Express Visuomotor Responses Reflect Knowledge of Both Target Locations and Contextual Rules during Reaches of Different Amplitudes.

When humans reach to visual targets, extremely rapid (∼90 ms) target-directed responses can be observed in task-relevant proximal muscles. Such express visuomotor responses are inflexibly locked in time and space to the target and have been proposed to reflect rapid visuomotor transformations conveyed subcortically via the tecto-reticulo-spinal pathway. Previously, we showed that express visuomotor responses are sensitive to explicit cue-driven information about the target, suggesting that the express pathway can be modulated by cortical signals affording contextual prestimulus expectations. Here, we show that the express visuomotor system incorporates information about the physical hand-to-target distance and contextual rules during visuospatial tasks requiring different movement amplitudes. In one experiment, we recorded the activity from two shoulder muscles as 14 participants (6 females) reached toward targets that appeared at different distances from the reaching hand. Increasing the reaching distance facilitated the generation of frequent and large express visuomotor responses. This suggests that both the direction and amplitude of veridical hand-to-target reaches are encoded along the putative subcortical express pathway. In a second experiment, we modulated the movement amplitude by asking 12 participants (4 females) to deliberately undershoot, overshoot, or stop (control) at the target. The overshoot and undershoot tasks impaired the generation of large and frequent express visuomotor responses, consistent with the inability of the express pathway to generate responses directed toward nonveridical targets as in the anti-reach task. Our findings appear to reflect strategic, cortically driven modulation of the express visuomotor circuit to facilitate rapid and effective response initiation during target-directed actions.SIGNIFICANCE STATEMENT Express (∼90 ms) arm muscle responses that are consistently tuned toward the location of visual stimuli suggest a subcortical contribution to target-directed visuomotor behavior in humans, potentially via the tecto-reticulo-spinal pathway. Here, we show that express muscle responses are modulated appropriately to reach targets at different distances, but generally suppressed when the task required nonveridical responses to overshoot/undershoot the real target. This suggests that the tecto-reticulo-spinal pathway can be exploited strategically by the cerebral cortex to facilitate rapid initiation of effective responses during a visuospatial task.

Evidence for an Intricate Relationship Between Express Visuomotor Responses, Postural Control and Rapid Step Initiation in the Lower Limbs.

Recent work has described express visuomotor responses (EVRs) on the upper limb. EVRs are directionally-tuned bursts of muscle activity that occur within 100 ms of visual stimulus appearance, facilitating rapid reaching. Rapid stepping responses are also important in daily life, and while there is evidence of EVR expression on lower limbs, it is unknown whether lower-limb EVRs are influenced by increased postural demands. Here, we investigate the interaction between stepping-related EVRs and anticipatory postural adjustments (APAs) that typically precede step initiation. 16 healthy young subjects rapidly stepped towards visual targets presented in front of the left or right foot. We recorded bilateral surface EMG of gluteus medius (GM), a muscle involved in both APAs and stepping, and bilateral ground reaction forces. Two conditions were introduced: an anterolateral or anteromedial stepping condition with reduced or increased postural demands, respectively. In the anterolateral stepping condition, EVRs were robustly and strongly present in stance-side GM, and ground reaction forces revealed strongly decreased expression of APAs. Larger EVRs preceded shorter RTs, consistent with EVRs facilitating step initiation. In contrast, in the anteromedial stepping condition, EVRs were largely absent, and ground reaction forces revealed the consistent expression of APAs. When occasionally present, EVRs in the anteromedial stepping condition preceded larger APAs and longer RTs. Thus, while EVRs in lower limbs can facilitate rapid stepping, their expression is normally suppressed when postural stability is low. Failing to appropriately suppress EVRs in such situations disrupts postural stability, necessitating larger compensatory APAs and leading to longer stepping RTs.

Done in 65 ms: Express Visuomotor Responses in Upper Limb Muscles in Rhesus Macaques.

