主动控制感在大脑表征中的时空标记

主动控制感是主动动作过程中产生的控制自身动作, 进而控制外部环境的主观体验。构成动作主动控制感的核心要素是主观意图与结果反馈。本研究试图通过操控这两个核心要素的不同属性, 借助脑磁图等技术, 探寻主动控制感在大脑额-顶为主的脑网络中前-后馈的作用方式及时空特异性标记, 并建构新的认知神经理论模型。这将有利于理解人类动作的产生及后效、为相关精神类疾病的临床诊断提供更加客观的参照标准。     

The phenomenology of agency and intention in the face of paralysis and insentience

Studies of perception have focussed on sensation, though more recently the perception of action has, once more, become the subject of investigation. These studies have looked at acute experimental situations. The present paper discusses the subjective experience of those with either clinical syndromes of loss of movement or sensation (spinal cord injury, sensory neuronopathy syndrome or motor stroke), or with experimental paralysis or sensory loss. The differing phenomenology of these is explored and their effects on intention and agency discussed. It is shown that sensory loss can have effects on the focussing of motor command and that for some a sense of agency can return despite paralysis.

Feedback and intention during motor-skill learning: a connection with prospective memory

The intention to complete an action in the future can improve the learning of this action, but it is unknown whether this effect persists when feedback is manipulated during encoding. In Experiment 1, participants were instructed to learn a motor skill with or without intending to reproduce this learning in the future, and feedback on their movements was administrated by self-decision, that is, participants asked for feedback whenever they wanted it. The results showed that intention increased the frequency with which feedback was requested, but did not improve motor performance. In Experiment 2, participants had to learn the task with high or few feedbacks, which they could not control. In these conditions, intention was beneficial in promoting motor learning only for a low feedback schedule. We suggest that the beneficial effect of intention on learning can be overshadowed or emphasised by the feedback processing during encoding. These findings are discussed in light of theories surrounding prospective memory.     

How do we know what we are doing? Time, intention and awareness of action

Time is a fundamental dimension of consciousness. Many studies of the "sense of agency" have investigated whether we attribute actions to ourselves based on a conscious experience of intention occurring prior to action, or based on a reconstruction after the action itself has occurred. Here, we ask the same question about a lower level aspect of action experience, namely awareness of the detailed spatial form of a simple movement. Subjects reached for a target, which unpredictably jumped to the side on some trials. Participants (1) expressed their expectancy of a target shift during the upcoming movement, (2) pointed at the target as quickly and accurately as possible before returning to the start posiment to the target shift if required and (3) reproduced the spatial path of the movement they had just made, as accurately as possible, to give an indication of their awareness of the pointing movement. We analysed the spatial disparity between the initial and the reproduced movements on those with a target shift. A negative disparity value, or undershoot, suggests that motor awareness merely reflects a sluggish record of coordinated motor performance, while a positive value, or overshoot, suggests that participants' intention to point to the shifting target contributes more to their awareness of action than their actual pointing movement. Undershoot and overshoot thus measure the reconstructive (motoric) and the preconstuctive (intentional) aspects of action awareness, respectively. We found that trials on which subjects strongly expected a target shift showed greater overshoot and less undershoot than trials with lower expectancy. Conscious expectancy therefore strongly influences the experience of the detailed motor parameters of our actions. Further, a delay inserted either between the expectancy judgement and the pointing movement, or between the pointing movement and the reproduction of the movement, had no effect on visuomotor adjustment but strongly influenced action awareness. Delays during either interval boosted undershoots, suggesting increased reliance on a time-limited sensory memory for action. The experience of action is thus strongly influenced by prior thoughts and expectations, but only over a short time period. Thus, awareness of our actions is a dynamic and relatively flexible mixture of what we intend to do, and what our motor system actually does.     

Visual and haptic feedback contribute to tuning and online control during object manipulation

The authors employed a virtual environment to investigate how humans use haptic and visual feedback in a simple, rhythmic object-manipulation task. The authors hypothesized that feedback would help participants identify the appropriate resonant frequency and perform online control adjustments. The 1st test was whether sensory feedback is needed at all; the 2nd was whether the motor system combines visual and haptic feedback to improve performance. Task performance was quantified in terms of work performed on the virtual inertia, ability to identify the correct rhythm, and variability of movement. Strict feedforward control was found to be ineffective for this task, even when participants had previous knowledge of the rhythm. Participants (N = 11) performed far better when feedback was available (11 times more work, 2.2 times more precise frequency, 30% less variability; p < .05 for all 3 performance measures). Using sensory feedback, participants were able to rapidly identify 4 different spring-inertia systems without foreknowledge of the corresponding resonant frequencies. They performed over 20% more work with 24% less variability when provided with both visual and haptic feedback than they did with either feedback channel alone (p < .05), providing evidence that they integrated online sensory channels. Whereas feedforward control alone led to poor performance, feedback control led to fast tuning or calibration of control according to the resonant frequency of the object, and to better control of the rhythmic movement itself.     

