EEG dipole analysis of motor-priming foreperiod activity reveals separate sources for motor and spatial attention components

OBJECTIVE: This study employed EEG source localisation procedures to study the contribution of motor preparatory and attentional processing to foreperiod activity in an S1-S2 motor priming task. METHODS: Behavioural and high-density event-related potential (ERP) data were recorded in an S1-S2 priming task where participants responded to S2 with a left or right-hand button press. S1 either provided information about response hand (informative) or ambiguous information (uninformative). RESULTS: Responses were significantly faster in informative trials compared with uninformative trials. Dipole source analysis of foreperiod lateralized ERPs revealed sources of motor preparatory activity in the dorsolateral premotor cortex (PMd) in line with previous work. In addition, two spatial attention components (ADAN, LDAP) were identified with generators in the PMd and occipitotemporal visual areas in the middle temporal (MT) region, respectively. Separation of motor-related and attentional PMd source locations was reliable along the rostral-caudal axis. CONCLUSIONS: The presence of attentional components in a motor priming paradigm supports the premotor theory of attention which suggests a close link between attention and motor preparatory processes. Separation of components in the premotor cortex is in accord with a functional division of PMd into rostral (higher-order processing) and caudal (motor-related processing) areas as suggested by imaging work. SIGNIFICANCE: A prime for response preparation is a trigger for separate, but closely linked, attention-related activity in premotor areas.     

ERP correlates of shared control mechanisms involved in saccade preparation and in covert attention

We investigated whether attention shifts and eye movement preparation are mediated by shared control mechanisms, as claimed by the premotor theory of attention. ERPs were recorded in three tasks where directional cues presented at the beginning of each trial instructed participants to direct their attention to the cued side without eye movements (Covert task), to prepare an eye movement in the cued direction without attention shifts (Saccade task) or both (Combined task). A peripheral visual Go/Nogo stimulus that was presented 800 ms after cue onset signalled whether responses had to be executed or withheld. Lateralised ERP components triggered during the cue-target interval, which are assumed to reflect preparatory control mechanisms that mediate attentional orienting, were very similar across tasks. They were also present in the Saccade task, which was designed to discourage any concomitant covert attention shifts. These results support the hypothesis that saccade preparation and attentional orienting are implemented by common control structures. There were however systematic differences in the impact of eye movement programming and covert attention on ERPs triggered in response to visual stimuli at cued versus uncued locations. It is concluded that, although the preparatory processes underlying saccade programming and covert attentional orienting may be based on common mechanisms, they nevertheless differ in their spatially specific effects on visual information processing.     

Modifying the cortical processing for motor preparation by repetitive transcranial magnetic stimulation

To investigate the effects of repetitive transcranial magnetic stimulation (rTMS) on the central processing of motor preparation, we had subjects perform a precued-choice reaction time (RT) task. They had to press one of two buttons as quickly as possible after a go signal specifying both the hand to be used and the button to press. A precue preceding this signal conveyed full, partial, or no advance information (hand and/or button), such that RT shortened with increasing amount of information. We applied 1200 to 2400 pulses of 1-Hz rTMS over various cortical areas and compared the subjects' performances at various times before and after this intervention. rTMS delayed RT at two distinct phases after stimulation, one within 10 min and another with a peak at 20 to 30 min and lasting for 60 to 90 min, with no significant effects on error rates or movement time. The effect was significantly larger on left- than on right-hand responses. RT was prominently delayed over the premotor and motor cortices with similar effects across different conditions of advance information, suggesting that preparatory processes relatively close to the formation of motor output were influenced by rTMS. In contrast, the effect of rTMS over the supplementary motor area and the anterior parietal cortex varied with the amount of advance information, indicating specific roles played by these areas in integrating target and effector information. The primary motor area, especially of the left hemisphere, may take over this processing, implementing motor output based on the information processed in other areas.     

