Imperative Signal Research Articles

Unilateral caudate inactivation increases motor impulsivity in rhesus monkeys.

Impulsivity, the tendency to react quickly and without consideration of consequences, is correlated with asymmetry in the volume of the caudate nucleus in human patients. In this study, we sought to determine whether the induction of functional asymmetry in the caudate nucleus of monkeys would produce phenomenologically comparable behavior. We found that unilateral suppression of the ventral caudate nucleus increases impulsive behavior in rhesus monkeys. Impulsivity was modeled by the subjects' inability to maintain hold of a touch-sensitive bar until presentation of an imperative signal. Two methods were used to suppress activity in the caudate region. First, muscimol was locally infused. Second, a viral construct expressing the hM4Di DREADD (designer receptor exclusively activated by designer drug) was injected at the same site. Clozapine N-oxide and deschloroclozapine activate the DREADD to suppress neuronal activity. Both methods of suppression, pharmacological and chemogenetic, increased the rate of early bar releases, a behavior we interpret to indicate impulsivity. Thus, we demonstrate a causal relationship between caudate asymmetry and impulsivity.

Two forms of short-interval intracortical inhibition in human motor cortex

BackgroundPulses of transcranial magnetic stimulation (TMS) with a predominantly anterior-posterior (AP) or posterior-anterior (PA) current direction over the primary motor cortex appear to activate distinct excitatory inputs to corticospinal neurons. In contrast, very few reports have examined whether the inhibitory neurons responsible for short-interval intracortical inhibition (SICI) are sensitive to TMS current direction. ObjectivesTo investigate whether SICI evaluated with AP and PA conditioning stimuli (CSPA and CSAP) activate different inhibitory pathways. SICI was always assessed using a PA-oriented test stimulus (TSPA). MethodsUsing two superimposed TMS coils, CSPA and CSAP were applied at interstimulus intervals (ISI) of 1–5 ms before a TSPA, and at a range of different intensities. Using a triple stimulation design, we then tested whether SICI at ISI of 3 ms using opposite directions of CS (SICICSPA3 and SICICSAP3) interacted differently with three other forms of inhibition, including SICI at ISI of 2 ms (SICICSPA2), cerebellum-motor cortex inhibition (CBI 5 ms) and short-latency afferent inhibition (SAI 22 ms). Finally, we compared the effect of tonic and phasic voluntary contraction on SICICSPA3 and SICICSAP3. ResultsCSAP produced little SICI at ISIs = 1 and 2 ms. However, at ISI = 3 ms, both CSAP and CSPA were equally effective at the same percent of maximum stimulator output. Despite this apparent similarity, combining SICICSPA3 or SICICSAP3 with other forms of inhibition led to quite different results: SICICSPA3 interacted in complex ways with CBI, SAI and SICICSPA2, whereas the effect of SICICSAP3 appeared to be quite independent of them. Although SICICSPA and SICICSAP were both reduced by the same amount during voluntary tonic contraction compared with rest, in a simple reaction time task SICICSAP was disinhibited much earlier following the imperative signal than SICICSPA. ConclusionsSICICSPA appears to activate a different inhibitory pathway to that activated by SICICSAP. The difference is behaviourally relevant since the pathways are controlled differently during volitional contraction. The results may explain some previous pathological data and open the possibility of testing whether these pathways are differentially recruited in a range of tasks.

Coordinating Initial and Final Action Goals in Planning Grasp-to-Rotate Movements: An ERP Study

