Response triggering by an acoustic stimulus increases with stimulus intensity and is best predicted by startle reflex activation

In a simple reaction time task, the presentation of a startling acoustic stimulus has been shown to trigger the prepared response at short latency, known as the StartReact effect. However, it is unclear under what conditions it can be assumed that the loud stimulus results in response triggering. The purpose of the present study was to examine how auditory stimulus intensity and preparation level affect the probability of involuntary response triggering and the incidence of activation in the startle reflex indicator of sternocleidomastoid (SCM). In two reaction time experiments, participants were presented with an irrelevant auditory stimulus of varying intensities at various time points prior to the visual go-signal. Responses were independently categorized as responding to either the auditory or visual stimulus and those with or without SCM activation (i.e., SCM+/−). Both the incidence of response triggering and proportion of SCM+ trials increased with stimulus intensity and presentation closer to the go-signal. Data also showed that participants reacted to the auditory stimulus at a much higher rate on trials where the auditory stimulus elicited SCM activity versus those that did not, and a logistic regression analysis confirmed that SCM activation is a reliable predictor of response triggering for all conditions.

Does exposure to startle impact voluntary reaching movements in individuals with severe-to-moderate stroke?

When movements of individuals with stroke (iwS) are elicited by startling acoustic stimulus (SAS), reaching movements are faster, further, and directed away from the body. However, these startle-evoked movements also elicit task-inappropriate flexor activity, raising concerns that chronic exposure to startle might also induce heightened flexor activity during voluntarily elicited movement. The objective of this study is to evaluate the impact of startle exposure on voluntary movements during point-to-point reaching in individuals with moderate and severe stroke. We hypothesize that startle exposure will increase task-inappropriate activity in flexor muscles, which will be associated with worse voluntarily initiated reaching performance (e.g. decreased distance, displacement, and final accuracy). Eleven individuals with moderate-to-severe stroke (UEFM = 8–41/66 and MAS = 0–4/4) performed voluntary point-to-point reaching with 1/3 of trials elicited by an SAS. We used electromyography to measure activity in brachioradialis (BR), biceps (BIC), triceps lateral head (TRI), pectoralis (PEC), anterior deltoid (AD), and posterior deltoid (PD). Conversely to our hypothesis, exposure to startle did not increase abnormal flexion but rather antagonist activity in the elbow flexors and shoulder horizontal adductors decreased, suggesting that abnormal flexor/extensor co-contraction was reduced. This reduction of flexion led to increased reaching distance (18.2% farther), movement onset (8.6% faster), and final accuracy (16.1% more accurate) by the end of the session. This study offers the first evidence that exposure to startle in iwS does not negatively impact voluntary movement; moreover, exposure may improve volitionally activated reaching movements by decreasing abnormal flexion activity.

Bimanual but not unimanual finger movements are triggered by a startling acoustic stimulus: evidence for increased reticulospinal drive for bimanual responses

The relative contributions of reticulospinal and corticospinal pathways to movement initiation are relatively unknown but appear to depend on the involved musculature. Here, we show that unimanual finger movements, which are predominantly initiated via corticospinal pathways, are not triggered at short latency by a startling acoustic stimulus (SAS), while bimanual finger movements are triggered by the SAS. This distinction is attributed to increased reticulospinal drive for bilateral responses.

Understanding the dual-task costs of walking: a StartReact study

The need to perform multiple tasks more or less simultaneously is a common occurrence during walking in daily life. Performing tasks simultaneously typically impacts task performance negatively. Hypothetically, such dual-task costs may be explained by a lowered state of preparation due to competition for attentional resources, or alternatively, by a ‘bottleneck’ in response initiation. Here, we investigated both hypotheses by comparing ‘StartReact’ effects during a manual squeezing task under single-task (when seated) and dual-task (when walking) conditions. StartReact is the acceleration of reaction times by a startling stimulation (a startling acoustic stimulus was applied in 25% of trials), attributed to the startling stimulus directly releasing a pre-prepared movement. If dual-task costs are due to a lowered state of preparation, we expected trials both with and without an accompanying startling stimulus to be delayed compared to the single-task condition, whereas we expected only trials without a startling stimulus to be delayed if a bottleneck in response initiation would underlie dual-task costs. Reaction times of the manual squeezing task in the flexor digitorum superficialis and extensor carpi radialis muscle were significantly delayed (approx. 20 ms) when walking compared to the seated position. A startling acoustic stimulus significantly decreased reaction times of the squeezing task (approx. 60 ms) both when walking and sitting. Dual-task costs during walking are, therefore, likely the result of lowered task preparation because of competition for attentional resources.