Prior information of stimulus location: effects on ERP measures of visual selection and response selection

This paper examines the effects of prior information of the location of an upcoming stimulus on event-related EEG potentials associated with the focusing of attention. Results of two tasks, reported in a previous publication (Praamstra, P., Boutsen, L., Humphreys, G.W., 2005. Frontoparietal control of spatial attention and motor intention in human EEG. J. Neurophysiol. 94, 764-774), were compared: one in which spatial attention was cued to the stimulus location and one in which the cue was non-informative. Only informative directional cues elicited directing-attention EEG potentials in the delay period between cue and target. Notwithstanding these electrophysiological signs of an attentional orientation prior to the occurrence of the target, there were no reaction time effects related to the presence of advance spatial information. By contrast, the advance information did have effects on EEG potentials following the target stimulus. The N2pc, reflecting an attentional selection mechanism in extrastriate cortex, was reduced in amplitude with advance spatial information. The N2cc, coinciding in time with the N2pc but measured over the motor cortex, was preempted by the advance spatial information. These results support that the N2cc is not due to overlap of the N2pc with movement execution-related activity. It is proposed that the neural activity underlying this EEG potential arises from the dorsal premotor cortex and serves an executive-attentional function that helps to ensure that the selection of a manual response is not biased by the direction of spatial attention.     

Impairment of willed actions and use of advance information for movement preparation in schizophrenia

OBJECTIVES: To assess willed actions in patients with schizophrenia using reaction time (RT) tasks that differ in the degree to which they involve volitionally controlled versus stimulus driven responses. METHODS: Ten patients diagnosed with schizophrenia and 13 normal controls of comparable age were tested. Subjects performed a visual simple RT (SRT), an uncued four choice reaction time (CRT), and a fully cued four choice RT task. A stimulus 1(S1)-stimulus 2(S2) paradigm was used. The warning signal/precue (S1) preceded the imperative stimulus (S2) by either 0 (no warning signal or precue) 200, 800, 1600, or 3200 ms. RESULTS: The patients with schizophrenia had significantly slower RTs and movement times than normal subjects across all RT tasks. The unwarned SRT trials were significantly faster than the uncued CRT trials for both groups. For both groups, fully cued CRTs were significantly faster than the uncued CRTs. However, the S1-S2 interval had a differential effect on CRTs in the two groups. For the normal subjects fully cued CRTs and SRTs were equivalent when S1-S2 intervals were 800 ms or longer. A similar pattern of effects was not seen in the patients with schizophrenia, for whom the fully cued CRT were unexpectedly equivalent to SRT for the 200 ms interval and expectedly for the 1600 ms S1-S2 interval, but not the 3200 or 800 ms intervals. CONCLUSIONS: Patients with schizophrenia were able to use advance information inherent in SRT or provided by the precue in fully cued CRT to speed up RT relative to uncued CRT. However, in the latter task, in which the volitional demands of preprogramming are higher since a different response has to be prepared on each trial, patients showed some unusual and inconsistent interval effects suggesting instability of attentional set. It is possible that future studies using RT tasks with higher volitional demands in patients with predominance of negative signs may disclose greater deficits in willed action in schizophrenia.     

Changes in electromyographic responses to muscle stretch, related to the programming of movement parameters.

Three experiments are reported that used the advance information paradigm which consists of providing subjects with either no or partial information about an upcoming movement. Subjects moved handles to control the vertical displacements of CRT beams, to point to eight targets. The illumination of different combinations of these targets prior to movement execution provided advance information about which hand, movement direction, or movement extent would be required. Reaction time (RT), integrated EMG activity in the forearm extensor and flexor muscles, and M1, M2, and M3 components of the stretch reflex responses triggered in these muscles were analysed as a function of the precued movement parameter. Compared to the no-information condition, RT decreased in all precue conditions; however, the reduction was greater when direction than when hand was precued, and greater when hand than extent was precued. The EMG activity of forearm muscles increased during the preparatory period in all precue conditions, but generally did not differ among them. An overall facilitation of the stretch reflex components was observed in all precue conditions. This facilitation: (1) was greater for flexor than extensor muscles, (2) was similar regardless of the degree of extent precued, (3) differed for the M2 and M3 components depending on whether the responding hand precued was ipsilateral or contralateral. When the precued movement direction was considered, similar changes in the M3 component were found in extensor and flexor muscles. M3 was facilitated when the muscle was precued as an agonist and was inhibited when it was precued as an antagonist. Collectively these data provide support for a motor programming conception of movement organization.     

