Laterality indices consensus initiative (LICI): A Delphi expert survey report on recommendations to record, assess, and report asymmetry in human behavioural and brain research.

Laterality indices (LIs) quantify the left-right asymmetry of brain and behavioural variables and provide a measure that is statistically convenient and seemingly easy to interpret. Substantial variability in how structural and functional asymmetries are recorded, calculated, and reported, however, suggest little agreement on the conditions required for its valid assessment. The present study aimed for consensus on general aspects in this context of laterality research, and more specifically within a particular method or technique (i.e., dichotic listening, visual half-field technique, performance asymmetries, preference bias reports, electrophysiological recording, functional MRI, structural MRI, and functional transcranial Doppler sonography). Experts in laterality research were invited to participate in an online Delphi survey to evaluate consensus and stimulate discussion. In Round 0, 106 experts generated 453 statements on what they considered good practice in their field of expertise. Statements were organised into a 295-statement survey that the experts then were asked, in Round 1, to independently assess for importance and support, which further reduced the survey to 241 statements that were presented again to the experts in Round 2. Based on the Round 2 input, we present a set of critically reviewed key recommendations to record, assess, and report laterality research for various methods.

Head and eye movements are each facilitated by the offset of a central fixation point in a virtual gap paradigm.

Eye movements exhibit reduced latencies when the point of fixation is extinguished prior to, or coincident with, the appearance of a peripheral target. Two independent components are responsible for this facilitation. If the offset occurs before target onset, it presents a warning which stimulates response preparation and execution. If offset occurs prior to or coincident with target onset, it triggers the release of fixation-maintenance neurons in the superior colliculus that can delay saccadic responses. While the warning effect facilitates responses regardless of effector, the fixation release effect is thought to be specific to the oculomotor system. Head movements, like saccades, contribute significantly to gaze shifts and may be generated directly by the SC. While head movements have been shown to benefit from the warning effect, it is unknown if, and to what degree, they are affected by the release of fixation-maintenance neurons responsible for inhibiting saccades. To address this issue, we measured head and eye response latencies in a virtual reality-based gap paradigm, turning off the fixation point either 200 ms before (temporal gap condition), coincident with (step condition), or 1000 ms after (temporal overlap/baseline condition) target onset. Our results indicate that head movements, like saccades, are facilitated by both the warning and release components of the gap paradigm. Further, rotational kinematics during gap trials differed significantly from those observed in step and overlap trials (higher, earlier peak velocities). These results are discussed with respect to the theorized structure and organisation of the superior colliculus in humans.

The Quiet Eye and Expertise: Sustained Fixations Do Not Transfer to Unpracticed Throws Among Expert Dart Players.

The length of the last visual fixation before the critical final phase of a movement-the quiet eye (QE) fixation-is positively correlated with expertise and success. The present study tested the potential for intraskill transfer of QE durations in order to determine whether it is intrinsically linked to expertise development or is a separable skill that may be employed to improve performance under novel circumstances. The authors tracked highly skilled dart throwers' gazes while they executed familiar (highly practiced) and familiar yet novel (distance/effector-modified) sport-specific actions. QE duration was significantly reduced when performing in unfamiliar conditions, suggesting that QE does not transfer to atypical conditions and may therefore be a result of-rather than a contributor to-expertise development. These results imply that intraskill transfer of QE is limited and, consistent with the inhibition hypothesis of QE development, argue against the value of teaching QE as an independent means of improving performance.

The left cerebral hemisphere may be dominant for the control of bimanual symmetric reach-to-grasp movements.