How rapidly can the brain transform vision into action? Work in humans has established that the transformation for visually-guided reaching can be remarkably rapid, with the first phase of upper limb muscle recruitment, the express visuomotor response, beginning within less than 100 ms of visual target presentation. Such short-latency responses limit the opportunities for extensive cortical processing, leading to the hypothesis that they are generated via the subcortical tecto-reticulo-spinal pathway. Here, we examine whether nonhuman primates (NHPs) exhibit express visuomotor responses. Two male macaques made visually-guided reaches in a behavioral paradigm known to elicit express visuomotor responses in humans, while we acquired intramuscular recordings from the deltoid muscle. Across several variants of this paradigm, express visuomotor responses began within 65 ms (range: 48-91 ms) of target presentation. Although the timing of the express visuomotor response did not co-vary with reaction time, larger express visuomotor responses tended to precede shorter latency reaches. Further, we observed that the magnitude of the express visuomotor response could be muted by contextual context, although this effect was quite variable. Overall, the response properties in NHPs resemble those in humans. Our results establish a new benchmark for visuomotor transformations underlying visually-guided reaches, setting the stage for experiments that can directly compare the role of cortical and subcortical areas in reaching when time is of the essence.

Single-Trial Dynamics of Competing Reach Plans in the Human Motor Periphery.

Contemporary motor control theories propose competition between multiple motor plans before the winning command is executed. While most competitions are completed before movement onset, movements are often initiated before the competition has been resolved. An example of this is saccadic averaging, wherein the eyes land at an intermediate location between two visual targets. Behavioral and neurophysiological signatures of competing motor commands have also been reported for reaching movements, but debate remains about whether such signatures attest to an unresolved competition, arise from averaging across many trials, or reflect a strategy to optimize behavior given task constraints. Here, we recorded EMG activity from an upper limb muscle (m. pectoralis) while 12 (8 female) participants performed an immediate response reach task, freely choosing between one of two identical and suddenly presented visual targets. On each trial, muscle recruitment showed two distinct phases of directionally tuned activity. In the first wave, time-locked ∼100 ms of target presentation, muscle activity was clearly influenced by the nonchosen target, reflecting a competition between reach commands that was biased in favor of the ultimately chosen target. This resulted in an initial movement intermediate between the two targets. In contrast, the second wave, time-locked to voluntary reach onset, was not biased toward the nonchosen target, showing that the competition between targets was resolved. Instead, this wave of activity compensated for the averaging induced by the first wave. Thus, single-trial analysis reveals an evolution in how the nonchosen target differentially influences the first and second wave of muscle activity.SIGNIFICANCE STATEMENT Contemporary theories of motor control suggest that multiple motor plans compete for selection before the winning command is executed. Evidence for this is found in intermediate reach movements toward two potential target locations, but recent findings have challenged this notion by arguing that intermediate reaching movements reflect an optimal response strategy. By examining upper limb muscle recruitment during a free-choice reach task, we show early recruitment of a suboptimal averaged motor command to the two targets that subsequently transitions to a single motor command that compensates for the initially averaged motor command. Recording limb muscle activity permits single-trial resolution of the dynamic influence of the nonchosen target through time.

Symbolic cues enhance express visuomotor responses in human arm muscles at the motor planning rather than the visuospatial processing stage.

Humans can produce "express" (∼100 ms) arm muscle responses that are inflexibly locked in time and space to visual target presentations, consistent with subcortical visuomotor transformations via the tecto-reticulo-spinal pathway. These express visuomotor responses are sensitive to explicit cue-driven expectations, but it is unclear at what stage of sensory-to-motor transformation such modulation occurs. Here, we recorded electromyographic activity from shoulder muscles as participants reached toward one of four virtual targets whose physical location was partially predictable from a symbolic cue. In an experiment in which targets could be veridically reached, express responses were inclusive of the biomechanical requirements for reaching the cued locations and not systematically modulated by cue validity. In a second experiment, movements were restricted to the horizontal plane so that the participants could perform only rightward or leftward reaches, irrespective of target position on the vertical axis. Express muscle responses were almost identical for targets that were validly cued in the horizontal direction, regardless of cue validity in the vertical dimension. Together, these findings suggest that the cue-induced enhancements of express responses are dominated by effects at the level of motor plans and not solely via facilitation of early visuospatial target processing. Notably, direct corticotectal and corticoreticular projections exist that are well-placed to modulate prestimulus motor preparation state in subcortical circuits. Our results could reflect a neural mechanism by which contextually relevant motor responses to compatible visual inputs are rapidly released via subcortical circuits that are sufficiently along the sensory-to-motor continuum.NEW & NOTEWORTHY Express arm muscle responses to suddenly appearing visual targets for reaching rapid have been attributed to the tecto-reticulo-spinal pathway in humans. We demonstrate that symbolic cues before target presentation can modulate such express arm muscle responses compatibly with the biomechanics of the cued reaching direction and the cue validity. This implies cortically mediated modulation of one or more sensorimotor transformation nodes of the subcortical express pathway.