Action goals have often been investigated in previous studies within a single action. However, most of the manual actions (such as prehension) are not restricted to a single action towards the object but can involve multiple follow-up actions to achieve a further purpose. The coordination of the initial (grip posture) and final (task purpose) action goals within such complex actions is still not fully understood. In the present experiment, the neural mechanisms underlying the goal coordination were investigated with the help of event-related potentials (ERP). With the “first cue – second cue – imperative signal” design, the action goals were presented separately in different sequences (either “final-initial” or “initial-final”), and participants were instructed to plan and execute a grasp-to-rotate movement with either free-choice or specified grasping. Results revealed that shorter reaction times were needed for the final-initial than for the initial-final trials only when the movement requires a free-choice grasping. At the moment when the goal information was incomplete (at the first cue), final goals evoked a larger anterior P2 than initial goals, whereas initial goals elicited a larger anterior N2 and a more robust frontal negativity (400–-550 ms) than final goals. When the goal information was complete (at the second cue), we only found a larger P2 for final goals than for initial goals in free-choice grasping. Moreover, a larger N2 was also found for the specified than for the free-choice grasping in initial-final trials. These neurophysiological results indicate that final goals are more critical than initial grip postures in planning prehensile movements. The initial and final action goals seem to be preferably coordinated in a hierarchical manner, that is, the final task purpose is processed with precedence, whereas the initial grip posture is selected depending on the final task purpose.

From nursing Virgins to brelfies: the project of maternal femininity

ABSTRACT Contemporary debates about breastfeeding are saturated with ideas about what constitutes ‘good’ and ‘bad’ mothering; ‘achieved’ or ‘failed’ femininity frequently derived from Christian representations of the Virgin Mother. In this article we trace deeply entrenched values about the maternal breast through the representation of the Madonna del Latte, specifically concentrating on breast ownership and the depiction of unattainable motherhood. The myth of maternal femininity is then explored in relation to contemporary visual discourses of the nursing mother in which the narrative of unattainable or failed maternal femininity is challenged. The performance work of Megan Marsh-McGlone and the phenomenon of sharing ‘brelfies’ on social media are read as indicative of a new relationship to the Mary myth in which mothers expose the cultural fear of achieved maternal femininity. Ultimately, we argue that what is required of mothers is that they are held in a process of becoming maternal and that representations that subvert this imperative signal a shift in the landscape of Marian iconography.

Age-related changes in TMS-based gamma-aminobutyric acid (GABA) modulation during a unimanual simple and choice reaction time task.

Age-related changes in TMS-based gamma-aminobutyric acid (GABA) modulation during a unimanual simple and choice reaction time task.

Investigating the effect of anticipating a startling acoustic stimulus on preparatory inhibition

Motor-evoked potentials (MEPs) to transcranial magnetic stimulation (TMS) show a profound suppression when elicited during the instructed-delay of reaction time (RT) tasks. One predominant hypothesis is that this phenomenon, called "preparatory inhibition", reflects the operation of processes that suppress motor activity to withhold prepared (but delayed) responses, a form of impulse control. In addition, a startling acoustic stimulus (SAS) -a loud and narrow sound- can trigger the release of prepared responses in RT tasks. We predicted that, if such premature release is clearly forbidden, then anticipating a SAS during delay periods may be associated with increased preparatory inhibition for greater impulse control. Subjects performed a behavioural (n=16) and TMS (n=11) experiment. Both used a choice RT task that required subjects to choose a response based on a preparatory cue but to only release it after an imperative signal. SAS and TMS pulses were elicited at the end of the delay period and subjects were asked to do their best to only release their response after the imperative signal, even in the presence of SAS. SAS could be either rare or frequent, in separate blocks. Consistent with the literature, SAS shortened RTs, especially when they occurred frequently. Moreover, MEPs were suppressed when subjects delayed prepared responses but this preparatory inhibition did not depend on whether SAS were frequent or rare. The stronger RT shortening with frequent rather than rare SAS may be due to increased attention and/or reduced reactive inhibition to SAS, leaving preparatory inhibition unaffected.

StartReact effects in first dorsal interosseous muscle are absent in a pinch task, but present when combined with elbow flexion.

