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Transcranial magnetic stimulation (TMS) has emerged as an important technique in cognitive neuroscience, permitting causal inferences about the contribution of a given brain area to behavior. Despite widespread use, exactly how TMS influences neural activity throughout an interconnected network, and how such influences ultimately change behavior, remain unclear. The oculomotor system of nonhuman primates (NHPs) offers a potential animal model to bridge this gap. Here, based on results suggesting that neck muscle activity provides a sensitive indicator of oculomotor activation, we show that single pulses of TMS over the frontal eye fields (FEFs) in awake NHPs evoked rapid (within ˜25 ms) and fairly consistent (˜50-75% of all trials) expression of a contralateral head-turning synergy. This neck muscle response resembled that evoked by subsaccadic electrical microstimulation of the FEF. Systematic variation in TMS location revealed that this response could also be evoked from the dorsolateral prefrontal cortex (dlPFC). Combining TMS with an oculomotor task revealed state dependency, with TMS evoking larger neck muscle responses when the stimulated area was actively engaged. Together, these results advance the suitability of the NHP oculomotor system as an animal model for TMS. The polysynaptic neck muscle response evoked by TMS of the prefrontal cortex is a quantifiable trial-by-trial reflection of oculomotor activation, comparable to the monosynaptic motor-evoked potential evoked by TMS of primary motor cortex. Our results also speak to a role for both the FEF and dlPFC in head orienting, presumably via subcortical connections with the superior colliculus. [ABSTRACT FROM AUTHOR]

Visual stimulus presentation activates the oculomotor network without requiring a gaze shift. Here, we demonstrate that primate neck muscles are recruited during such reflexive covert orienting in a manner that parallels activity recorded from the superior colliculus (SC). Our results indicate the presence of a brainstem circuit whereby reflexive covert orienting is prevented from shifting gaze, but recruits neck muscles, predicting that similarities between SC and neck muscle activity should extend to other cognitive processes that are known to influence SC activity. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The superior colliculus (SC) has been increasingly implicated in the rapid processing of evolutionarily relevant stimuli like faces, but the behavioural relevance of such processing is unclear. The SC has also been implicated in the generation of express visuomotor responses (EVR), which are very short-latency (~80 ms) bursts of muscle activity time-locked to visual target presentation. These observations led us to investigate the influence of faces on EVRs. We recorded upper limb muscle activity from healthy participants as they reached toward targets in the presence of a distractor. In some experiments, faces were used as stimuli. Across blocks of trials, we varied the instruction as to which stimulus served as the target or distractor. Doing so allowed us to assess the impact of instruction on muscle recruitment given identical visual stimuli. We found that responses were uniquely modulated in tasks involving high-contrast faces, promoting reaches toward or away from a face depending on instruction. Follow-up experiments confirmed that the phenomenon required highly salient repeated faces and was not observed to non-facial stimuli nor to faces expressing different affects. This study extends the hypothesis that the SC mediates the EVR by demonstrating that faces impact muscle recruitment at short latencies that precede cortical activity for face perception. Our results constitute direct evidence for the behavioural relevance of face detection in the brainstem, and also implicate a role for top-down cortical pre-setting of the EVR depending on task context., (© 2024 The Author(s). European Journal of Neuroscience published by Federation of European Neuroscience Societies and John Wiley & Sons Ltd.)

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., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

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., (Copyright © 2023 the authors.)

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., (Copyright © 2023 Cecala et al.)

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., (Copyright © 2023 the authors.)

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., (© 2020. Springer Science+Business Media, LLC, part of Springer Nature.)

A distributed network of cortical and subcortical brain areas controls our oculomotor behavior. This network includes the superior colliculus (SC), which coordinates an ancient visual grasp reflex via outputs that ramify widely within the brainstem and spinal cord, accessing saccadic and other premotor and autonomic circuits. In this Review, we discuss recent results correlating subliminal SC activity in the absence of saccades with diverse components of the visual grasp reflex, including neck and limb muscle recruitment, pupil dilation, and microsaccade propensity. Such subtle manifestations of covert orienting are accessible in the motor periphery and may provide the next generation of oculomotor biomarkers in health and disease., (Copyright © 2014 Elsevier Inc. All rights reserved.)

Adaptive adjustments of strategies help optimize behavior in a dynamic and uncertain world. Previous studies in the countermanding (or stop-signal) paradigm have detailed how reaction times (RTs) change with trial sequence, demonstrating adaptive control of movement generation. Comparatively little is known about the adaptive control of movement cancellation in the countermanding task, mainly because movement cancellation implies the absence of an outcome and estimates of movement cancellation require hundreds of trials. Here, we exploit a within-trial proxy of movement cancellation based on recordings of neck muscle activity while human subjects attempted to cancel large eye-head gaze shifts. On a subset of successfully cancelled trials where gaze remains stable, small head-only movements to the target are actively braked by a pulse of antagonist neck muscle activity. The timing of such antagonist muscle recruitment relative to the stop signal, termed the "antagonist latency," tended to decrease or increase after trials with or without a stop-signal, respectively. Over multiple time scales, fluctuations in the antagonist latency tended to be the mirror opposite of those occurring contemporaneously with RTs. These results provide new insights into the adaptive control of movement cancellation at an unprecedented resolution, suggesting it can be as prone to dynamic adjustment as movement generation. Adaptive control in the countermanding task appears to be governed by a dynamic balance between movement cancellation and generation: shifting the balance in favor of movement cancellation slows movement generation, whereas shifting the balance in favor of movement generation slows movement cancellation.