Expectancy induces dynamic modulation of corticospinal excitability

Behavioral studies using motor preparation paradigms have revealed that increased expectancy of a response signal shortens reaction times (RTs). Neurophysiological data suggest that in such paradigms, not only RT but also neuronal activity in the motor structures involved is modulated by expectancy of behaviorally relevant events. Here, we directly tested whether expectancy of a response signal modulates excitability of the corticospinal system used in the subsequent movement. We combined single- and paired-pulse transcranial magnetic stimulation (TMS) over the primary motor cortex with a simple RT task with variable preparatory delays. We found that, in line with typical behavioral observations, the subjects' RTs decreased with increasing response signal expectancy. TMS results revealed a modulation of corticospinal excitability in correspondence with response signal expectancy. Besides an increased excitability over the time-course of the preparatory delay, corticospinal excitability transiently increased whenever a response signal was expected. Paired-pulse TMS showed that this modulation is unlikely to be mediated by excitability changes in interneuronal inhibitory or facilitatory networks in the primary motor cortex. Changes in corticospinal synchronization or other mechanisms involving spinal circuits are candidates mediating the modulation of corticospinal excitability by expectancy.     

Anticipatory changes of long-latency stretch responses during preparation for directional hand movements

We have investigated, in 9 normal subjects, the time course of amplitude changes in the automatic long latency stretch reflex of wrist flexors during the preparatory period as a function of the precued direction of hand movement. Subjects maintained right hand position against a weak force and learned to compensate precisely for brief ramp stretch (250 deg./s, 50 ms) which imposed wrist extension. This probe stretch was applied randomly at different times during the 1 s preparatory period of a reaction time task. The warning signal gave directional advance information (DAI) about the voluntary movement that had to be performed. The EMG activity, recorded between 20 and 80 ms after the stretch started, was measured in terms of 3 successive components, M1, M2 and M3, identified on the basis of their respective latency.There was no significant change in the M1 component. Following a warning signal which precued an extension movement, M2 was depressed prior to the response signal. The time course of M3 was clearly different according to DAI: it increased following a warning signal which precued a flexion movement and decreased in the alternative case. This difference reached statistical significance 400 ms before the response signal. In fast-performing subjects the differential development of M3 was more marked than for slow-performing subjects. This underlines its preparatory significance. These results suggest that the neuronal pathways involved in the M3 response to stretch: i, are partly different from those conveying the earlier components, and ii, include structures which take part in the pre-programming of rapid movements.     

Does your gaze direction and head orientation shift my visual attention?

The effects of another person's gaze direction and head orientation on the observer's attentional processes were investigated. Subjects responded to visual, laterally presented reaction signals. The presentation of the reaction signal was preceded by a facial cue stimulus signaling a direction which was either congruent, neutral, or incongruent with the laterality of the reaction signal. A head (front and profile views) with an averted gaze affected the response times in comparison to the front view of a face with a straight gaze. In contrast, a profile view of a head with a compatible gaze direction did not result in such an effect. The results indicate that visual information from the other individual's gaze direction and head orientation is integrated, and the integrated information is fed to the brain areas subserving visual attention orienting.     

Attentional modulation of reward processing in the human brain

Although neural signals of reward anticipation have been studied extensively, the functional relationship between reward and attention has remained unclear: Neural signals implicated in reward processing could either reflect attentional biases towards motivationally salient stimuli, or proceed independently of attentional processes. Here, we sought to disentangle reward and attention‐related neural processes by independently modulating reward value and attentional task demands in a functional magnetic resonance imaging study in healthy human participants. During presentation of a visual reward cue that indicated whether monetary reward could be obtained in a subsequent reaction time task, participants either attended to the reward cue or performed an unrelated attention‐demanding task at two different levels of difficulty. In ventral striatum and ventral tegmental area, neural responses were modulated by reward anticipation irrespective of attentional demands, thus indicating attention‐independent processing of reward cues. By contrast, additive effects of reward and attention were observed in visual cortex. Critically, reward‐related activations in right anterior insula strongly depended on attention to the reward cue. Dynamic causal modelling revealed that the attentional modulation of reward processing in insular cortex was mediated by enhanced effective connectivity from ventral striatum to anterior insula. Our results provide evidence for distinct functional roles of the brain regions involved in the processing of reward‐indicating information: While subcortical structures signal the motivational salience of reward cues even when attention is fully engaged elsewhere, reward‐related responses in anterior insula depend on available attentional resources, likely reflecting the conscious evaluation of sensory information with respect to motivational value. Hum Brain Mapp 35:3036–3051, 2014. © 2013 Wiley Periodicals, Inc.     

