Mental rotation task of hands: differential influence number of rotational axes

Various studies on the hand laterality judgment task, using complex sets of stimuli, have shown that the judgments during this task are dependent on bodily constraints. More specific, these studies showed that reaction times are dependent on the participant’s posture or differ for hand pictures rotated away or toward the mid-sagittal plane (i.e., lateral or medial rotation, respectively). These findings point to the use of a cognitive embodied process referred to as motor imagery. We hypothesize that the number of axes of rotation of the displayed stimuli during the task is a critical factor for showing engagement in a mental rotation task, with an increased number of rotational axes leading to a facilitation of motor imagery. To test this hypothesis, we used a hand laterality judgment paradigm in which we manipulated the difficulty of the task via the manipulation of the number of rotational axes of the shown stimuli. Our results showed increased influence of bodily constraints for increasing number of axes of rotation. More specifically, for the stimulus set containing stimuli rotated over a single axis, no influence of biomechanical constraints was present. The stimulus sets containing stimuli rotated over more than one axes of rotation did induce the use of motor imagery, as a clear influence of bodily constraints on the reaction times was found. These findings extend and refine previous findings on motor imagery as our results show that engagement in motor imagery critically depends on the used number of axes of rotation of the stimulus set.     

Corticospinal facilitation during first and third person imagery

Motor imagery can be defined as the covert rehearsal of movement. Previous research with transcranial magnetic stimulation (TMS) has demonstrated that motor imagery increases the corticospinal excitability of the primary motor cortex in the area corresponding to the representation of the muscle involved in the imagined movement. This research, however, has been limited to imagery of oneself in motion. We extend the TMS research by contrasting first person imagery and third person imagery of index finger abduction-adduction movements. Motor evoked potentials were recorded from first dorsal interosseous (FDI) and abductor digiti minimi (ADM) during single pulse TMS. Participants performed first and third person motor imagery, visual imagery, and static imagery. Visual imagery involved non biological motion while static imagery involved a first person perspective of the unmoving hand. Relative to static imagery, excitability during imagined movement increased in FDI but not ADM. The facilitation in first person imagery adds to previous findings. A greater facilitation of MEPs recorded from FDI was found in third person imagery where the action was clearly attributable to another person. We interpret this novel result in the context of observed action and imagined observation of self action, and attribute the result to activation of mirror systems for matching the imagined action with an inner visuo-motor template.     

Developmental changes of the biomechanical effect in motor imagery

Motor imagery has been investigated in childhood and early adolescence, but not across adolescence stages; moreover, available evidence did not clarify whether the involvement of motor information in mental rotation of body parts becomes stronger or weaker during development. In the present study, we employed the hand laterality task to assess motor imagery in ninety-seven typically developing adolescents divided into three age groups (i.e., 11–12, 14–15, and 17–18 years); mental rotation of objects and letters were also assessed. As a specific marker of the motor involvement in mental rotation of body parts, we assessed the so-called biomechanical effect, that is, the advantage for judging hand pictures showing physically comfortable positions with respect to hand pictures showing physically impossible or awkward positions. Results demonstrated that the biomechanical effect did not significantly affect early adolescents’ performance, whereas it became significant in 14- to 15-year-old participants and even more stronger in 17- to 18-year-old participants; this pattern did not depend on an increase in processing speed to mentally rotate both corporeal and non-corporeal (objects and letters) stimuli. The present findings demonstrated that: (1) motor imagery undergoes a continuous and progressive refinement throughout adolescence, and (2) full exploitation of motor information to mentally transform corporeal stimuli can be attained in late adolescence only.     

