Mirror neuron activation as a function of explicit learning: changes in mu-event-related power after learning novel responses to ideomotor compatible, partially compatible, and non-compatible stimuli.

Questions regarding the malleability of the mirror neuron system (MNS) continue to be debated. MNS activation has been reported when people observe another person performing biological goal-directed behaviors, such as grasping a cup. These findings support the importance of mapping goal-directed biological behavior onto one's motor repertoire as a means of understanding the actions of others. Still, other evidence supports the Associative Sequence Learning (ASL) model which predicts that the MNS responds to a variety of stimuli after sensorimotor learning, not simply biological behavior. MNS activity develops as a consequence of developing stimulus-response associations between a stimulus and its motor outcome. Findings from the ideomotor literature indicate that stimuli that are more ideomotor compatible with a response are accompanied by an increase in response activation compared to less compatible stimuli; however, non-compatible stimuli robustly activate a constituent response after sensorimotor learning. Here, we measured changes in the mu-rhythm, an EEG marker thought to index MNS activity, predicting that stimuli that differ along dimensions of ideomotor compatibility should show changes in mirror neuron activation as participants learn the respective stimulus-response associations. We observed robust mu-suppression for ideomotor-compatible hand actions and partially compatible dot animations prior to learning; however, compatible stimuli showed greater mu-suppression than partially or non-compatible stimuli after explicit learning. Additionally, non-compatible abstract stimuli exceeded baseline only after participants explicitly learned the motor responses associated with the stimuli. We conclude that the empirical differences between the biological and ASL accounts of the MNS can be explained by Ideomotor Theory.

Motor-Evoked Potentials for Early Individual Elements of an Action Sequence During Planning Reflect Parallel Activation Processes.

Several computational models make predictions about the activation states of individual elements of an action sequence during planning and execution; however, the neural mechanisms of action planning are still poorly understood. Simple chaining models predict that only the first response in an action sequence should be active during planning. Conversely, some parallel activation models suggest that during planning, a serial inhibition process places the individual elements of the action into a serial order across a winner-takes-all competitive choice gradient in which earlier responses are more active, and hence likely to be selected for execution compared with later responses. We triggered transcranial magnetic stimulation pulses at 200 or 400 ms after the onset of a five-letter word, in which all but one response was planned and typed with the left hand, except for a single letter which required a right index finger response exclusively at one of five serial positions. We measured the resulting motor-evoked potentials at the right index finger as a marker for the activation state of that planned response. We observed no difference in motor-evoked potential amplitude across any serial position when a right index finger response was planned at 200 ms after the onset of the word; however, we observed a graded pattern of activation at 400 ms, with earlier positions that required a right index finger response showing greater motor-evoked potentials amplitude compared with later positions. These findings provide empirical support for competitive queuing computational models of action planning.

Dorsal stream activity and connectivity associated with action priming of ambiguous apparent motion.

Theories proposing a bidirectional influence between action and perception are well supported by behavioral findings. In contrast to the growing literature investigating the brain mechanisms by which perception influences action, there is a relative dearth of neural evidence documenting how action may influence perception. Here we show that action priming of apparent motion perception is associated with increased functional connectivity between dorsal cortical regions connecting vision with action. Participants manually rotated a joystick in a clockwise or counter-clockwise direction while viewing ambiguous apparent rotational motion. Actions influenced perception when the perceived direction of the ambiguous display was the same as manual rotation. For comparison, participants also rotated the joystick while viewing non-ambiguous apparent motion and in the absence of apparent motion. In a final control condition, participants viewed ambiguous apparent motion without manual rotation. Actions influence on perception was accompanied by a significant increase in alpha and beta band event related desynchronization (ERD) in contralateral primary motor cortex, superior parietal lobe and middle occipital gyrus. Increased ERD across these areas was accompanied by an increase in gamma band phase locking between primary motor, parietal, striate and extrastriate regions. Similar patterns were not observed when action was compatible with perception, but did not influence it. These data demonstrate that action influences perception by strengthening the interaction across a broad sensorimotor network for the putative purpose of integrating compatible action outcomes and sensory information into a single coherent percept.