ObjectiveTo provide a neurophysiological tool for assessing sensorimotor pathways, which may differ for those involving distal muscles in simple tasks from those involving distal muscles in a kinetic chain task, or proximal muscles in both.MethodsWe compared latencies and magnitudes of motor responses in a reaction time paradigm in a proximal (biceps brachii, BB) and a distal (first dorsal interosseous, FDI) muscle following electrical stimuli used as imperative signal (IS) delivered to the index finger. These stimuli were applied during different motor tasks: simple tasks involving either one muscle, e.g. flexing the elbow for BB (FLEX), or pinching a pen for FDI (PINCH); combined tasks engaging both muscles by pinching and flexing simultaneously (PINCH-FLEX). Stimuli were of varying intensity and occasionally elicited a startle response, and a StartReact effect.ResultsIn BB, response latencies decreased gradually and response amplitudes increased progressively with increasing IS intensities for non-startling trials, while for trials containing startle responses, latencies were uniformly shortened and response amplitudes similarly augmented across all IS intensities in both FLEX and PINCH-FLEX. In FDI, response latencies decreased gradually and response amplitudes increased progressively with increasing IS intensities in both PINCH and PINCH-FLEX for non-startling trials, but, unlike in BB for the simple task, in PINCH for trials containing startle responses as well. In PINCH-FLEX, FDI latencies were uniformly shortened and amplitudes similarly increased across all stimulus intensities whenever startle signs were present.ConclusionsOur results suggest the presence of different sensorimotor pathways supporting a dissociation between simple tasks that involve distal upper limb muscles (FDI in PINCH) from simple tasks involving proximal muscles (BB in FLEX), and combined tasks that engage both muscles (FDI and BB in PINCH-FLEX), all in accordance with differential importance in the control of movements by cortical and subcortical structures.SignificanceSimple assessment tools may provide useful information regarding the differential involvement of sensorimotor pathways in the control of both simple and combined tasks that engage proximal and distal muscles.

Using a Double-Coil TMS Protocol to Assess Preparatory Inhibition Bilaterally.

Transcranial magnetic stimulation (TMS) applied over the primary motor cortex (M1), elicits motor-evoked potentials (MEPs) in contralateral limb muscles which are valuable indicators of corticospinal excitability (CSE) at the time of stimulation. So far, most studies have used single-coil TMS over one M1, yielding MEPs in muscles of a single limb—usually the hand. However, tracking CSE in the two hands simultaneously would be useful in many contexts. We recently showed that, in the resting state, double-coil stimulation of the two M1 with a 1 ms inter-pulse interval (double-coil1 ms TMS) elicits MEPs in both hands that are comparable to MEPs obtained using single-coil TMS. To further evaluate this new technique, we considered the MEPs elicited by double-coil1 ms TMS in an instructed-delay choice reaction time task where a prepared response has to be withheld until an imperative signal is displayed. Single-coil TMS studies have repetitively shown that in this type of task, the motor system is transiently inhibited during the delay period, as evident from the broad suppression of MEP amplitudes. Here, we aimed at investigating whether a comparable inhibitory effect can be observed with MEPs elicited using double-coil1 ms TMS. To do so, we compared the amplitude as well as the coefficient of variation (CV) of MEPs produced by double-coil1 ms or single-coil TMS during action preparation. We observed that MEPs were suppressed (smaller amplitude) and often less variable (smaller CV) during the delay period compared to baseline. Importantly, these effects were equivalent whether single-coil or double-coil1 ms TMS was used. This suggests that double-coil1 ms TMS is a reliable tool to assess CSE, not only when subjects are at rest, but also when they are involved in a task, opening new research horizons for scientists interested in the corticospinal correlates of human behavior.

The Neural Specificity of Movement Preparation During Actual and Imagined Movements.

Current theories consider motor imagery, the mental representation of action, to have considerable functional overlap with the processes involved in actual movement preparation and execution. To test the neural specificity of motor imagery, we conducted a series of 3 experiments using transcranial magnetic stimulation (TMS). We compared changes in corticospinal excitability as people prepared and implemented actual or imagined movements, using a delayed response task in which a cue indicated the forthcoming response. TMS pulses, used to elicit motor-evoked responses in the first dorsal interosseous muscle of the right hand, were applied before and after an imperative signal, allowing us to probe the state of excitability during movement preparation and implementation. Similar to previous work, excitability increased in the agonist muscle during the implementation of an actual or imagined movement. Interestingly, preparing an imagined movement engaged similar inhibitory processes as that observed during actual movement, although the degree of inhibition was less selective in the imagery conditions. These changes in corticospinal excitability were specific to actual/imagined movement preparation, as no modulation was observed when preparing and generating images of cued visual objects. Taken together, inhibition is a signature of how actions are prepared, whether they are imagined or actually executed.