Movements or targets: What makes an action in action–effect learning?

According to ideomotor theory, actions become linked to the sensory feedback they contingently produce, so that anticipating the feedback automatically evokes the action it typically results from. Numerous recent studies have provided evidence in favour of such action–effect learning but left an important issue unresolved. It remains unspecified to what extent action–effect learning is based on associating effect-representations to representations of the performed movements or to representations of the targets at which the behaviour aimed at. Two experiments were designed to clarify this issue. In an acquisition phase, participants learned the contingency between key presses and effect tones. In a following test phase, key–effect and movement–effect relations were orthogonally assessed by changing the hand–key mapping for one half of the participants. Experiment 1 showed precedence for target–effect over movement–effect learning in a forced-choice RT task. In Experiment 2, target–effect learning was also shown to influence the outcome of response selection in a free-choice task. Altogether, the data indicate that both movement–effect and target–effect associations contribute to the formation of action–effect linkages—provided that movements and targets are likewise contingently related to the effects.     

Changes in predictive motor control in drop-jumps based on uncertainties in task execution

Drop-jumps are controlled by predictive and reactive motor strategies which differ with respect to the utilization of sensory feedback. With reaction, sensory feedback is integrated while performing the task. With prediction, sensory information may be used prior to movement onset. Certainty about upcoming events is important for prediction. The present study aimed at investigating how uncertainties in the task execution affect predictive motor control in drop-jumps. Ten healthy subjects (22±1 years, M±SD) participated. The subjects performed either (i) drop-jumps by knowing that they might had to switch to a landing movement upon an auditory cue, which was sometimes elicited prior to touch-down (uncertainty). In (ii), subjects performed drop-jumps by knowing that there would be no auditory cue and consequently no switch of the movement (certainty). The m. soleus EMG prior to touch-down was higher when subjects knew there would be no auditory cue compared to when subjects performed the same task but switching from drop-jump to landing was possible (uncertainty). The EMG was reversed in the late concentric phase, meaning that it was higher in the high uncertainty task. The results of the present study showed that the muscular activity was predictively adjusted according to uncertainties in task execution. It is argued that tendomuscular stiffness was the variable responsible for the adjustment of muscular activity. The required tendomuscular stiffness was higher in drop-jumps than in landings. Consequently, when it was not certain whether to jump or to land, muscular activity and therefore tendomuscular stiffness was reduced.     

Interplay between action and movement intentions during social interaction

Observing the movements of another person influences the observer's intention to execute similar movements. However, little is known about how action intentions formed prior to movement planning influence this effect. In the experiment reported here, we manipulated the congruency of movement intentions and action intentions in a pair of jointly acting individuals (i.e., a participant paired with a confederate coactor) and investigated how congruency influenced performance. Overall, participants initiated actions faster when they had the same action intention as the coactor (i.e., when participants and the coactor were pursuing the same conceptual goal). Participants' responses were also faster when their and the coactor's movement intentions were directed to the same spatial location, but only when participants had the same action intention as the coactor. These findings suggest that observers use the same representation to implement their own action intentions that they use to infer other people's action intentions and also that a dynamic, multitiered intentional mechanism is involved in the processing of other people's actions.     

Effects of motor intention on the perception of somatosensory events: A behavioural and functional magnetic resonance imaging study

The intention to execute a movement can modulate our perception of sensory events, and this modulation is observed ahead of both ocular and upper limb movements. However, theoretical accounts of these effects, and also the empirical data, are often contradictory. Accounts of “active touch”, and the premotor theory of attention, have emphasized how movement intention leads to enhanced perceptual processing at the target of a movement, or on the to-be-moved effector. By contrast, recent theories of motor control emphasize how internal “forward” model (FM) estimates may be used to cancel or attenuate sensory signals that arise as a result of self-generated movements. We used behavioural and functional brain imaging (functional magnetic resonance imaging, fMRI) to investigate how perception of a somatosensory stimulus differed according to whether it was delivered to a hand that was about to execute a reaching movement or the alternative, nonmoving, hand. The results of our study demonstrate that a somatosensory stimulus delivered to a hand that is being prepared for movement is perceived to have occurred later than when that same stimulus is delivered to a nonmoving hand. This result indicates that it takes longer for a tactile stimulus to be detected when it is delivered to a moving limb and may correspond to a change in perceptual threshold. Our behavioural results are paralleled by the results of our fMRI study that demonstrated that there were significantly reduced blood-oxygen-level-dependent (BOLD) responses within the parietal operculum and insula following somatosensory stimulation of the hand being prepared for movement, compared to when an identical stimulus was delivered to a nonmoving hand. These findings are consistent with the prediction of FM accounts of motor control that postulate that central sensory suppression of somatosensation accompanies self-generated limb movements, and with previous reports indicating that effects of sensory suppression are observed in higher order somatosensory regions.     