When the left hand does not know what the left hand is doing: response mode affects mental rotation of hands

Mentally simulating a movement is known to share temporal and kinematic characteristics with the execution of the same movement, and this is thought to be reflected in the sharing of neural resources between the two activities. A powerful method of implicitly facilitating such mental simulation (or motor imagery) in individuals is to present them with a picture of a hand and ask them to identify its laterality (i.e. left or right). The mental rotation undertaken in order to complete this hand laterality recognition task (HLRT) provides an effective form of motor imagery, and the task has become an influential tool in clinical and experimental studies. However, performance on the task is modified by numerous factors, and there is a suggestion that the method of response demanded by different versions of the task may have a modulating effect. Here, we compared performance on the HLRT when responding verbally or manually in a group of unimpaired right-handed participants. For manual responses, we also compared performance when participants responded unimanually, using the index and middle fingers of their dominant or non-dominant hand. Performance was poorer for the manual compared to the verbal condition both in terms of accuracy and response time. Furthermore, for manual responses, the requirement to make a response with a specific limb selectively disrupted the ability to recognise an image of the corresponding limb. The disruption is considered to reflect difficulty in concurrently planning two actions with the same limb (manual response and mental rotation). Implications for the interpretation of existing and future studies are discussed.     

On the link between action planning and motor imagery: a developmental study

We examined the link between action planning and motor imagery in 6- and 8-year-old children. Action planning efficiency was assessed with a bar transport task. Motor imagery and visual imagery abilities were measured using a hand mental rotation task and a number (i.e., non-body stimuli) mental rotation task, respectively. Overall, results showed that performance varied with age in all tasks, performance being progressively refined with development. Importantly, action planning performance was correlated with motor imagery, whereas no relationship was evident between action planning and visual imagery at any age. The results showed that the ability to engage sensorimotor mechanisms when solving a motor imagery task was concomitant with action planning efficiency. The present work is the first demonstration that evaluating the consequences of the upcoming action in grasping depends on children’s abilities to mentally simulate the response options to choose the most efficient grasp.     

Self-touch affects motor imagery: a study on posture interference effect

Several studies showed that mental rotation of body parts is interfered with by manipulation of the subjects’ posture. However, the experimental manipulations in such studies, e.g., to hold one arm flexed on one’s own chest, activated not only proprioceptive but also self-tactile information. Here, we tested the hypothesis that the combination of self-touch and proprioception is more effective than proprioception alone in interfering with motor imagery. In Experiment 1 right- and left-handers were required to perform the hand laterality task, while holding one arm (right or left) flexed with the hand in direct contact with their chest (self-touch condition, STC) or with the hand placed on a wooden smooth surface in correspondence with their chest (no self-touch condition, NoSTC); in a third neutral condition, subjects kept both arms extended (neutral posture condition, NPC). Right-handers were slower when judging hand laterality in STC with respect to NoSTC and NPC, particularly when the sensory manipulation involved their dominant arm. No posture-related effect was observed in left-handers. In Experiment 2, by applying the same sensory manipulations as above to both arms, we verified that previous results were not due to a conflict between perceived position of the two hands. These data highlighted a complex interaction between body schema and motor imagery, and underlined the role of hand dominance in shaping such interaction.     

Corticomotor facilitation associated with observation,imagery and imitation of hand actions: a comparative study in young and old adults

In the present report, we extent our previous findings (Clark et al. in Neuropsychologia 42:105–122, 2004) on corticomotor facilitation associated with covert (observation and imagery) and overt execution (action imitation) of hand actions to better delineate the selectivity of the effect in the context of an object-oriented action. A second aim was to examine whether the pattern of facilitation would be affected by age. Corticomotor facilitation was determined in two groups of participants (young n = 21, 24 ± 2 years; old n = 19, 62 ± 6 years) by monitoring changes in the amplitude and latency of motor evoked potentials (MEPs) elicited in hand muscles by transcranial magnetic stimulation. MEP responses were measured from both the first dorsal interosseous (FDI, task selective muscle) and the abductor digiti minimi (ADM) of the right hand while participants attended to four different video presentations. Each of four videos provided specific instructions for participants to either: (1) close their eyes and relax (REST), (2) observe the action attentively (OBS), (3) close their eyes and mentally simulate the action (IMAG), or (4) imitate the action (IMIT). The action depicted in the videos represented a male subject cutting a piece of material with scissors. In the young group, the pattern of results revealed selective facilitation in the FDI in conditions involving either covert (OBS and IMAG) or overt action execution (IMIT). In the ADM, only overt execution with action imitation was associated with significant MEP facilitation. In the old group, a similar pattern of results was observed, although the modulation was less selective than that seen in the young group. In fact, older individuals often exhibited concomitant facilitation in both the FDI and ADM during either covert (OBS and IMAG conditions) or overt action execution (IMIT condition). Taken together, these results further corroborate the notion that the corticomotor system is selectively active when actions are covertly executed through internal simulation triggered by observation or by motor imagery, as proposed by Jeannerod (Neuroimage 14:S103–S109, 2001). With aging, the ability to produce corticomotor facilitation in association with covert action execution appears to be largely preserved, although there seems to be a loss in selectivity. This lack of selectivity may, in turn, reflect age-related alterations in the function of the corticospinal system, which may impair the ability to individuate finger movements either in the covert or overt stage of action execution.     