The role of Dopamine in Preparatory Inhibition: What can we learn from Parkinson’s disease?

The role of Dopamine in Preparatory Inhibition: What can we learn from Parkinson’s disease?

Learning stage-dependent effect of M1 disruption on value-based motor decisions

The present study aimed at characterizing the impact of M1 disruption on the implementation of implicit value information in motor decisions, at both early stages (during reinforcement learning) and late stages (after consolidation) of action value encoding. Fifty subjects performed, over three consecutive days, a task that required them to select between two finger responses according to the color (instruction) and to the shape (implicit, undisclosed rule) of an imperative signal: considering the implicit rule in addition to the instruction allowed subjects to earn more money. We investigated the functional contribution of M1 to the implementation of the implicit rule in subjects' motor decisions. Continuous theta burst stimulation (cTBS) was applied over M1 either on Day 1 or on Day 3, producing a temporary lesion either during reinforcement learning (cTBSLearning group) or after consolidation of the implicit rule, during decision-making (cTBSDecision group), respectively. Interestingly, disrupting M1 activity on Day 1 improved the reliance on the implicit rule, plausibly because M1 cTBS increased dopamine release in the putamen in an indirect way. This finding corroborates the view that cTBS may affect activity in unstimulated areas, such as the basal ganglia. Notably, this effect was short-lasting; it did not persist overnight, suggesting that the functional integrity of M1 during learning is a prerequisite for the consolidation of implicit value information to occur. Besides, cTBS over M1 did not impact the use of the implicit rule when applied on Day 3, although it did so when applied on Day 2 in a recent study where the reliance on the implicit rule declined following cTBS (Derosiere et al., 2017). Overall, these findings indicate that the human M1 is functionally involved in the consolidation and implementation of implicit value information underlying motor decisions. However, M1 contribution seems to vanish as subjects become more experienced in using the implicit value information to make their motor decisions.

S138 Sensory tricks, startle and prepulse inhibition in blepharospasm

Objectives Dystonia is a disorder that manifests as an involuntary muscle contraction that may be generated by abnormal processing of sensory inputs. Abnormalities of motor control occur mainly in circuits of some complexity. An important feature in blepharospasm (BSP) is the need for a fine control of the level of eye closure, which may be abnormal in BSP patients. Methods We have examined the response of the orbicularis oculi (OOc) to various external interventions. For the study of the blink reflex, electrical stimuli were applied to the supraorbital nerve of each side or mechanical stimuli were applied to glabella and chin. For the study of prepulse inhibition, a conditioning electrical prepulse was applied to the median nerve or auditory stimuli were delivered 100 ms prior to the supraorbital nerve shock. For a startle, a loud auditory stimulus was delivered unexpectedly or together with an imperative somatosensory signal carrying the instruction to react either closing or opening the eyes. Results The blink reflex was abnormally larger and was inhibited less by a prepulse in patients than in healthy subjects. The degree of prepulse inhibition correlated with the degree of transient alleviation of BSP with a sensory trick. A startle sound together with the instruction to react accelerated reaction and movement times in closing and opening the eyes in both healthy subjects and patients. Discussion The combination of various electrophysiological tests characterize the clinical phenomenology in patients with BSP. Conclusion Prepulse inhibition in BSP patients correlate with the benefit of sensory tricks.