Previous research has established that the left cerebral hemisphere is dominant for the control of continuous bimanual movements. The lateralisation of motor control for discrete bimanual movements, in contrast, is underexplored. The purpose of the current study was to investigate which (if either) hemisphere is dominant for discrete bimanual movements. Twenty-one participants made bimanual reach-to-grasp movements towards pieces of candy. Participants grasped the candy to either place it in their mouths (grasp-to-eat) or in a receptacle near their mouths (grasp-to-place). Research has shown smaller maximum grip apertures (MGAs) for unimanual grasp-to-eat movements than unimanual grasp-to-place movements when controlled by the left hemisphere. In Experiment 1, participants made bimanual symmetric movements where both hands made grasp-to-eat or grasp-to-place movements. We hypothesised that a left hemisphere dominance for bimanual movements would cause smaller MGAs in both hands during bimanual grasp-to-eat movements compared to those in bimanual grasp-to-place movements. The results revealed that MGAs were indeed smaller for bimanual grasp-to-eat movements than grasp-to-place movements. This supports that the left hemisphere may be dominant for the control of bimanual symmetric movements, which agrees with studies on continuous bimanual movements. In Experiment 2, participants made bimanual asymmetric movements where one hand made a grasp-to-eat movement while the other hand made a grasp-to-place movement. The results failed to support the potential predictions of left hemisphere dominance, right hemisphere dominance, or contralateral control.

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.

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.

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.

Children's bilateral advantage for grasp-to-eat actions becomes unimanual by age 10 years.

Studies have shown that infants tend to develop a lateralized hand preference for hand-to-mouth actions earlier than they do a preference for many other grasp-to-place or grasp-to-manipulate tasks, years even before direction of hand preference can be reliably determined. This observation has led to a series of studies contrasting the kinematics of grasp-to-eat and grasp-to-place actions in adults. These studies have described a robust kinematic asymmetry between left- and right-handed grasp-to-eat maximum grip apertures (MGAs) that has been interpreted as a right-hand advantage for feeding that may have led to right-handedness as observed on a global scale. The current study examines grasp-to-eat and grasp-to-place kinematics in two groups of typically developing children aged 7 to 12 years. It was found that the previously described task difference is present in both hands among younger children and that the effect does not become lateralized until the end of the first decade of life. Additional kinematics of both the dominant and non-dominant hands are described in detail to augment a growing catalogue of reach-to-grasp action descriptions for typically developing children. The maturation of the right-hand advantage for grasp-to-eat actions is discussed in terms of an inherent right-hand/left-hemisphere bias for such actions that may have influenced the development of population-level right-handedness in humans.

Evidence for right-hand feeding biases in a left-handed population.

We have recently shown that actions with similar kinematic requirements, but different end-state goals may be supported by distinct neural networks. Specifically, we demonstrated that when right-handed individuals reach-to-grasp food items with intent to eat, they produce smaller maximum grip apertures (MGAs) than when they grasp the same item with intent to place it in a location near the mouth. This effect was restricted to right-handed movements; left-handed movements showed no difference between tasks. The current study investigates whether (and to which side) the effect may be lateralized in left-handed individuals. Twenty-one self-identified left-handed participants grasped food items of three different sizes while grasp kinematics were captured via an Optotrak Certus motion capture array. A main effect of task was identified wherein the grasp-to-eat action generated significantly smaller MGAs than did the grasp-to-place action. Further analysis revealed that similar to the findings in right-handed individuals, this effect was significant only during right-handed movements. Upon further inspection however, we found individual differences in the magnitude and direction of the observed lateralization. These results underscore the evolutionary significance of the grasp-to-eat movement in producing population-level right-handedness in humans as well as highlighting the heterogeneity of the left-handed population.

Eating interrupted: the effect of intent on hand-to-mouth actions.

Evidence from recent neurophysiological studies on nonhuman primates as well as from human behavioral studies suggests that actions with similar kinematic requirements but different end-state goals are supported by separate neural networks. It is unknown whether these different networks supporting seemingly similar reach-to-grasp actions are lateralized, or if they are equally represented in both hemispheres. Recently published behavioral evidence suggests certain networks are lateralized to the left hemisphere. Specifically, when participants used their right hand, their maximum grip aperture (MGA) was smaller when grasping to eat food items than when grasping to place the same items. Left-handed movements showed no difference between tasks. The present study investigates whether the differences between grasp-to-eat and grasp-to-place actions are driven by an intent to eat, or if placing an item into the mouth (sans ingestion) is sufficient to produce asymmetries. Twelve right-handed adults were asked to reach to grasp food items to 1) eat them, 2) place them in a bib, or 3) place them between their lips and then toss them into a nearby receptacle. Participants performed each task with large and small food items, using both their dominant and nondominant hands. The current study replicated the previous finding of smaller MGAs for the eat condition during right-handed but not left-handed grasps. MGAs in the eat and spit conditions did not significantly differ from each other, suggesting that eating and bringing a food item to the mouth both utilize similar motor plans, likely originating within the same neural network. Results are discussed in relation to neurophysiology and development.