Judging hand laterality from my or your point of view: Interactions between motor imagery and visual perspective

Motor imagery tasks (hand laterality judgment) are usually performed with respect to a self-body (egocentric) representation, but manipulations of stimulus features (hand orientation) can induce a shift to other's body (allocentric) reference frame. Visual perspective taking tasks are also performed in self-body perspective but a shift to an allocentric frame can be triggered by manipulations of context features (e.g., another person present in the to-be-judged scene). Combining hand laterality task and visual perspective taking, we demonstrated that both stimulus and context features can modulate motor imagery performance. In Experiment 1, participants judged laterality of a hand embedded in a human or non-human silhouette. Results showed that observing a human silhouette interfered with judgments on “egocentric hand stimuli” (right hand, fingers up). In Experiment 2, participants were explicitly required to judge laterality of a hand embedded in a human silhouette from their own (egocentric group) or from the silhouette's perspective (allocentric group). Consistent with previous results, the egocentric group was significantly faster than the allocentric group in judging fingers-up right hand stimuli. These findings showed that concurrent activation of egocentric and allocentric frames during mental transformation of body parts impairs participants’ performance due to a conflict between motor and visual mechanisms.     

Implementation of specific motor expertise during a mental rotation task of hands

Mental rotation of the hands classically induces kinesthetic effects according to the direction of the rotation, with faster response times to the hands’ medial rotations compared with lateral rotations, and is thus commonly used to induce engagement in motor imagery (MI). In the present study, we compared the performances of table tennis players (experts on hand movements), who commonly execute and observe fast hand movements, to those of soccer players (non-experts on hand movements) on a mental rotation task of hands. Our results showed a significant effect of the direction of rotation (DOR) confirming the engagement of the participants in MI. In addition, only hand movement experts were faster when the task figures corresponded to their dominant hand compared with the non-dominant hand, revealing a selective effect of motor expertise. Interestingly, the effect of the DOR collapsed in hand movement experts only when the task figures corresponded to their dominant hand, but it is noteworthy that lateral and medial rotations of the right-hand stimuli were not faster than medial rotations of the left-hand stimuli. These results are discussed in relation to possible strategies during the task. Overall, the present study highlights the embodied nature of the mental rotation task of hands by revealing selective effects of motor expertise.     

Simultaneous action execution and observation optimise grasping actions

Action observation and execution share overlapping neural resonating mechanisms. In the present study, we sought to examine the effect of the activation of this system during concurrent movement observation and execution in a prehension task, when no a priori information about the requirements of grasping action was available. Although it is known that simultaneous activation by observation and execution influences motor performance, the importance of the delays of these two events and the specific effect of movement observation itself (and not the prediction of the to-be-observed movement) on action performance are poorly known. Fine-grained kinematic analysis of both the transport and grasp components of the movement should provide knowledge about the influence of movement observation on the precision and the performance of the executed movement. The experiment involved two real participants who were asked to grasp a different side of a single object that was composed of a large and a small part. In the first experiment, we measured how the transport component and the grasp component were affected by movement observation. We tested whether this influence was greater if the observed movement occurred just before the onset of movement (200 ms) or well before the onset of movement (1 s). In a second experiment, to reproduce the previous experiment and to verify the specificity of the grasping movements, we also included a condition consisting of pointing towards the object. Both experiments showed two main results. A general facilitation of the transport component was found when observing a simultaneous action, independent of its congruency. Moreover, a specific facilitation of the grasp component was present during the observation of a congruent action when movement execution and observation were nearly synchronised. While the general facilitation may arise from a competition between the two participants as they reached for the object, the specific facilitation of the grasp component seems to be directly related to mirror neuron system activity induced by action observation itself. Moreover, the time course of the events appears to be an essential factor for this modulation, implying the transitory activation of the mirror neuron system.     

