Sex differences in visuospatial cognition- a female advantage in jigsaw puzzle solving.

Mentally visualizing objects, understanding relationships between two- or three- dimensional objects, and manipulating objects in space are some examples of visuospatial abilities. Numerous studies have shown that male participants outperform female participants in visuospatial tasks, particularly in mental rotation. One exception is solving jigsaw puzzles. Performance by seven- to eight-year-old girls was found to be superior to that of boys of the same age (Kocijan et al. 2017). No study, however, has confirmed this finding in an adult population, where sex differences are often detectable. Seventy-nine young adult participants were given four different jigsaw puzzles and the Shepard and Metzler mental rotation test (MRT) with two main goals: First, to investigate possible sex differences in jigsaw puzzle solving, and second, to explore a potential relationship between mental rotation and jigsaw puzzle solving. We hypothesized that female participants would outperform males in the jigsaw puzzles but males would outperform females in the MRT. The findings confirmed this hypothesis. Notably, the male performance in jigsaw puzzle solving was attributed to their sex and mediated by their higher MRT scores. These results yielded two key insights. First, they indicate a dissociation between these two visuospatial abilities, jigsaw puzzle solving and mental rotation; and second, female and male participants capitalize on their distinct cognitive strengths when solving visuospatial tasks.

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.

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.

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.

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.

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.

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.

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.

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.

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.

Kinematics of ventrally mediated grasp-to-eat actions: right-hand advantage is dependent on dorsal stream input.

Studies have suggested a left-hemisphere specialization for visually guided grasp-to-eat actions by way of task-dependent kinematic asymmetries (i.e., smaller maximum grip apertures for right-handed grasp-to-eat movements than for right-handed grasp-to-place movements or left-handed movements of either type). It is unknown, however, whether this left-hemisphere/right-hand kinematic advantage is reliant on the dorsal "vision-for-action" visual stream. The present study investigates the kinematic differences between grasp-to-eat and grasp-to place actions performance during closed-loop (i.e., dorsally mediated) and open-loop delay (i.e., ventrally mediated) conditions. Twenty-one right-handed adult participants were asked to reach to grasp small food items to (1) eat them, or (2) place them in a container below the mouth. Grasps were performed in both closed-loop and open-loop delay conditions, in separate sessions. We show that participants displayed the right-hand grasp-to-eat kinematic advantage in the closed-loop condition, but not in the open-loop delay condition. As no task-dependent kinematic differences were found in ventrally mediated grasps, we posit that the left-hemisphere/right-hand advantage is dependent on dorsal stream processing.

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.

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.

The inimitable mouth: task-dependent kinematic differences are independent of terminal precision.

Previous studies in our lab have described kinematic difference between grasp-to-eat and grasp-to-place movements, whereby participants produce smaller maximum grip apertures (MGAs) when grasping to bring the item to the mouth than when grasping to bring the item to a container near the mouth. This task difference is limited to right-handed movements, regardless of handedness; it has, therefore, been interpreted as evidence of left-hemisphere lateralization of the grasp-to-eat and other hand-to-mouth grasping movements. However, the difference in end-goal aperture may have accounted for both the kinematic signature (smaller MGAs) and their lateralized expression. Specifically, if the right hand is more sensitive to the precision requirements of secondary movements, it may have produced more precise MGAs for actions whose ultimate goal is the small-aperture mouth rather than a comparatively large aperture container. The current study addresses this question by replacing the previously-used bib with a small drinking glass whose aperture more closely resembles that of the mouth. 25 adult participants reached-to-grasp small cereal items to either (a) eat them, or (b) place them into a small-aperture glass hanging beneath their chin. Results once more showed a lateralised kinematic signature in the form of smaller MGAs for the eat action, demonstrating that the signature is not a result of lateralized sensitivity to a movement's secondary precision requirements. We discuss these results in terms of their impact on predominant theories regarding visual guidance of grasping movements.

A kinematic examination of hand perception.

Previous research has found that the perception of our hands is inaccurate. This distorted representation has several constant characteristics including an overestimation of hand width and an underestimation of finger length. In this study, we further investigate this phenomenon by exploring the boundaries of hand representation. Participants placed one hand underneath a table top so it was occluded from view. Using their free hand, participants were instructed to point to the location where they believed the tips and bases of each of their fingers were. These ten landmarks were recorded using a motion capture system. One group of participants pointed to the landmarks in a random order (as done in previous studies) while another group pointed to them in a systematic fashion (from the tip of the thumb sequentially through to the pinky). Furthermore, to explore if having a frame of reference facilitates hand perception, some participants initiated each of their estimations directly from the previous landmark while others initiated them from a home spot located outside the span of the hand. Results showed that the participants who pointed in the systematic order made numerous accurate judgments of hand size and were overall more precise than participants who pointed in a random order. Including a frame of reference however, had no effect on the judgments. The results also showed asymmetries in hand perception. These findings are discussed in relation to different possible internal body representations and hemispheric asymmetries in body perception.

The destination defines the journey: an examination of the kinematics of hand-to-mouth movements.

Long-train electrical stimulation of the motor and premotor cortices of nonhuman primates can produce either hand-to-mouth or grasp-to-inspect movements, depending on the precise location of stimulation. Furthermore, single-neuron recording studies identify discrete neuronal populations in the inferior parietal and ventral premotor cortices that respond uniquely to either grasp-to-eat or grasp-to-place movements, despite their identical mechanistic requirements. These studies demonstrate that the macaque motor cortex is organized around producing functional, goal-oriented movements, rather than simply fulfilling muscular prerequisites of action. In humans, right-handed hand-to-mouth movements have a unique kinematic signature; smaller maximum grip apertures are produced when grasping to eat than when grasping to place identical targets. This is evidence that the motor cortex in humans is also organized around producing functional movements. However, in both macaques and humans, grasp-to-eat/hand-to-mouth movements have always been elicited using edible targets and have (necessarily) been paired with mouth movement. It is therefore unknown whether the kinematic distinction is a natural result of grasping food and/or is simply attributable to concurrent opening of the mouth while grasping. In experiment 1, we used goal-differentiated grasping tasks, directed toward edible and inedible targets, to show that the unique kinematic signature is present even with inedible targets. In experiment 2, we used the same goal-differentiated grasping tasks, either coupled with or divorced from an open-mouth movement, to show that the signature is not attributable merely to a planned opening of the mouth during the grasp. These results are discussed in relation to the role of hand-to-mouth movements in human development, independently of grasp-to-eat behavior.

Is that graspable? Let your right hand be the judge.

A right-hand preference for visually-guided grasping has been shown on numerous accounts. Grasping an object requires the integration of both visual and motor components of visuomotor processing. It has been suggested that the left hemisphere plays an integral role in visuomotor functions. The present study serves to investigate whether the visual processing of graspable objects, without any actual reaching or grasping movements, yields a right-hand (left-hemisphere) advantage. Further, we aim to address whether such an advantage is automatically evoked by motor affordances. Two groups of right-handed participants were asked to categorize objects presented on a computer monitor by responding on a keypad. The first group was asked to categorize visual stimuli as graspable (e.g. apple) or non-graspable (e.g. car). A second group categorized the same stimuli but as nature-made (e.g. apple) or man-made (e.g. car). Reaction times were measured in response to the visually presented stimuli. Results showed a right-hand advantage for graspable objects only when participants were asked to respond to the graspable/non-graspable categorization. When participants were asked to categorize objects as nature-made or man-made, a right-hand advantage for graspable objects did not emerge. The results suggest that motor affordances may not always be automatic and might require conscious representations that are appropriate for object interaction.