Face/Agent Interference in Individual and Social Context

Two experiments studied the impact of irrelevant background faces on Go responses to imperative signals superimposed on them. Two major results were obtained. The first concerns a response time advantage of one's own face and a related advantage of familiar over unfamiliar faces. When the imperative signal was superimposed on the responding agent's own face, or the face of a sibling or friend, Go responses were faster as compared to unfamiliar faces. The second concerns the impact of social context. A slight increase of the own-face advantage was observed when the task was performed together with a co-actor. Importantly, this increase was only observed for unfamiliar, not familiar co-actors. The implications of these results are discussed in a framework for stimulus-based response selection which relies on two factors, face familiarity and face/agent compatibility.

P159 Specific and non-specific inhibition of interneurons circuits during motor preparation and suppression

Introduction Corticospinal excitability is transiently suppressed during motor preparation (Greenhouse et al., 2015 ) and when cancelling a prepared movement (Hoshiyama et al., 1997 ). Because MEPs are suppressed even in task-irrelevant muscles Greenhouse et al., 2015 , Hoshiyama et al., 1997 ), it has been proposed that there is a “global” inhibitory signal (Greenhouse et al., 2015) . Objectives We tested the whether the inhibition is global or specific to different excitatory interneuron circuits in primary motor cortex (M1). Materials & methods A controllable pulse parameter TMS (cTMS; Rogue Resolutions Ltd.) device was used to activate distinct interneuron circuits in M1 by changing the coil orientation (posterior–anterior, PA; anterior–posterior, AP) and TMS pulse width (30–120 μs). Motor evoked potentials (MEPs) were elicited in the first dorsal interosseous muscle(s) using short AP- (AP30) and long PA-directed (PA120) pulses at various stages during: (1) a left/right choice reaction time task (CRTT) in which a non-informative warning cue appeared 0.5 s prior to the imperative signal indicated the required response; and (2) a Go/No-Go task in which a non-informative warning cue appeared 2 s prior to a Go or No-Go signal indicating to respond or not respond. Because TMS pulse specificity is greatest at low stimulus intensities, experiments were conducted with weak voluntary contraction (5–10% maximum) to reduce stimulus threshold. Results AP30 pulses evoked MEPs with a longer latency than PA120 pulses, confirming that different excitatory interneurons were recruited by the different pulse. [RH1] In the CRTT, the amplitude of AP30-evoked MEPs was suppressed at the time of the imperative stimulus in both responding and non-responding hands, whereas responses to PA120 stimuli were unaffected (Fig. 1A) Download : Download high-res image (366KB) Download : Download full-size image .[RH2] In the Go/No-Go task, AP30 and PA120 MEPs were suppressed to a similar extent in the No-Go condition near to the expected onset of movement (Fig. 1B). Conclusion During motor preparation for the CRTT there was a selective suppression of one interneuron circuit in M1. In contrast, a global inhibition of corticospinal excitability was observed following a decision not to move. The implication is that the mechanisms underlying each form of inhibition are at least partially distinct.

Effect of Aging on Motor Inhibition during Action Preparation under Sensory Conflict.

Motor behaviors often require refraining from selecting options that may be part of the repertoire of natural response tendencies but that are in conflict with ongoing goals. The presence of sensory conflict has a behavioral cost but the latter can be attenuated in contexts where control processes are recruited because conflict is expected in advance, producing a behavioral gain compared to contexts where conflict occurs in a less predictable way. In the present study, we investigated the corticospinal correlates of these behavioral effects (both conflict-driven cost and context-related gain). To do so, we measured motor-evoked potentials (MEPs) elicited by transcranial magnetic stimulation (TMS) over the primary motor cortex (M1) of young and healthy older adults performing the Eriksen Flanker Task. Subjects performed button-presses according to a central arrow, flanked by irrelevant arrows pointing in the same (congruent trial) or opposite direction (incongruent trial). Conflict expectation was manipulated by changing the probability of congruent and incongruent trials in a given block. It was either high (mostly incongruent blocks, MIB, 80% incongruent trials) or low (mostly congruent blocks, MCB, 80% congruent). The MEP data indicate that the conflict-driven behavioral cost is associated with a strong increase in inappropriate motor activity regardless of the age of individuals, as revealed by larger MEPs in the non-responding muscle in incongruent than in congruent trials. However, this aberrant facilitation disappeared in both groups of subjects when conflict could be anticipated (i.e., in the MIBs) compared to when it occurred in a less predictably way (MCBs), probably allowing the behavioral gain observed in both the young and the older individuals. Hence, the ability to overcome and anticipate conflict was surprisingly preserved in the older adults. Nevertheless, some control processes are likely to evolve with age because the behavioral gain observed in the MIB context was associated with an attenuated suppression of MEPs at the time of the imperative signal (i.e., before conflict is actually detected) in older individuals, suggesting altered motor inhibition, compared to young individuals. In addition, the behavioral analysis suggests that young and older adults rely on different strategies to cope with conflict, including a change in speed-accuracy tradeoff.