Passive reading and motor imagery about hand actions and tool-use actions: an fMRI study

Recent studies have shown that motor activations in action verb comprehension can be modulated by task demands (e.g., motor imagery vs. passive reading) and the specificity of action verb meaning. However, how the two factors work together to influence the involvement of the motor system during action verb comprehension is still unclear. To address the issue, the current study investigated the brain activations in motor imagery and passive reading of verbs about hand actions and tool-use actions. Three types of Chinese verbs were used, including hand-action verbs and two types of tool-use verbs emphasizing either the hand or tools information. Results indicated that all three types of verbs elicited common activations in hand motor areas during passive reading and motor imagery. Contrast analyses showed that in the hand verbs and the tool verbs where the hand information was emphasized, motor imagery elicited stronger effects than passive reading in the superior frontal gyrus, supplemental motor area and cingulate cortex that are related to motor control and regulation. For tool-use verbs emphasizing tools information, the motor imagery task elicited stronger activity than passive reading in occipital regions related to visual imagery. These results suggest that motor activations during action verb comprehension can be modulated by task demands and semantic features of action verbs. The sensorimotor simulation during language comprehension is flexible and determined by the interactions between linguistic and extralinguistic contexts.     

The effect of unconditional stimulus modality and intensity on blink startle and electrodermal responses

Attentional accounts of blink facilitation during Pavlovian conditioning predict enhanced reflexes if reflex and unconditional stimuli (US) are from the same modality. Emotional accounts emphasize the importance of US intensity. In Experiment 1, we crossed US modality (tone vs. shock) and intensity in a 2 × 2 between-subjects design. US intensity but not US modality affected blink facilitation. In Experiment 2, we demonstrated that the results from Experiment I were not due to the motor task requirements employed. In Experiment 3, we used a within-subjects design to investigate the effects of US modality and intensity. Contrary to predictions derived from an attentional account, blink facilitation was larger during conditional stimuli that preceded shock than during those that preceded tones. The present results are not consistent with an attentional account of blink facilitation during Pavlovian conditioning in humans.     

Lateralization in motor facilitation during action observation: a TMS study

Action observation facilitates corticospinal excitability. This is presumably due to a premotor neural system that is active when we perform actions and when we observe actions performed by others. It has been speculated that this neural system is a precursor of neural systems subserving language. If this theory is true, we may expect hemispheric differences in the motor facilitation produced by action observation, with the language-dominant left hemisphere showing stronger facilitation than the right hemisphere. Furthermore, it has been suggested that body parts are recognized via cortical regions controlling sensory and motor processing associated with that body part. If this is true, then corticospinal facilitation during action observation should be modulated by the laterality of the observed body part. The present study addressed these two issues using TMS for each motor cortex separately as participants observed actions being performed by a left hand, a right hand, or a control stimulus on the computer screen. We found no overall difference between the right and left hemisphere for motor-evoked potential (MEP) size during action observation. However, when TMS was applied to the left motor cortex, MEPs were larger while observing right hand actions. Likewise, when TMS was applied to the right motor cortex, MEPs were larger while observing left hand actions. Our data do not suggest left hemisphere superiority in the facilitating effects of action observation on the motor system. However, they do support the notion of a sensory-motor loop according to which sensory stimulus properties (for example, the image of a left hand or a right hand) directly affect motor cortex activity, even when no motor output is required. The pattern of this effect is congruent with the pattern of motor representation in each hemisphere.     