Crossing the hands disrupts tactile spatial attention but not motor attention: evidence from event-related potentials

During covert shifts of tactile spatial attention both somatotopic and external reference frames are employed to encode hand location. When participants cross their hands these frames of references produce conflicting spatial codes which disrupt tactile attentional selectivity. Because attentional shifts are triggered not only in Attention tasks but also during covert movement preparation, the present study aimed at investigating the reference frame employed during such 'motor shifts of attention'. Event related brain potentials (ERPs) were recorded during a Motor task where a visual cue (S1) indicated the relevant hand for a manual movement prior to a tactile Go/Nogo stimulus (S2). For comparison, we ran a tactile Attention task where the same cue (S1) now indicated the relevant hand for a tactile discrimination (S2). Both tasks were performed under uncrossed and crossed hands conditions. In both Attention and Motor tasks similar lateralized components were observed following S1 presentation. Anterior and posterior ERP components indicative of covert attention shifts were exclusively guided by an external reference frame, while a later central negativity operated according to a somatotopic reference frame in both tasks. In the Motor task, this negativity reflected selective activation of the motor cortex in preparation for movement execution. In the Attention task, this component might reflect activity in the somatosensory cortex in preparation for the subsequent tactile discrimination. The presence of multiple and conflicting spatial codes resulted in disruption of tactile attentional selection in the Attention task where attentional modulations of tactile processing were delayed and attenuated with crossed hands as indicated by the analysis of ERPs elicited by S2. In contrast, attentional modulations of S2 processing in the Motor task were largely unaffected by the hand posture manipulation, suggesting that motor attention employs primarily one spatial coordinate system.     

Task-specific sensory and motor preparatory activation revealed by contingent magnetic variation

The present report studied the magnetic counterpart (CMV) of the auditory contingent negative variation (CNV). The ear where the target auditory stimulus would be presented was cued with a visual central arrow at a validity of 84%. The subject's behavioral response and the magnetoencephalographic (MEG) and electroencephalographic (EEG) signals were recorded. The central cue diminished reaction times (RTs) to the auditory target in the valid conditions with respect to the invalid conditions, indicating that the attentional manipulation was effective. The averaged magnetic field power during the preparatory period was significantly higher than baseline, suggesting the simultaneous presence of a magnetic counterpart of the electric CNV--the CMV. The field maps of the CMV grand averages showed two different and well-established periods: an early one with a magnetic field distribution that suggests a central source, and a late one with a field topography comparable to a low-intensity auditory-evoked field (M1). Single-dipole analysis of the preparatory phase in the subject's magnetic resonance images (MRI) demonstrated the presence of dipolar activity in the posterior cingulate (PCC) and posterior parietal cortices (PPC), superior temporal gyrus (STG) and motor cortices (MC). The lateralization of this activity depended on the orientation of the central cue. These results suggest that the action and perceptual-related areas needed to process the expected subsequent imperative task are recruited during the preparatory periods, influencing the behavioral RTs.     

Orienting visuospatial attention generates manual reaction time asymmetries in target detection and pointing

Right-handers exhibit a left hand advantage in response preparation when pointing to targets. These manual asymmetries are generally attributed to a right hemisphere specialization for spatial processing. More precisely, the left hand reaction time (RT) advantage was recently supposed to reflect specifically the right hemisphere superiority for movement planning. This study proposes to investigate a possible attentional origin for manual RT asymmetries. In a first experiment, we used the covert orienting of attention paradigm to measure subjects' RTs when reaching at targets (pointing task) both in valid, neutral and invalid conditions, either in the left or in the right visual fields and with the left and the right hand. In a second experiment, we applied the same paradigm to a detection task (key-pressing). Results revealed that orienting of attention to spatial locations was more time consuming when responding with the right than with the left hand, whether movement planning was required or not. It is suggested that the right hemisphere dominance for orienting of visuospatial attention account, partly at least, for the RT asymmetries classically observed in manual aiming.     