Response characteristics of vestibular evoked myogenic potentials recorded over splenius capitis in young adults and adolescents.

BACKGROUND AND OBJECTIVES: Examine vestibular evoked myogenic potential (VEMP) responses recorded from surface electrodes over Splenius Capitis (SPC) in a seated position. SPECIFIC AIMS: (1) validate response characteristics of VEMP recordings from surface electrodes over Sternocleidomastoid (SCM) and over SCP and (2) assess age effects on responses in adolescents and young adults. MATERIALS AND METHODS: Simultaneous surface VEMP was recorded bilaterally from electrodes placed over the dorsal neck musculature at a location known from previous work to record from SPC in 15 healthy participants during trials with head rotation toward and away from the stimulated ear. VEMP was also recorded from electrodes over SCM, ipsilateral to the stimulus ear, in the same participants in a supine, head lift/turn position. RESULTS: Response amplitudes significantly increased with contraction strength and decreased with age. Participants were able to maintain sufficient contraction strength (amplitude) with head rotation to reliably measure over SPC. Normalized response amplitudes measured from electrodes over contralateral SPC were largest with head rotation contralateral to the stimulus ear. Normalized amplitudes and peak latencies were comparable to the same measures from SCM obtained in supine, head lift/turn position. CONCLUSIONS: Otolith generated myogenic responses can be recorded seated from electrodes over the dorsal neck with head rotation contralateral to the stimulus ear. In this position, contralateral recordings are consistent with responses known from previous work to arise from SPC; ipsilateral recordings may include crosstalk from activated muscles nearby, including ipsilateral SCM. Overall, techniques targeting contralateral SPC during contralateral head turn may provide additional methods of recording VEMPs.

Response characteristics of vestibular evoked myogenic potentials recorded over splenius capitis in young adults and adolescents

Background and objectives: Examine vestibular evoked myogenic potential (VEMP) responses recorded from surface electrodes over Splenius Capitis (SPC) in a seated position. Specific aims: (1) validate response characteristics of VEMP recordings from surface electrodes over Sternocleidomastoid (SCM) and over SCP and (2) assess age effects on responses in adolescents and young adults. Materials and methods: Simultaneous surface VEMP was recorded bilaterally from electrodes placed over the dorsal neck musculature at a location known from previous work to record from SPC in 15 healthy participants during trials with head rotation toward and away from the stimulated ear. VEMP was also recorded from electrodes over SCM, ipsilateral to the stimulus ear, in the same participants in a supine, head lift/turn position. Results: Response amplitudes significantly increased with contraction strength and decreased with age. Participants were able to maintain sufficient contraction strength (amplitude) with head rotation to reliably measure over SPC. Normalized response amplitudes measured from electrodes over contralateral SPC were largest with head rotation contralateral to the stimulus ear. Normalized amplitudes and peak latencies were comparable to the same measures from SCM obtained in supine, head lift/turn position. Conclusions: Otolith generated myogenic responses can be recorded seated from electrodes over the dorsal neck with head rotation contralateral to the stimulus ear. In this position, contralateral recordings are consistent with responses known from previous work to arise from SPC; ipsilateral recordings may include crosstalk from activated muscles nearby, including ipsilateral SCM. Overall, techniques targeting contralateral SPC during contralateral head turn may provide additional methods of recording VEMPs.(AU)