The Effects of Textured Insoles on Cortical Activity and Quiet Bipedal Standing With and Without Vision: An EEG Study.

Wearing textured insoles (TIs) can reduce static postural sway, but the neurophysiological mechanisms by which these changes occur are not well understood. To address this issue, cortical activity was investigated in this study using electroencephalography (EEG) recordings from 19 scalp locations, in 15 healthy young adults (5 females; mean age = 27 ± 4.09 years) during quiet bipedal standing, under different insole conditions (textured versus smooth), with and without vision. Compared to smooth insoles (SIs), TIs significantly reduced postural sway in two measures; anterior-posterior range and standard deviation. In the EEG data, whole-head analyses showed cortical activity in the upper alpha power band was significantly reduced for textured compared to SIs. Exploratory analyses revealed this effect was significant both with and without vision, and was more pronounced over the parietal, compared to central regions, and over central compared to frontal regions. This trend was observed in low alpha and theta bands, but the effect of insole type was not significant. Textured insoles thus appear to affect not only balance outcomes but also cortical activity. The cortical activity adaptation may represent greater information becoming readily available at the cortical level, enhancing the representation of the body in space.

Parallel regulation of past, present, and future actions during sequencing.

Past, present, and future actions must be regulated online to produce sequences of actions, but the regulation process is not well understood because of measurement limitations. We provide the first direct tests of the parallel action regulation hypothesis during sequencing in humans. We used transcranial magnetic stimulation to probe the level of excitation for flexion of the right index finger during typing. Motor evoked potentials (MEPs) were recorded at the onset of typing 5-letter words and nonwords. A single letter typed by the right index finger varied across letter positions 1 to 5. MEP amplitude was largest for the upcoming action in the second position and decreased monotonically across future serial positions, suggesting a serial inhibition process regulates all future actions in parallel during sequencing. This is the most direct human evidence to date corroborating models of sequence production that assume parallel regulation of actions. (PsycINFO Database Record

Spatial knowledge during skilled action sequencing: Hierarchical versus nonhierarchical representations.

Typists can type 4 to 5 keystrokes per second at around 95% accuracy, yet they appear to have poor declarative knowledge of key locations. Logan and Crump (2011, Psychology of Learning and Motivation, Vol. 54, pp. 1-27) accounted for this paradox by proposing that typing is hierarchically organized into two loops, with an outer loop that transforms sentences into words and passes each word, one at a time, to an inner loop that transforms each word into its constituent keystrokes; however, the nature of the inner loop's spatial knowledge is not well understood. Key locations may be learned through the experiences of locating and traversing between keys. In daily life, people tend to type structured language, and, as a consequence, certain keys and key-to-key transitions are experienced more frequently than others. Here, we asked whether or not this knowledge is structured hierarchically. For example, knowledge of key locations may be nested within representations of words, or the inner loop may rely on knowledge that is independent from higher level structures. To test this, we had people type English, English-like, and random strings during normal, partially occluded, and occluded typing. In both partially occluded and occluded typing, error rates were higher while typing random strings compared to English and English-like strings, whereas there was no difference in error rates between English and English-like strings. This suggests that typists' spatial knowledge of the keyboard is not driven by hierarchical word-level representations, but instead is likely driven by a collection of individual processes, such as knowledge of the sequential structure of language acquired by typing more frequently occurring letters.

The dynamic range of response set activation during action sequencing.