Investigation of timing preparation during response initiation and execution using a startling acoustic stimulus.

The purpose of the current study was to examine the processes involved in the preparation of timing during response initiation and execution through the use of a startling acoustic stimulus (SAS). In Experiment 1, participants performed a delayed response task in which a two key-press movement was to be initiated 200ms after an imperative signal (IS) with either a short (200ms) or long (500ms) interval between key-presses. On selected trials, a SAS was presented to probe the preparation processes associated with the initiation delay and execution of the inter-key interval. The SAS resulted in a significant decrease in the initiation time, which was attributed to a speeding of pacemaker pulses used to time the delay interval, caused by an increased activation due to the SAS. Conversely, the SAS delayed the short inter-key interval, which was attributed to temporary interference with cortical processing. In Experiment 2, participants performed a 500-ms delayed response task involving two key-presses 200ms apart. In this condition, the SAS resulted in significantly decreased initiation time and a delayed inter-key interval (p=.053). Collectively, these results support a different timeline for the preparation of the delay interval, which is thought to be prepared in advance of the IS, and the inter-key interval, which is thought to be prepared following the IS. This conclusion provides novel information with regard to timing preparation that is consistent with models in which response preparation, initiation, and execution are considered separate and dissociable processes.

Modulation of force and velocity for accurate tasks by a startle

Background Some skilfulfast activities require control of trajectory and force to fulfill the task. It is known that voluntary reactions can be speeded up by a startling auditory stimulus (SAS) delivered at the same time as the imperative signal (IS), a phenomenon termed Start-React (Valls-Sole et al., 1999). The aim of the present study was to examine how the performance of a task that demands accuracy, mainly in terms of velocity during the trajectory or force control when holding an object, is changed by a SAS timed to occur either at the time of IS or at a pre-determined time interval when movement has already begun and motor control of accurate reaching is expected. Methods Nine subjects performed an accurate elbow extension between two spots placed at an angular distance of 30° when holding with their right hand a pen that monitored, through two strain gauge systems, the force of the subject’s fingers during the hold, and the force at which pen-tip was exposed with table contact. According to block of trials, this end spot had three possible diameters: 5, 10, and 20 mm. Kinematic and force parameters were measured for three conditions, control, with SAS at IS or SAS delivered when movement has already started. Results In SAS trials, there were no evident differences with respect to control trials in the general outline but there were differences in the timing of the events. In SAS-IS trials time to peak velocity was 166 ms (SD = 41 ms), time to peak force was 95 ms (SD = 43 ms) both shorter than in control or SAS delivered when movement began. Peak velocity (315 ± 56 o/s) and peak force (8 ± 2 arbitr. units) were larger for SAS-IS trials than for control or SAS delivered when movement started. Conclusions Startle can advance a task that requires accuracy with velocity and force modulations, but when task onset is already launched there are not evident changes in velocity or force parameters, that seem already prepared.