Effects of motor intention on the perception of somatosensory events: a behavioural and functional magnetic resonance imaging study

The intention to execute a movement can modulate our perception of sensory events, and this modulation is observed ahead of both ocular and upper limb movements. However, theoretical accounts of these effects, and also the empirical data, are often contradictory. Accounts of "active touch", and the premotor theory of attention, have emphasized how movement intention leads to enhanced perceptual processing at the target of a movement, or on the to-be-moved effector. By contrast, recent theories of motor control emphasize how internal "forward" model (FM) estimates may be used to cancel or attenuate sensory signals that arise as a result of self-generated movements. We used behavioural and functional brain imaging (functional magnetic resonance imaging, fMRI) to investigate how perception of a somatosensory stimulus differed according to whether it was delivered to a hand that was about to execute a reaching movement or the alternative, nonmoving, hand. The results of our study demonstrate that a somatosensory stimulus delivered to a hand that is being prepared for movement is perceived to have occurred later than when that same stimulus is delivered to a nonmoving hand. This result indicates that it takes longer for a tactile stimulus to be detected when it is delivered to a moving limb and may correspond to a change in perceptual threshold. Our behavioural results are paralleled by the results of our fMRI study that demonstrated that there were significantly reduced blood-oxygen-level-dependent (BOLD) responses within the parietal operculum and insula following somatosensory stimulation of the hand being prepared for movement, compared to when an identical stimulus was delivered to a nonmoving hand. These findings are consistent with the prediction of FM accounts of motor control that postulate that central sensory suppression of somatosensation accompanies self-generated limb movements, and with previous reports indicating that effects of sensory suppression are observed in higher order somatosensory regions.     

Effect of distorted visual feedback on the sense of agency

It has been hypothesized that an internal model is involved in controlling and recognizing one's own actions (action attribution). This results from a comparison process between the predicted sensory feedback of the action and its real sensory consequences. The aim of the present study is to distinguish the respective importance of two action parameters (time and direction) on such an attribution judgment. We used a device that allows introduction of discordance between the movements actually performed and the sensory feedback displayed on a computer screen. Participants were asked to judge whether they were viewing 1) their own movements, 2) their own movements modified (spatially or temporally displaced), or 3) those of another agent (i.e, the experimenter). In fact, in all conditions they were only shown their own movements either unaltered or modified by varying amounts in space or time. Movements were only attributed to another agent when there was a high spatial discordance between participants' hand movements and sensory feedback. This study is the first to show that the direction of movements is a cardinal feature in action attribution, whereas temporal properties of movements play a less important role.     

On the signals underlying conscious awareness of action

To investigate whether conscious judgments of movement onset are based solely on pre-movement signals (i.e., premotor or efference copy signals) or whether sensory feedback (i.e., reafferent) signals also play a role, participants judged the onset of finger and toe movements that were either active (i.e., self initiated) or passive (i.e., initiated by the experimenter). Conscious judgments were made by reporting the position of a rotating clock hand presented on a computer screen and were then compared to the actual measured time of movement onset. In line with previous studies, judgment errors were found to be anticipatory for both finger and toe movements. There was a significant difference between judgment errors for active and passive movements, with judgments of active movements being more anticipatory than judgments of passive movements. This is consistent with a pre-movement (from here on referred to as an “efferent”) account of action awareness because premotor and efference copy signals are only present in active movements, whereas the main source of movement information in passive movements is sensory feedback which is subject to time delays of conduction (and hence predicts later judgment times for passive movements). However, judgments of active toe movement onset time were less anticipatory than judgments of active finger movement onset time. This pattern of results is not consistent with a pure efferent account of conscious awareness of action onset - as this account predicts more anticipatory judgments for toe movements compared to finger movements. Instead, the data support the idea that conscious judgments of movement onset are based on efferent (i.e., premotor, efference copy) and reafferent (i.e., feedback from the movement) components.     

