateralization and neural organization.

Sensorimotor lateralization scaffolds cognitive specialization.

In this chapter, we review hemispheric differences for sensorimotor function and cognitive abilities. Specifically, we examine the left-hemisphere specialization for visuomotor control and its interplay with language, executive function, and musical training. Similarly, we discuss right-hemisphere lateralization for haptic processing and its relationship to spatial and numerical processing. We propose that cerebral lateralization for sensorimotor functions served as a foundation for the development of higher cognitive abilities and their hemispheric functional specialization. We further suggest that sensorimotor and cognitive functions are inextricably linked. Based on the studies discussed in this chapter our view is that sensorimotor control serves as a loom upon which the fibers of language, executive function, spatial, and numerical processing are woven together to create the fabric of cognition.

Bimanual joint action: correlated timing or "bimanual" movements accomplished by two people.

A crew of two rowing together in perfect synchrony is an example of a task that requires each performer to maintain meticulous timing when coordinating their movements with the other. At the individual level, temporal coordination of the limbs has been observed in bimanual pointing movements even when made to targets of different distance. Timing of the arms is not independent; rather there is a natural temporal coupling. The aim of this experiment was to investigate whether the temporal characteristics of pointing movements can be observed under joint conditions. Sixteen pairs of participants made short and long, unimanual and bimanual pointing movements. In the unimanual and bimanual solo conditions, participants made the movements alone. In the joint condition, each participant contributed one arm to the joint "bimanual" movements. Absolute temporal coupling at movement initiation and termination was measured by the differences in reaction time and total response time. Relative temporal coupling at movement initiation and termination was measured by correlating reaction time and total response time of the left and right limbs. Pointing movements had synchronous movement termination in the bimanual solo conditions and asynchronous termination in the unimanual solo and bimanual joint conditions. The initiation and termination of the arms were not correlated in the unimanual solo condition (initiation r = 0.01, termination r = 0.03). Small-to-medium correlations (r = 0.19, r = 0.24) were observed in the bimanual joint condition, and they were larger than the unimanual solo condition (p = 0.022, p = 0.063). As expected, there were large correlations in the bimanual solo conditions (r = 0.91, r = 0.81). Our findings suggest that absolute temporal coupling does not occur between individuals, but there is evidence for relative temporal coupling in the bimanual joint condition.

Complexity of movement preparation and the spatiotemporal coupling of bimanual reach-to-grasp movements.

There is a movement preparation cost for bimanual asymmetric reaching movements compared to bimanual symmetric movements. This is likely caused by the complex spatiotemporal coupling of bimanual asymmetric movements. The spatiotemporal coupling of bimanual reach-to-grasp movements has been investigated, but not the potential movement preparation costs. The purpose of the present study was to investigate the relationship between movement preparation costs and spatiotemporal coupling of reach-to-grasp movements. Twenty-four participants made unimanual, bimanual symmetric, and bimanual asymmetric reach-to-grasp movements in four-choice reaction time tasks. There was a movement preparation cost for bimanual symmetric reach-to-grasp movements compared to unimanual movements, which was not previously seen for reaching movements. Coordinating two symmetric grasps probably caused this bimanual symmetric cost, as we have previously shown that there is no bimanual symmetric cost for reaching movements. It was also surprising that the complexity of movement preparation was comparable for bimanual symmetric and asymmetric reach-to-grasp movements. However, the spatial coupling of bimanual asymmetric movements at movement initiation suggested that they were prepared as bimanual symmetric movements. Online control was then used to modify these symmetric reach-to-grasp movements into asymmetric movements. Preparing bimanual symmetric reach-to-grasp movements in advance instead of asymmetric movements likely prevented a bimanual asymmetric cost.