Preshaping and continuous evolution of motor cortical representations during movement preparation

While a goal-directed movement is prepared, motor cortical neurons selectively change their activity in relation to prior information about movement direction. Only little is known, however, about the neuronal representation of partial information about this parameter. We investigated this question by training monkeys in a multidirectional centre-out pointing task. A preparatory signal provided prior information about one, two or three possible adjacent targets, thus manipulating the level of certainty about movement direction. After a 1-s delay, the response signal specified one of the precued targets to indicate the actual movement to be performed. Based on the directional tuning curves of individual motor cortical neurons determined during the reaction time interval, we constructed distributions of the population activation (DPAs), which we were then able to estimate as well during the preparatory period. We found that these distributions were preshaped by prior information, with peaks of activation centred over the range of precued movement directions. These peaks sharpened as the response signal approached, and shifted to the specified movement direction subsequent to that signal. Wider ranges of precued movement directions were represented by broader DPAs. Trials in which monkeys produced short reaction times were characterized by narrower distributions than trials with long reaction times. Our study thus provides evidence for (i) a graded preshaping of the neuronal population representation of movement direction by partial information about this parameter, and (ii) the continuous evolution of the preshaped population representation during the preparatory period towards movement initiation.     

Attention, intention, and strategy in preparatory control

The neural mechanisms underlying different forms of preparatory control were examined using event-related fMRI. Preparatory brain activation was monitored in relation to different types of advance information: (1) random task cues indicating which of two possible tasks to perform upon subsequent target presentation; (2) task-ambiguous target stimuli; or (3) targets for which the correct response could be pre-determined. Three types of activation pattern were observed in different brain regions. First, more posterior regions of lateral prefrontal cortex (LPFC) and parietal cortex were activated by both advance task cues and advance targets, but with increased and more sustained activation for the latter. Second, more anterior regions of LPFC and parietal cortex were selectively activated by advance targets. Importantly, in these regions preparatory activation was not further modulated by the availability of advance response information. In contrast, preparatory activation in a third set of brain regions, including medial frontal cortex, reflected the utilization of advance response information, but by only a subset of participants. These results suggest three types of preparatory control: attentional (stimulus-oriented), intentional (action-oriented), and a possibly strategic component that might determine inter-individual differences in response readiness. Notably, the absence of regions selectively or even preferentially activated during cue-based preparation argues against certain conceptualizations of task-selective attention under cued task-switching conditions.     

Covert manual response preparation triggers attentional shifts: ERP evidence for the premotor theory of attention

The premotor theory of attention claims that the preparation of goal-directed action and shifts of attention are closely linked, because they are controlled by shared sensorymotor mechanisms. Until now, support for this theory has come primarily from studies demonstrating links between saccade programming and attention shifts. The present event-related brain potential (ERP) study demonstrated that attentional orienting processes are also elicited during the covert preparation of unimanual responses. ERPs were recorded in the interval between a visual response-hand selection cue and a subsequent visual Go/Nogo signal when participants prepared to lift their left or right index finger. Lateralised ERP components elicited during response preparation were very similar to components previously observed during instructed endogenous attention shifts, indicating that analogous attentional orienting processes are activated in both cases. Somatosensory ERP components (P90, N140) were enhanced when task-irrelevant tactile probes were delivered during response preparation to the hand involved in an anticipated response, even when probes were presented well in advance of response execution. These results suggest that attentional shifts are triggered during unimanual response preparation, as predicted by the premotor theory. This link between manual response programming and attention is consistent with the hypothesis that common mechanisms are involved in the control of attention and action.     

Dissociating effector and movement direction selection during the preparation of manual reaching movements: evidence from lateralized ERP components.