Antecedentes y objetivos: Examinamos las respuestas de los potenciales evocados miogénicos vestibulares (PEMV) recogidas de los electrodos de superficie sobre el músculo esplenio (ME) en posición sentada. Objetivos específicos: 1) validar las características de los registros de la respuesta de los PEMV recogidos de los electrodos de superficie sobre el músculo esternocleidomastoideo (SCM) y el ME, y 2) evaluar los efectos de la edad en adolescentes y adultos jóvenes. Materiales y métodos: Se registraron simultáneamente los PEMV bilaterales de los electrodos situados en la musculatura dorsal del cuello, en un sitio conocido de un estudio anterior para obtener registros del ME en 15 participantes sanos durante los ensayos, con rotación de cabeza hacia y fuera del oído estimulado. También se registraron los PEMV de los electrodos situados sobre el SCM, en posición ipsilateral al oído estimulado, en los mismos participantes, en posición supina y con elevación/giro de cabeza. Resultados: Las amplitudes de la respuesta se incrementaron significativamente con la fuerza de la contracción y disminuyeron con la edad. Los participantes fueron capaces de mantener suficiente fuerza de contracción (amplitud) con la rotación de cabeza, para realizar mediciones fiables sobre el ME. Las amplitudes de la respuesta normalizada medidas en los electrodos sobre el ME contralateral fueron mayores con la rotación de cabeza contralateral al oído estimulado. Las amplitudes normalizadas y las latencias máximas fueron comparables a las mismas medidas del SCM obtenidas en posición supina, y elevación/giro de cabeza. Conclusiones: Las respuestas miogénicas generadas por otolitos pueden registrarse en posición sentada a partir de los electrodos situados en la parte dorsal del cuello, contralateral al oído estimulado. En esta posición, los registros contralaterales son coherentes con las respuestas conocidas de un estudio previo, derivadas del ME; los registros ipsilaterales pueden incluir interferencias de los músculos activados cercanos, incluyendo el SCM ipsilateral. En general, las técnicas centradas en el ME contralateral durante el giro de cabeza contralateral pueden aportar métodos adicionales de registro de los PEMV.(AU)

Express arm responses appear bilaterally on upper-limb muscles in an arm choice reaching task.

When required, humans can generate very short latency reaches toward visual targets, such as catching a falling cellphone. During such rapid reaches, express arm responses are the first wave of upper limb muscle recruitment, occurring ∼80-100 ms after target appearance. There is accumulating evidence that express arm responses arise from signaling along the tecto-reticulo-spinal tract, but the involvement of the reticulospinal tract has not been well studied. Since the reticulospinal tract projects bilaterally, we studied whether express arm responses would be generated bilaterally. Human participants (n = 14; 7 females) performed visually guided reaches in a modified emerging target paradigm where either arm could intercept the target. We recorded electromyographic activity bilaterally from the pectoralis major muscle. Our analysis focused on target locations where participants reached with the right arm on some trials, and the left arm on others. In support of the involvement of the reticulospinal tract, express arm responses persisted bilaterally regardless of which arm reached to the target. The latency and magnitude of the express arm response did not depend on whether the arm was chosen to reach or not. However, on the reaching arm, the magnitude of the express arm response was correlated to the level of anticipatory activity. The bilateral generation of express arm responses supports the involvement of the reticulospinal tract. We surmise that the correlation between anticipatory activity and the magnitude of express arm responses on the reaching arm arises from convergence of cortically derived signals with a parallel subcortical pathway mediating the express arm response.NEW & NOTEWORTHY Express arm responses have been proposed to arise from the tecto-reticulo-spinal tract originating within the superior colliculus, but the involvement of the reticulospinal tract has not been well studied. Here, we show these responses appear bilaterally in a task where either arm can reach to a newly appearing stimulus. Our results suggest that the most rapid visuomotor transformations for reaching are performed by a subcortical pathway.

Completing the puzzle: Why studies in non-human primates are needed to better understand the effects of non-invasive brain stimulation.

Brain stimulation is a core method in neuroscience. Numerous non-invasive brain stimulation (NIBS) techniques are currently in use in basic and clinical research, and recent advances promise the ability to non-invasively access deep brain structures. While encouraging, there is a surprising gap in our understanding of precisely how NIBS perturbs neural activity throughout an interconnected network, and how such perturbed neural activity ultimately links to behaviour. In this review, we will consider why non-human primate (NHP) models of NIBS are ideally situated to address this gap in knowledge, and why the oculomotor network that moves our line of sight offers a particularly valuable platform in which to empirically test hypothesis regarding NIBS-induced changes in brain and behaviour. NHP models of NIBS will enable investigation of the complex, dynamic effects of brain stimulation across multiple hierarchically interconnected brain areas, networks, and effectors. By establishing such links between brain and behavioural output, work in NHPs can help optimize experimental and therapeutic approaches, improve NIBS efficacy, and reduce side-effects of NIBS.