Biases in the subjective timing of perceptual events: Libet et al. (1983) revisited

We report two experiments in which participants had to judge the time of occurrence of a stimulus relative to a clock. The experiments were based on the control condition used by Libet, Gleason, Wright, and Pearl [Libet, B., Gleason, C. A., Wright, E. W., & Pearl, D. K. (1983). Time of conscious intention to act in relation to onset of cerebral activities (readiness-potential): The unconscious initiation of a freely voluntary act. Brain 106, 623-642] to correct for any bias in the estimation of the time at which an endogenous event, the conscious intention to perform a movement, occurred. Participants' responses were affected systematically by the sensory modality of the stimulus and by the speed of the clock. Such findings demonstrate the variability in judging the time at which an exogenous event occurs and, by extension, suggest that such variability may also apply to the judging the time of occurrence of endogenous events. The reliability of participants' estimations of when they formed the conscious intention to perform a movement in Libet et al.'s (1983) study is therefore questionable.     

Reduced timing variability during bimanual coupling: A role for sensory information

On a repetitive tapping task, the within-hand variability of intertap intervals is reduced when participants tap with two hands as compared to one-hand tapping. Because this bimanual advantage can be attributed to timer variance (Wing-Kristofferson model, 1973a, b), separate timers have been proposed for each hand, whose outputs are then averaged (Helmuth & Ivry, 1996). An alternative notion is that action timing is based on its sensory reafferences (Aschersleben & Prinz, 1995; Prinz, 1990). The bimanual advantage is then due to increased sensory reafference. We studied bimanual tapping with the continuation paradigm. Participants first synchronized their taps with a metronome and then continued without the pacing signal. Experiment 1 replicated the bimanual advantage. Experiment 2 examined the influence of additional sensory reafferences. Results showed a reduction of timer variance for both uni- and bimanual tapping when auditory feedback was added to each tap. Experiment 3 showed that the bimanual advantage decreased when auditory feedback was removed from taps with the left hand. Results indicate that the sensory reafferences of both hands are used and integrated into timing. This is consistent with the assumption that the bimanual advantage is at least partly due to the increase in sensory reafference. A reformulation of the Wing-Kristofferson model is proposed to explain these results, in which the timer provides action goals in terms of sensory reafferences.     

Stimulation-induced behavioral inhibition: A new model for understanding physical violence

Physical violence is widely considered to result from action carried out with the intention of causing injury; that is, from aggression. However, the “hypothesis” of aggression is inapplicable in all but a few instances as well as inappropriate for many destructive rage-associated responses directed at inanimate objects. This paper outlines a new perspective on physical violence, reinterpreting many behaviors hitherto labeled aggressive as stimulation-seeking behaviors (SSBs) above an arbitrary level of intensity. It is further proposed that: 1) physical violence is a by-product of SSB, driven, in part by brain catecholaminergic (CA) systems, and the direct result of exchanges of energy that exceed the body's tolerance threshold; 2) allegedly discrete categories of motor-motivational behavior represent overlapping bands of intensity on a continuous spectrum of SSB; and 3) the sensory input derived from SSB is fed back into the central nervous system where it activates, brain serotonergic and/or cholinergic systems, which in turn inhibit CA systems, resulting in a general state of behavioral quiescence. In addition to accounting for a number of previously unexplained observations, the model suggests that physical violence could be prevented by providing groups at high risk with extensive opportunities for therapeutic sensory stimulation to substitute for that derived from excessive SSB. For people at especially high risk, portable devices could be developed that would allow the user to self-administer desired levels of sensory stimulation at moments of intense anger, thereby preventing potentially dangerous outbursts of SSB prior to the onset of the behavior.     

Stimulation-induced behavioral inhibition: a new model for understanding physical violence.

Physical violence is widely considered to result from action carried out with the intention of causing injury; that is, from aggression. However, the "hypothesis" of aggression is inapplicable in all but a few instances as well as inappropriate for many destructive, rage-associated responses directed at inanimate objects. This paper outlines a new perspective on physical violence, reinterpreting many behaviors hitherto labeled aggressive as stimulation-seeking behaviors (SSBs) above an arbitrary level of intensity. It is further proposed that: 1) physical violence is a by-product of SSB, driven in part by brain catecholaminergic (CA) systems, and the direct result of exchanges of energy that exceed the body's tolerance threshold; 2) allegedly discrete categories of motor-motivational behavior represent overlapping bands of intensity on a continuous spectrum of SSB; and 3) the sensory input derived from SSB is fed back into the central nervous system where it activates brain serotonergic and/or cholinergic systems, which in turn inhibit CA systems, resulting in a general state of behavioral quiescence. In addition to accounting for a number of previously unexplained observations, the model suggests that physical violence could be prevented by providing groups at high risk with extensive opportunities for therapeutic sensory stimulation to substitute for that derived from excessive SSB. For people at especially high risk, portable devices could be developed that would allow the user to self-administer desired levels of sensory stimulation at moments of intense anger, thereby preventing potentially dangerous outbursts of SSB prior to the onset of the behavior.     