Effect of observation combined with motor imagery of a skilled hand-motor task on motor cortical excitability: Difference between novice and expert

We examined the effects of observation combined with motor imagery (MI) of a skilled hand-motor task on motor cortex excitability, which was assessed by transcranial magnetic stimulation (TMS). Novices and experts at 3-ball cascade juggling (3BCJ) participated in this study. In one trial, the subjects observed a video clip of 3BCJ while imagining performing it. In addition, the subjects also imagined performing 3BCJ without video clip observation. Motor evoked potentials (MEPs) were recorded from the hand muscles that were activated by the task during each trial. In the novices, the MEP amplitude was significantly increased by video clip observation combined with MI. In contrast, MI without video clip observation significantly increased the MEP amplitude of the experts. These results suggest that action observation of 3BCJ increases the ability of novices to make their MI performing the task. Meanwhile, experts use their own motor program to recall their MI of the task.     

Facilitation of cortically evoked potentials with motor imagery during post-exercise depression of corticospinal excitability

This study examined whether muscle fatigue alters the facilitatory effect of motor imagery on corticospinal excitability. We aimed to determine if post-exercise depression of potentials evoked magnetically from the motor cortex is associated with alterations in internally generated movement plans. In experiment 1, motor-evoked potentials (MEPs) were recorded from two right hand and two right forearm muscles, at rest and during motor imagery of a maximal handgrip contraction, in eight neurologically normal subjects, before and after a 2-min maximal voluntary handgrip contraction. Resting MEP amplitude was facilitated by motor imagery in three of the four muscles, but consistently only in two. Motor imagery also reduced the trial-to-trial variability of resting MEPs. Following the exercise, resting MEP amplitude was depressed reliably in only one muscle engaged in the task, although two other muscles exhibited some depression. Motor imagery MEPs were smaller after exercise, but the degree of facilitation compared to the rest MEP was unchanged. In experiment 2, TMS intensity was increased after exercise-induced MEP depression so that the MEP amplitude matched the pre-exercise baseline. The amplitude of the MEP facilitated with motor imagery was not altered by MEP depression, nor was it increased when the TMS intensity was increased. These results suggest, at least with a simple motor task, that while post-exercise depression reduces corticospinal excitability, it does not appear to significantly affect the strength of the input to the motor cortex from those areas of the brain responsible for the storage and generation of internal representations of movement.     

Combined effects of planning and execution constraints on bimanual task performance

In this study we investigated the relative impact of planning and execution constraints on discrete bimanual task performance. In particular, in a bimanual CD-placement task, we compared people’s preference to end movements comfortably with their preference to move symmetrically. In “Experiment 1” we examined the degree of interlimb coupling as participants repositioned two CDs in a CD rack by simultaneously moving their arms mirror-symmetrically or asymmetrically into comfortable or uncomfortable end postures. Interlimb coupling was stronger when the arms moved symmetrically towards uncomfortable end postures. In “Experiment 2” participants were asked to realize specific end orientations of the CDs but they were free to choose an initial grip type and subsequent direction of forearm rotation. Surprisingly, the participants did not move their arms symmetrically but preferred to end in a comfortable posture with their right hand but not with their left hand. We conclude that in discrete bimanual task performance the tendency to end movements in a comfortable posture dominates over the tendency to synchronously activate homologous muscle pairs. The lateralized end-state comfort effect suggests a hemispheric specialization for motor planning.     

The effect of stimulus modality and task difficulty on attentional modulation of blink startle

The effects of the sensory modality of the lead stimulus and of task difficulty on attentional modulation of the electrical and acoustic blink reflex were examined. Participants performed a discrimination and counting task with either two acoustic, two visual, or two tactile lead stimuli. In Experiment 1, facilitation of the electrically elicited blink was greater during task-relevant than during task-irrelevant lead stimuli. Increasing task difficulty enhanced magnitude facilitation for acoustic lead stimuli. In Experiment 2, acoustic blink facilitation was greater during task-relevant lead stimuli, but was unaffected by task difficulty. Experiment 3 showed that a further increase in task difficulty did not affect acoustic blink facilitation during visual lead stimuli. The observation that blink reflexes are facilitated by attention in the present task domain is consistent across a range of stimulus modality and task difficulty conditions.     