Trial-by-trial modulation of express visuomotor responses induced by symbolic or barely detectable cues.

Human cerebral cortex can produce visuomotor responses that are modulated by contextual and task-specific constraints. However, the distributed cortical network for visuomotor transformations limits the minimal response time of that pathway. Notably, humans can generate express visuomotor responses in arm muscles that are inflexibly tuned to the target location and occur 80-120 ms from stimulus presentation [stimulus-locked responses (SLRs)]. This suggests a subcortical pathway for visuomotor transformations that might involve the superior colliculus and its downstream reticulo-spinal projections. Here we investigated whether cognitive expectations can modulate the SLR. In one experiment, we recorded surface electromyogram (EMG) from shoulder muscles as participants reached toward a visual target whose location was unpredictable in control conditions and partially predictable in cue conditions by interpreting a symbolic cue (75% validity). Valid symbolic cues led to earlier and larger SLRs than control conditions; invalid symbolic cues produced later and smaller SLRs than control conditions. This is consistent with a cortical top-down modulation of the putative subcortical SLR network. In a second experiment, we presented high-contrast targets in isolation (control) or ∼24 ms after low-contrast stimuli, which could appear at the same (valid cue) or opposite (invalid cue) location as the target and with equal probability (50% cue validity). We observed earlier SLRs than control with the valid low-contrast cues, whereas the invalid cues led to the opposite results. These findings may reflect bottom-up attentional mechanisms, potentially evolving subcortically via the superior colliculus. Overall, our results support both top-down and bottom-up modulations of the putative subcortical SLR network in humans.NEW & NOTEWORTHY Express visuomotor responses in humans appear to reflect subcortical sensorimotor transformation of visual inputs, potentially conveyed via the tecto-reticulo-spinal pathway. Here we show that the express responses are influenced by both symbolic and barely detectable spatial cues about stimulus location. The symbolic cue-induced effects suggest cortical top-down modulation of the putative subcortical visuomotor network. The effects of barely detectable cues may reflect exogenous facilitation mechanisms of the tecto-reticulo-spinal pathway.

High-contrast, moving targets in an emerging target paradigm promote fast visuomotor responses during visually guided reaching.

Humans have a remarkable capacity to rapidly interact with the surrounding environment, often by transforming visual input into motor output on a moment-to-moment basis. But what visual features promote rapid reaching? High-contrast, fast-moving targets elicit strong responses in the superior colliculus (SC), a structure associated with express saccades and implicated in rapid electromyographic (EMG) responses on upper limb muscles. To test the influence of stimulus properties on rapid reaches, we had human subjects perform visually guided reaches to moving targets varied by speed (experiment 1) or speed and contrast (experiment 2) in an emerging target paradigm that has recently been shown to robustly elicit fast visuomotor responses. Our analysis focused on stimulus-locked responses (SLRs) on upper limb muscles. SLRs appear within <100 ms of target presentation, and as the first wave of muscle recruitment they have been hypothesized to arise from the SC. Across 32 subjects studied in both experiments, 97% expressed SLRs in the emerging target paradigm, whereas only 69% expressed SLRs in an immediate response paradigm toward static targets. Faster-moving targets (experiment 1) evoked large-magnitude SLRs, whereas high-contrast fast-moving targets (experiment 2) evoked short-latency, large-magnitude SLRs. In some instances, SLR magnitude exceeded the magnitude of movement-aligned activity. Both large-magnitude and short-latency SLRs were correlated with short-latency reach reaction times. Our results support the hypothesis that, in scenarios requiring expedited responses, a subcortical pathway originating in the SC elicits the earliest wave of muscle recruitment, expediting reaction times.NEW & NOTEWORTHY How does the brain rapidly transform vision into action? Here, by recording upper limb muscle activity, we find that high-contrast and fast-moving targets are highly effective at evoking rapid visually guided reaches. We surmise that a brain stem circuit originating in the superior colliculus contributes to the most rapid reaching responses. When time is of the essence, cortical areas may serve to prime this circuit and elaborate subsequent phases of recruitment.

Response characteristics of vestibular evoked myogenic potentials recorded over splenius capitis in young adults and adolescents.