Perceptual, cognitive and motor responses to contact heat stimuli

Objective Afferent inputs in humans lead to a trace of excitability in the corresponding sensory channel, perception, cognition and prepared motor responses. We investigated how the events related to each of these aspects of sensory processing are affected by stimulus intensity. Methods Nine healthy subjects performed a wrist extension reaction time (RT) task experiment in front of the Libet’s clock in order to assess the time at which they perceived the contact heat stimulus used as imperative signal (awareness = AW). We recorded also the thermoalgesic evoked potentials (N2/P2). Stimuli were either strong (52 °C) or mild (42 °C). We monitored RT, N2 latency, amplitude of N2–P2, and the time of AW for each stimulation condition. Results Usual order of responses was RT, AW, N2. N2–P2 amplitude had a negative correlation with AW ( p Conclusions Reaction time is executed as soon as the imperative signal is processed, while a late cognitive response does not depend on stimulus intensity. Key message The test used allows for separation of perceptual levels in humans.

Comparison of the two cerebral hemispheres in inhibitory processes operative during movement preparation

Neuroimaging and neuropsychological studies suggest that in right-handed individuals, the left hemisphere plays a dominant role in praxis, relative to the right hemisphere. However hemispheric asymmetries assessed with transcranial magnetic stimulation (TMS) has not shown consistent differences in corticospinal (CS) excitability of the two hemispheres during movements. In the current study, we systematically explored hemispheric asymmetries in inhibitory processes that are manifest during movement preparation and initiation. Single-pulse TMS was applied over the left or right primary motor cortex (M1LEFT and M1RIGHT, respectively) to elicit motor-evoked potentials (MEPs) in the contralateral hand while participants performed a two-choice reaction time task requiring a cued movement of the left or right index finger. In Experiments 1 and 2, TMS probes were obtained during a delay period following the presentation of the preparatory cue that provided partial or full information about the required response. MEPs were suppressed relative to baseline regardless of whether they were elicited in a cued or uncued hand. Importantly, the magnitude of these inhibitory changes in CS excitability was similar when TMS was applied over M1LEFT or M1RIGHT, irrespective of the amount of information carried by the preparatory cue. In Experiment 3, there was no preparatory cue and TMS was applied at various time points after the imperative signal. When CS excitability was probed in the cued effector, MEPs were initially inhibited and then rose across the reaction time interval. This function was similar for M1LEFT and M1RIGHT TMS. When CS excitability was probed in the uncued effector, MEPs remained inhibited throughout the RT interval. However, MEPs in right FDI became more inhibited during selection and initiation of a left hand movement, whereas MEPs in left FDI remained relatively invariant across RT interval for the right hand. In addition to these task-specific effects, there was a global difference in CS excitability across experiments between the two hemispheres. When the intensity of stimulation was set to 115% of the resting threshold, MEPs were larger when the TMS probe was applied over the M1LEFT than over M1RIGHT. In summary, while the latter result suggests that M1LEFT is more excitable than M1RIGHT, the recruitment of preparatory inhibitory mechanisms is similar within the two cerebral hemispheres.

Startle activation is additive with voluntary cortical activation irrespective of stimulus modality

When a startling acoustic stimulus (SAS) is presented during a simple reaction time (RT) task, it can trigger the prepared response through an involuntary initiation pathway. Previous research modelling the effects of presenting a SAS at various intervals following a non-startling auditory imperative signal (IS) suggested that involuntary initiation-related neural activation is additive with the voluntary initiation processes. The current study tested the predictions of this additive model when the SAS and IS are of different modalities by using a visual rather than auditory go-signal. Because voluntary RT latencies are delayed for visual stimuli compared to acoustic stimuli, it was hypothesised that the time course of additive activation would be similarly delayed. Participants performed 150 RT trials requiring a targeted 20° wrist extension task with a SAS presented 0–125ms following a visual go-signal. Results were not different to those predicted by an additive model (p=0.979), yet were significantly different to those predicted by a horse-race model (p=0.037), indicating a joint contribution of voluntary and involuntary activation, even when the IS and SAS are of different modalities. Furthermore, the results indicated that voluntary RT differences due to stimulus modality are attributable to processes that occur prior to the increase in initiation-related activation.