Line bisection by eye and by hand reveal opposite biases

The vision-for-action literature favours the idea that the motor output of an action—whether manual or oculomotor—leads to similar results regarding object handling. Findings on line bisection performance challenge this idea: healthy individuals bisect lines manually to the left of centre and to the right of centre when using eye fixation. In case that these opposite biases for manual and oculomotor action reflect more universal compensatory mechanisms that cancel each other out to enhance overall accuracy, one would like to observe comparable opposite biases for other material. In the present study, we report on three independent experiments in which we tested line bisection (by hand, by eye fixation) not only for solid lines, but also for letter lines; the latter, when bisected manually, is known to result in a rightward bias. Accordingly, we expected a leftward bias for letter lines when bisected via eye fixation. Analysis of bisection biases provided evidence for this idea: manual bisection was more rightward for letter as compared to solid lines, while bisection by eye fixation was more leftward for letter as compared to solid lines. Support for the eye fixation observation was particularly obvious in two of the three studies, for which comparability between eye and hand action was increasingly adjusted (paper–pencil versus touch screen for manual action). These findings question the assumption that ocular motor and manual output are always inter-changeable, but rather suggest that at least for some situations ocular motor and manual output biases are orthogonal to each other, possibly balancing each other out.     

Suppression of corticospinal excitability during negative motor imagery

To investigate the effect of negative motor imagery on corticospinal excitability, we performed transcranial magnetic stimulation (TMS) studies in seven healthy subjects during imagination of suppressing movements. Subjects were asked to imagine suppression of TMS-induced twitching movement of their nondominant left hands by attempting to increase the amount of relaxation after receiving an auditory NoGo cue (negative motor imagery), but to imagine squeezing hands after a Go cue (positive motor imagery). Single- and paired-pulse TMS were triggered at 2 s after Go or NoGo cues. Motor-evoked potentials (MEPs) were recorded in the first dorsal interosseus (FDI), abductor pollicis brevis (APB), and abductor digiti minimi (ADM) muscles of the left hand. Paired-pulse TMS with subthreshold conditioning stimuli at interstimulus intervals of 2 (short intracortical inhibition) and 15 ms (intracortical facilitation) and that with suprathreshold conditioning stimuli at interstimulus interval of 80 ms (long intracortical inhibition) were performed in both negative motor imagery and control conditions. Compared with the control state (no imagination), MEP amplitudes of FDI (but not APB and ADM) were significantly suppressed in negative motor imagery, but those from all three muscles were unchanged during positive motor imagery. F-wave responses (amplitudes and persistence) were unchanged during both negative and positive motor imagery. During negative motor imagery, resting motor threshold was significantly increased, but short and long intracortical inhibition and intracortical facilitation were unchanged. The present results demonstrate that excitatory corticospinal drive is suppressed during imagination of suppressing movements.     

Dissociated effects of distractors on saccades and manual aiming

The remote distractor effect (RDE) is a robust phenomenon whereby target-directed saccades are delayed by the appearance of a distractor. This effect persists even when the target location is perfectly predictable. The RDE has been studied extensively in the oculomotor domain but it is unknown whether it generalises to other spatially oriented responses. In three experiments, we tested whether the RDE generalises to manual aiming. Experiment 1 required participants to move their hand or eyes to predictable targets presented alone or accompanied by a distractor in the opposite hemifield. The RDE was observed for the eyes but not for the hand. Experiment 2 replicated this dissociation in a more naturalistic task in which eye movements were not constrained during manual aiming. Experiment 3 confirmed the lack of manual RDE across a wider range of distractor delays (0, 50, 100, and 150?ms). Our data imply that the RDE is specific to the oculomotor system, at least for non-foveal distractors. We suggest that the oculomotor RDE reflects competitive interactions between target and distractor representations in the superior colliculus, which are not necessarily shared by manual aiming.