BACKGROUND AND OBJECTIVES: Examine vestibular evoked myogenic potential (VEMP) responses recorded from surface electrodes over Splenius Capitis (SPC) in a seated position. SPECIFIC AIMS: (1) validate response characteristics of VEMP recordings from surface electrodes over Sternocleidomastoid (SCM) and over SCP and (2) assess age effects on responses in adolescents and young adults. MATERIALS AND METHODS: Simultaneous surface VEMP was recorded bilaterally from electrodes placed over the dorsal neck musculature at a location known from previous work to record from SPC in 15 healthy participants during trials with head rotation toward and away from the stimulated ear. VEMP was also recorded from electrodes over SCM, ipsilateral to the stimulus ear, in the same participants in a supine, head lift/turn position. RESULTS: Response amplitudes significantly increased with contraction strength and decreased with age. Participants were able to maintain sufficient contraction strength (amplitude) with head rotation to reliably measure over SPC. Normalized response amplitudes measured from electrodes over contralateral SPC were largest with head rotation contralateral to the stimulus ear. Normalized amplitudes and peak latencies were comparable to the same measures from SCM obtained in supine, head lift/turn position. CONCLUSIONS: Otolith generated myogenic responses can be recorded seated from electrodes over the dorsal neck with head rotation contralateral to the stimulus ear. In this position, contralateral recordings are consistent with responses known from previous work to arise from SPC; ipsilateral recordings may include crosstalk from activated muscles nearby, including ipsilateral SCM. Overall, techniques targeting contralateral SPC during contralateral head turn may provide additional methods of recording VEMPs.

Frontal eye field inactivation alters the readout of superior colliculus activity for saccade generation in a task-dependent manner.

Saccades require a spatiotemporal transformation of activity between the intermediate layers of the superior colliculus (iSC) and downstream brainstem burst generator. The dynamic linear ensemble-coding model (Goossens and Van Opstal 2006) proposes that each iSC spike contributes a fixed mini-vector to saccade displacement. Although biologically-plausible, this model assumes cortical areas like the frontal eye fields (FEF) simply provide the saccadic goal to be executed by the iSC and brainstem burst generator. However, the FEF and iSC operate in unison during saccades, and a pathway from the FEF to the brainstem burst generator that bypasses the iSC exists. Here, we investigate the impact of large yet reversible inactivation of the FEF on iSC activity in the context of the model across four saccade tasks. We exploit the overlap of saccade vectors generated when the FEF is inactivated or not, comparing the number of iSC spikes for metrically-matched saccades. We found that the iSC emits fewer spikes for metrically-matched saccades during FEF inactivation. The decrease in spike count is task-dependent, with a greater decrease accompanying more cognitively-demanding saccades. Our results show that FEF integrity influences the readout of iSC activity in a task-dependent manner. We propose that the dynamic linear ensemble-coding model be modified so that FEF inactivation increases the gain of a readout parameter, effectively increasing the influence of a single iSC spike. We speculate that this modification could be instantiated by FEF and iSC pathways to the cerebellum that could modulate the excitability of the brainstem burst generator.

The influence of temporal predictability on express visuomotor responses.

Humans are able to generate target-directed visuomotor responses in less than 100 ms after stimulus onset. These "express" responses have been termed stimulus-locked responses (SLRs) and are proposed to be modulated by visuomotor transformations performed subcortically via the superior colliculus. Unfortunately, these responses have proven difficult to detect consistently across individuals. The recent report of an effective paradigm for generating SLRs in 100% of participants appears to change this. The task required the interception of a target moving at a constant velocity that emerged from behind a barrier. Here, we aimed to reproduce the efficacy of this paradigm for eliciting SLRs and to test the hypothesis that its effectiveness derives from the predictability of target onset time as opposed to target motion per se. In one experiment, we recorded surface electromyogram (EMG) from shoulder muscles as participants made reaches to intercept temporally predictable or unpredictable targets. Consistent with our hypothesis, predictably timed targets produced more frequent and stronger SLRs than unpredictably timed targets. In a second experiment, we compared different temporally predictable stimuli and observed that transiently presented targets produced larger and earlier SLRs than sustained moving targets. Our results suggest that target motion is not critical for facilitating the SLR expression and that timing predictability does not rely on extrapolation of a physically plausible motion trajectory. These findings provide support for a mechanism whereby an internal timer, probably located in cerebral cortex, primes the processing of both visual input and motor output within the superior colliculus to produce SLRs.NEW & NOTEWORTHY Express stimulus-driven responses in humans have been proposed to be originated subcortically via the superior colliculus. These short-latency responses are facilitated by the presentation of dynamic visual stimuli. Here, we show that this facilitation is related to the predictable target timing, regardless of its kinematic attributes. We propose that the superior colliculus can be primed to generate express stimulus-driven motor responses via cortical top-down projection.