We show that theories of response scheduling for sequential action can be discriminated on the basis of their predictions for the dynamic range of response set activation during sequencing, which refers to the momentary span of activation states for completed and to-be-completed actions in a response set. In particular, theories allow that future actions in a plan are partially activated, but differ with respect to the width of the range, which refers to the number of future actions that are partially activated. Similarly, theories differ on the width of the range for recently completed actions that are assumed to be rapidly deactivated or gradually deactivated in a passive fashion. We validate a new typing task for measuring momentary activation states of actions across a response set during action sequencing. Typists recruited from Amazon Mechanical Turk copied a paragraph by responding to a "go" signal that usually cued the next letter but sometimes cued a near-past or future letter (n-3, -2, -1, 0, +2, +3). The activation states for producing letters across go-signal positions can be inferred from RTs and errors. In general, we found evidence of graded parallel activation for future actions and rapid deactivation of more distal past actions. (PsycINFO Database Record

Working memory modulates neural efficiency over motor components during a novel action planning task: an EEG study.

Research shows neural efficiency of motor-related activity based on learning and expertise in a specific domain (e.g., guitar playing, sharp-shooting or a sport). However, it is unknown whether neural efficiency of motor-related activity, underlying action planning and maintenance, can be modulated by general cognitive ability alone. This study examined whether working memory span can influence motor-related neural activity during a novel motor task. Participants were divided into low- and high-span working memory groups based on their scores in an operation span task. Afterwards, participants learned different sequences of button responses corresponding to different abstract stimuli. The task required participants to briefly maintain an action plan in working memory to a stimulus that they would execute after responding to a subsequent stimulus. We used EEG to record changes in event related power in the mu- and beta-bands in left and right motor components during the interval where participants planned and maintained an action in working memory. Results showed decreases in mu- and beta-event related power for low-span participants and increases in mu- and beta-event related power for high-span participants over the left motor cluster while maintaining an action plan in working memory. Also, high-span participants were faster and more accurate in the task than low-span participants. This suggests that neural efficiency during a novel motor task can be influenced by working memory span, and that such differences are localized to the motor system.

Interference due to shared features between action plans is influenced by working memory span.

In this study, we examined the interactions between the action plans that we hold in memory and the actions that we carry out, asking whether the interference due to shared features between action plans is due to selection demands imposed on working memory. Individuals with low and high working memory spans learned arbitrary motor actions in response to two different visual events (A and B), presented in a serial order. They planned a response to the first event (A) and while maintaining this action plan in memory they then executed a speeded response to the second event (B). Afterward, they executed the action plan for the first event (A) maintained in memory. Speeded responses to the second event (B) were delayed when it shared an action feature (feature overlap) with the first event (A), relative to when it did not (no feature overlap). The size of the feature-overlap delay was greater for low-span than for high-span participants. This indicates that interference due to overlapping action plans is greater when fewer working memory resources are available, suggesting that this interference is due to selection demands imposed on working memory. Thus, working memory plays an important role in managing current and upcoming action plans, at least for newly learned tasks. Also, managing multiple action plans is compromised in individuals who have low versus high working memory spans.

Frequency of the first feature in action sequences influences feature binding.

We investigated whether binding among perception and action feature codes is a preliminary step toward creating a more durable memory trace of an action event. If so, increasing the frequency of a particular event (e.g., a stimulus requiring a movement with the left or right hand in an up or down direction) should increase the strength and speed of feature binding for this event. The results from two experiments, using a partial-repetition paradigm, confirmed that feature binding increased in strength and/or occurred earlier for a high-frequency (e.g., left hand moving up) than for a low-frequency (e.g., right hand moving down) event. Moreover, increasing the frequency of the first-specified feature in the action sequence alone (e.g., "left" hand) increased the strength and/or speed of action feature binding (e.g., between the "left" hand and movement in an "up" or "down" direction). The latter finding suggests an update to the theory of event coding, as not all features in the action sequence equally determine binding strength. We conclude that action planning involves serial binding of features in the order of action feature execution (i.e., associations among features are not bidirectional but are directional), which can lead to a more durable memory trace. This is consistent with physiological evidence suggesting that serial order is preserved in an action plan executed from memory and that the first feature in the action sequence may be critical in preserving this serial order.