OBJECTIVE: The present study investigated whether lateralized ERP components triggered during covert manual response preparation (ADAN, LDAP) reflect effector selection, the selection of movement direction, or both. METHODS: Event-related brain potentials were recorded during a response precueing paradigm where visual cues provided either partial (Experiment 1) or full (Experiment 2) information about the response hand and the direction for a subsequent reaching movement. RESULTS: ADAN and LDAP components were elicited even when only partial response information was available, demonstrating that they do not require the presence of a fully specified motor program. The ADAN was elicited in a similar fashion regardless of whether effector or movement direction information was provided, suggesting that the underlying mechanisms are equally sensitive to both types of response-related information. In contrast, the LDAP was larger in response to cues providing effector information, but was also reliably present when movement direction was available. CONCLUSIONS: ADAN and LDAP components reflect preparatory activity within anterior and posterior parts of the parieto-premotor sensorimotor network where different parameters for manual reaching movements are programmed independently. SIGNIFICANCE: These results support the claim of the premotor theory of attention that shared sensorimotor control mechanisms are involved in attention and motor programming.     

Fear-conditioned cues of impending pain facilitate attentional engagement.

AIMS OF STUDY: Selective attention to signals of impending pain allows the avoidance of bodily harm. In order to identify the attentional components involved in the selection of pain signals over competing demands, we used an emotional modification of an exogenous cueing task. METHODS: Fifty-two pain-free volunteers detected visual targets of which the location was correctly or incorrectly predicted by a spatial cue. Cues were emotionally modulated using differential classical conditioning. The conditioned cue (CS+) was sometimes followed by an electrocutaneous stimulus (UCS), thus becoming a pain signal, whereas the UCS never followed the other cue (CS-), referred to as safety signal. RESULTS: Analyses of response times showed that pain signals facilitated the directing of attention to their location in comparison to safety signals. In contrast, pain signals did not impair disengagement of attention from their location in comparison to safety signals. CONCLUSION: It is concluded that attention is more strongly engaged to a signal of impending pain compared with a cue signalling its absence. We explore why disengagement from the pain signal is not impaired compared to the safety signal. The findings are discussed in terms of the defensive importance of pain anticipation.     

Blocking of irrelevant memories by posterior alpha activity boosts memory encoding

In our daily lives, we are confronted with a large amount of information. Because only a small fraction can be encoded in long‐term memory, the brain must rely on powerful mechanisms to filter out irrelevant information. To understand the neuronal mechanisms underlying the gating of information into long‐term memory, we employed a paradigm where the encoding was directed by a “Remember” or a “No‐Remember” cue. We found that posterior alpha activity increased prior to the “No‐Remember” stimuli, whereas it decreased prior to the “Remember” stimuli. The sources were localized in the parietal cortex included in the dorsal attention network. Subjects with a larger cue‐modulation of the alpha activity had better memory for the to‐be‐remembered items. Interestingly, alpha activity reflecting successful inhibition following the “No‐Remember” cue was observed in the frontal midline structures suggesting preparatory inhibition was mediated by anterior parts of the dorsal attention network. During the presentation of the memory items, there was more gamma activity for the “Remember” compared to the “No‐Remember” items in the same regions. Importantly, the anticipatory alpha power during cue predicted the gamma power during item. Our findings suggest that top‐down controlled alpha activity reflects attentional inhibition of sensory processing in the dorsal attention network, which then finally gates information to long‐term memory. This gating is achieved by inhibiting the processing of visual information reflected by neuronal synchronization in the gamma band. In conclusion, the functional architecture revealed by region‐specific changes in the alpha activity reflects attentional modulation which has consequences for long‐term memory encoding. Hum Brain Mapp 35:3972–3987, 2013. © 2013 Wiley Periodicals, Inc.     

Reaction time and movement velocity abnormalities in Parkinson's disease under different task conditions

We examined reaction times, movement velocities, and the associated agonist and antagonist muscle behaviors in nine Parkinson's disease (PD) patients and eight normal subjects before and after medications, using a wrist extension task to changing locations of a visual target. Targets changing 500 msec before an auditory "go" signal act as a preparatory cue, while targets changing at the time of the go signal provide a combined auditory and visual stimulus. Late target changes allowed examination of (1) reaction times during an ongoing movement, and (2) movement in the presence and absence of visual targets. PD prolonged the time from the onset agonist electromyographic activity and reduction of antagonist activity to movement onset. Both were shortened by preparatory cues and combined visual and auditory go signals. PD slowed movement only in a subset of trials in which there was movement to a displayed target.