An Emerging Target Paradigm to Evoke Fast Visuomotor Responses on Human Upper Limb Muscles.

To reach towards a seen object, visual information has to be transformed into motor commands. Visual information such as the object's color, shape, and size are processed and integrated within numerous brain areas, then ultimately relayed to the motor periphery. In some instances, a reaction is needed as fast as possible. These fast visuomotor transformations, and their underlying neurological substrates, are poorly understood in humans as they have lacked a reliable biomarker. Stimulus-locked responses (SLRs) are short latency (<100 ms) bursts of electromyographic (EMG) activity representing the first wave of muscle recruitment influenced by visual stimulus presentation. SLRs provide a quantifiable output of rapid visuomotor transformations, but SLRs have not been consistently observed in all subjects in past studies. Here we describe a new, behavioral paradigm featuring the sudden emergence of a moving target below an obstacle that consistently evokes robust SLRs. Human participants generated visually guided reaches toward or away from the emerging target using a robotic manipulandum while surface electrodes recorded EMG activity from the pectoralis major muscle. In comparison to previous studies that investigated SLRs using static stimuli, the SLRs evoked with this emerging target paradigm were larger, evolved earlier, and were present in all participants. Reach reaction times (RTs) were also expedited in the emerging target paradigm. This paradigm affords numerous opportunities for modification that could permit systematic study of the impact of various sensory, cognitive, and motor manipulations on fast visuomotor responses. Overall, our results demonstrate that an emerging target paradigm is capable of consistently and robustly evoking activity within a fast visuomotor system.

Impairment but not abolishment of express saccades after unilateral or bilateral inactivation of the frontal eye fields.

Express saccades are a manifestation of a visual grasp reflex triggered when visual information arrives in the intermediate layers of the superior colliculus (SCi), which in turn orchestrates the lower level brainstem saccade generator to evoke a saccade with a very short latency (~100 ms or less). A prominent theory regarding express saccades generation is that they are facilitated by preparatory signals, presumably from cortical areas, which prime the SCi before the arrival of visual information. Here, we test this theory by reversibly inactivating a key cortical input to the SCi, the frontal eye fields (FEF), while monkeys perform an oculomotor task that promotes express saccades. Across three tasks with a different combination of potential target locations and unilateral or bilateral FEF inactivation, we found a spared ability for monkeys to generate express saccades, despite decreases in express saccade frequency during FEF inactivation. This result is consistent with the FEF having a facilitatory but not critical role in express saccade generation, likely because other cortical areas compensate for the loss of preparatory input to the SCi. However, we also found decreases in the accuracy and peak velocity of express saccades generated during FEF inactivation, which argues for an influence of the FEF on the saccadic burst generator even during express saccades. Overall, our results shed further light on the role of the FEF in the shortest-latency visually-guided eye movements.NEW & NOTEWORTHY Express saccades are the shortest-latency saccade. The frontal eye fields (FEF) are thought to promote express saccades by presetting the superior colliculus. Here, by reversibly inactivating the FEF either unilaterally or bilaterally via cortical cooling, we support this by showing that the FEF plays a facilitative but not critical role in express saccade generation. We also found that FEF inactivation lowered express saccade peak velocity, emphasizing a contribution of the FEF to express saccade kinematics.

Stimulus-Locked Responses on Human Upper Limb Muscles and Corrective Reaches Are Preferentially Evoked by Low Spatial Frequencies.