Manual asymmetries in the kinematics of a reach-to-grasp action.

In the present study, we manipulated the perceived demand of an ecologically valid task to investigate the possible presence of manual asymmetries in a reach-to-grasp action. Participants reached, grasped and sipped from a water glass under low (nearly empty) and high (nearly full) demand conditions. Participants reached to grasp in closed-loop, open-loop and delay visual conditions. Manual asymmetries were found in movement time, peak velocity and maximum grip aperture variability. Consistent with reach-to-point literature: (1) right-handed actions were completed in less time than left-handed actions in visually and memory-guided conditions; (2) right-handed movements were more accurate (i.e., produced more consistent maximum grip apertures) than left-handed movements in visually guided conditions. The results support a theory of left-hemisphere specialization for visual control of action.

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.

Hand preference across the lifespan: effects of end-goal, task nature, and object location.

In the present study we investigate age-related changes in hand preference for grasping and the influence of task demands on such preference. Children (2-11), young-adults (17-28) and older-adults (57-90) were examined in a grasp-to-eat and a grasp-to-construct task. The end-goal of these tasks was different (eat vs. construct) as was the nature of the task (unimanual vs. bimanual). In both tasks, ipsilateral and contralateral grasps were analyzed. Results showed a right-hand preference that did not change with age. Across the three age groups, a more robust right-hand preference was observed for the unimanual, grasp-to-eat task. To disentangle if the nature (unimanual) or the end-goal (grasp-to-eat) was the driver of the robust right-hand preference, a follow up experiment was conducted. Young-adult participants completed a unimanual grasp-to-place task. This was contrasted with the unimanual grasp-to-eat task and the bimanual grasp-to-construct task. Rates of hand preference for the grasp-to-eat task remained the highest when compared to the other two grasping tasks. Together, the results demonstrate that hand preference remains stable from childhood to older adulthood, and they suggest that a left hemisphere specialization exists for grasping, particularly when bringing food to the mouth.

On the evolution of handedness: evidence for feeding biases.

Many theories have been put forward to explain the origins of right-handedness in humans. Here we present evidence that this preference may stem in part from a right hand advantage in grasping for feeding. Thirteen participants were asked to reach-to-grasp food items of 3 different sizes: SMALL (Cheerios®), MEDIUM (Froot Loops®), and LARGE (Oatmeal Squares®). Participants used both their right- and left-hands in separate blocks (50 trials each, starting order counterbalanced) to grasp the items. After each grasp, participants either a) ate the food item, or b) placed it inside a bib worn beneath his/her chin (25 trials each, blocked design, counterbalanced). The conditions were designed such that the outward and inward movement trajectories were similar, differing only in the final step of placing it in the mouth or bib. Participants wore Plato liquid crystal goggles that blocked vision between trials. All trials were conducted in closed-loop with 5000 ms of vision. Hand kinematics were recorded by an Optotrak Certus, which tracked the position of three infrared diodes attached separately to the index finger, thumb, and wrist. We found a task (EAT/PLACE) by hand (LEFT/RIGHT) interaction on maximum grip aperture (MGA; the maximum distance between the index finger and thumb achieved during grasp pre-shaping). MGAs were smaller during right-handed movements, but only when grasping with intent to eat. Follow-up tests show that the RIGHT-HAND/EAT MGA was significantly smaller than all other hand/task conditions. Because smaller grip apertures are typically associated with greater precision, our results demonstrate a right-hand advantage for the grasp-to-eat movement. From an evolutionary perspective, early humans may have preferred the hand that could grasp food with more precision, thereby maximizing the likelihood of retrieval, consumption, and consequently, survival.