Spatial and Temporal Adaptations That Accompany Increasing Catching Performance During Learning

During stance, head extension increases postural sway, possibly due to interference with sensory feedback. The sit-to-stand movement is potentially destabilizing due to the development of momentum as the trunk flexes forward and the body transitions to a smaller base of support. It is unclear what role head orientation plays in the postural and movement characteristics of the sit-to-stand transition. The authors assessed how moving from sitting to standing with head-on-trunk extension compared with moving with the head neutral or flexed, or with moving with the head facing forward in space (which would involve head-on-trunk extension, but not head-in-space extension) in healthy, young participants. Head-on-trunk extension increased center of pressure variability, but decreased movement velocities, movement duration, and trunk flexion compared with flexed and neutral head-on-trunk orientations. Similarities in movement characteristics between head-on-trunk extension and the forward head-in-space orientation suggest that stabilizing the head in space does not fully counteract the postural and movement changes due to head-on-trunk extension. Findings suggest that proprioceptive feedback from the neck muscles contributes to the regulation of posture and movement, and therefore should not be overlooked in research on the role of sensory feedback in postural control.     

Is head-on-trunk extension a proprioceptive mediator of postural control and sit-to-stand movement characteristics?

During stance, head extension increases postural sway, possibly due to interference with sensory feedback. The sit-to-stand movement is potentially destabilizing due to the development of momentum as the trunk flexes forward and the body transitions to a smaller base of support. It is unclear what role head orientation plays in the postural and movement characteristics of the sit-to-stand transition. The authors assessed how moving from sitting to standing with head-on-trunk extension compared with moving with the head neutral or flexed, or with moving with the head facing forward in space (which would involve head-on-trunk extension, but not head-in-space extension) in healthy, young participants. Head-on-trunk extension increased center of pressure variability, but decreased movement velocities, movement duration, and trunk flexion compared with flexed and neutral head-on-trunk orientations. Similarities in movement characteristics between head-on-trunk extension and the forward head-in-space orientation suggest that stabilizing the head in space does not fully counteract the postural and movement changes due to head-on-trunk extension. Findings suggest that proprioceptive feedback from the neck muscles contributes to the regulation of posture and movement, and therefore should not be overlooked in research on the role of sensory feedback in postural control.     

Vision,proprioception and corollary discharge in a movement recall test

Movement recall was investigated in relation to the sensory processes involved in a triangle drawing task.Forty subjects in two groups, one with and one without visual feedback, performed a recall task involving movements of their index finger. All subjects attended different experimental sessions in which (1) all proprioceptive feedback was eliminated by the ischaemic block technique, (2) muscle spindle feedback was distorted by vibration of the muscles and tendons involved in the movement, and (3) proprioceptive feedback was normal.Within each session subjects were required firstly to recall triangular movements made for them passively by the experimenter, and secondly, to recall movements they had made actively. Results indicated comparable accuracy in recall of active movements in all conditions, and a decrement in passive recall dependent on the availability of the alternative sources of feedback. The results indicated a process of integrated contribution of all inputs to the perception of movement; redundancy in information when all channels are available; and a role of corollary discharge in recall of movements.     

Impaired embodiment and intersubjectivity

This paper considers the importance of the body for self-esteem, communication, and emotional expression and experience, through the reflections of those who live with various neurological impairments of movement and sensation; sensory deafferentation, spinal cord injury and Möbius Syndrome (the congenital absence of facial expression). People with severe sensory loss, who require conscious attention and visual feedback for movement, describe the imperative to use the same strategies to reacquire gesture, to appear normal and have embodied expression. Those paralysed after spinal cord injury struggle to have others see them as people rather than as people in wheelchairs and have been active in the disability movement, distinguishing between their medical impairment and the social induced disability others project onto them. Lastly those with Möbius reveal the importance of the face for emotional expression and communication and indeed for emotional experience itself. All these examples explore the crucial role of the body as agent for social and personal expression and self-esteem.