In situations requiring immediate action, humans can generate visually-guided responses at remarkably short latencies. Here, to better understand the visual attributes that best evoke such rapid responses, we recorded upper limb muscle activity while participants performed visually-guided reaches towards Gabor patches composed of differing spatial frequencies (SFs). We studied reaches initiated from a stable posture (experiment 1, a static condition), or during on-line reach corrections to an abruptly displaced target (experiment 2, a dynamic condition). In both experiments, we detail the latency and prevalence of stimulus-locked responses (SLRs), which are brief bursts of EMG activity that are time-locked to target presentation rather than movement onset. SLRs represent the first wave of EMG recruitment influenced by target presentation, and enable quantification of rapid visuomotor transformations. In both experiments, reach targets composed of low SFs elicited the shortest latency and most prevalent SLRs, with SLR latency increasing and SLR prevalence decreasing for reach targets composed of progressively higher SFs. SLRs could be evoked in either the static or dynamic condition, and when present in experiment 2, were associated with shorter latency and larger magnitude corrections. The results in experiment 2 are consistent with a linkage between the forces produced by SLRs and the earliest portion of on-line reach corrections. Overall, our results demonstrate that stimuli composed of low SFs preferentially evoke the most rapid visuomotor responses that, in the context of rapidly correcting an on-going reaching movement, are associated with earlier and larger on-line reach corrections.

Splenius capitis: sensitive target for the cVEMP in older and neurodegenerative patients.

BACKGROUND: The vestibular evoked myogenic potential (VEMP) is a technique used to assess vestibular function. Cervical VEMPs (cVEMPs) are obtained conventionally from the sternocleidomastoid (SCM) muscle; however, the dorsal neck muscle splenius capitis (SPL) has also been shown to be a reliable target alongside the SCM in young subjects. OBJECTIVE: This study aimed to compare cVEMPs from the SCM and SPL in two positions across young, older, and Parkinson's disease (PD) patients. METHOD: Experiments were carried out using surface EMG electrodes placed over the SCM and SPL. cVEMPs were measured using a 30 s, 126 dB sound stimulus with 222 individual tone bursts, while subjects were in a supine and head-turned posture (also known as the head elevation method), and in a seated head-turned posture. RESULTS: When comparing cVEMPs across positions, the incidence of supine and seated SCM-cVEMPs diminished significantly in older and PD patients in comparison with young subjects. However, no statistically significant differences in incidences were found in seated SPL-cVEMPs when comparing young, older and PD patients. SPL-cVEMPs were present significantly more often than seated SCM-cVEMPs in PD patients. CONCLUSIONS: SPL-cVEMPs are not altered to the same extent that SCM-cVEMPs are by aging and disease and its addition to cVEMP testing may reduce false-positive tests for vestibulopathy.

A consensus guide to capturing the ability to inhibit actions and impulsive behaviors in the stop-signal task.

Response inhibition is essential for navigating everyday life. Its derailment is considered integral to numerous neurological and psychiatric disorders, and more generally, to a wide range of behavioral and health problems. Response-inhibition efficiency furthermore correlates with treatment outcome in some of these conditions. The stop-signal task is an essential tool to determine how quickly response inhibition is implemented. Despite its apparent simplicity, there are many features (ranging from task design to data analysis) that vary across studies in ways that can easily compromise the validity of the obtained results. Our goal is to facilitate a more accurate use of the stop-signal task. To this end, we provide 12 easy-to-implement consensus recommendations and point out the problems that can arise when they are not followed. Furthermore, we provide user-friendly open-source resources intended to inform statistical-power considerations, facilitate the correct implementation of the task, and assist in proper data analysis.

A rapid visuomotor response on the human upper limb is selectively influenced by implicit motor learning.

How do humans learn to adapt their motor actions to achieve task success? Recent behavioral and patient studies have challenged the classic notion that motor learning arises solely from the errors produced during a task, suggesting instead that explicit cognitive strategies can act in concert with the implicit, error-based, motor learning component. In this study, we show that the earliest wave of directionally tuned neuromuscular activity that begins within ~100 ms of peripheral visual stimulus onset is selectively influenced by the implicit component of motor learning. In contrast, the voluntary neuromuscular activity associated with reach initiation, which evolves ~100-200 ms later, is influenced by both the implicit and explicit components of motor learning. The selective influence of the implicit, but not explicit, component of motor learning on the directional tuning of the earliest cascade of neuromuscular activity supports the notion that these components of motor learning can differentially influence descending motor pathways. NEW & NOTEWORTHY Motor learning can be driven both by an implicit error-based component and an explicit strategic component, but the influence of these components on the descending pathways that contribute to motor control is unknown. In this study, we show that the implicit component selectively influences a reflexive circuit that rapidly generates a visuomotor response on the human upper limb. Our results show that the substrates mediating implicit and explicit motor learning exert distinct influences on descending motor pathways.