StartReact effects are dependent on engagement of startle reflex circuits: support for a subcortically mediated initiation pathway

The “StartReact” effect refers to the rapid involuntary triggering of a prepared movement in response to a loud startling acoustic stimulus (SAS). This effect is typically confirmed by the presence of short-latency electromyographic activity in startle reflex-related muscles such as the sternocleidomastoid (SCM); however, there is debate regarding the specific neural pathways involved in the StartReact effect. Some research has implicated a subcortically mediated pathway, which would predict different response latencies depending on the presence of a startle reflex. Alternatively, other research has suggested that this effect involves the same pathways responsible for voluntary response initiation and simply reflects higher preparatory activation levels, and thus faster voluntary initiation. To distinguish between these competing hypotheses, the present study assessed preparation level during a simple reaction time (RT) task involving wrist extension in response to a control tone or a SAS. Premotor RT and startle circuitry engagement (as measured by SCM activation) were determined for each trial. Additionally, preparation level at the go signal on each trial was measured using motor-evoked potentials (MEP) elicited by transcranial magnetic stimulation (TMS). Results showed that SAS trial RTs were significantly shorter ( P = 0.009) in the presence of startle-related SCM activity. Nevertheless, preparation levels (as indexed by MEP amplitude) were statistically equivalent between trials with and without SCM activation. These results indicate that the StartReact effect relates to engagement of the startle reflex circuitry rather than simply being a result of an increased level of preparatory activation. NEW & NOTEWORTHY The neural mechanism underlying the early triggering of goal-directed actions by a startling acoustic stimulus (SAS) is unclear. We show that although significant reaction time differences were evident depending on whether the SAS elicited a startle reflex, motor preparatory activation was the same. Thus, in a highly prepared state, the short-latency responses associated with the StartReact effect appear to be related to engagement of startle reflex circuitry, not differences in motor preparatory level.

High-intensity transcranial magnetic stimulation reveals differential cortical contributions to prepared responses

Corticospinal output pathways have typically been considered to be the primary driver for voluntary movements of the hand/forearm; however, more recently, reticulospinal drive has also been implicated in the production of these movements. Although both pathways may play a role, the reticulospinal tract is thought to have stronger connections to flexor muscles than to extensors. Similarly, movements involuntarily triggered via a startling acoustic stimulus (SAS) are believed to receive greater reticular input than voluntary movements. To investigate a differential role of reticulospinal drive depending on movement type or acoustic stimulus, corticospinal drive was transiently interrupted using high-intensity transcranial magnetic stimulation (TMS) applied during the reaction time (RT) interval. This TMS-induced suppression of cortical drive leads to RT delays that can be used to assess cortical contributions to movement. Participants completed targeted flexion and extension movements of the wrist in a simple RT paradigm in response to a control auditory go signal or SAS. Occasionally, suprathreshold TMS was applied over the motor cortical representation for the prime mover. Results revealed that TMS significantly increased RT in all conditions. There was a significantly longer TMS-induced RT delay seen in extension movements than in flexion movements and a greater RT delay in movements initiated in response to control stimuli compared with SAS. These results suggest that the contribution of reticulospinal drive is larger for wrist flexion than for extension. Similarly, movements triggered involuntarily by an SAS appear to involve greater reticulospinal drive, and relatively less corticospinal drive, than those that are voluntarily initiated.NEW & NOTEWORTHY Through the use of the transcranial magnetic stimulation-induced silent period, we provide novel evidence for a greater contribution of reticulospinal drive, and a relative decrease in corticospinal drive, to movements involuntarily triggered by a startle compared with voluntary movements. These results also provide support for the notion that both cortical and reticular structures are involved in the neural pathway underlying startle-triggered movements. Furthermore, our results indicate greater reticulospinal contribution to wrist flexion than extension movements.

Response preparation and execution during intentional bimanual pattern switching

During continuous bimanual coordination, in-phase (IP; 0° relative phase) and anti-phase (AP; 180° relative phase) patterns can be stably performed without practice. Paradigms in which participants are required to intentionally switch between these coordination patterns have been used to investigate the interaction between the performer’s intentions and intrinsic dynamics of the body’s preferred patterns. The current study examined the processes associated with switching preparation and execution through the use of a startling acoustic stimulus (SAS) as the switch stimulus. A SAS is known to involuntarily trigger preprogrammed responses at a shortened latency and, thus, can be used to probe advance preparation. Participants performed cyclical IP and AP bimanual elbow extension-flexion movements in which they were required to switch patterns in response to an auditory switch cue, which was either nonstartling (80 dB) or a SAS (120 dB). Results indicated that reaction time to the switch stimulus (i.e., switch onset) was significantly reduced on startle trials, indicative of advance preparation of the switch response. Similarly, switching time was reduced on startle trials, which was attributed to increased neural activation caused by the SAS. Switching time was also shorter for AP to IP trials, but only when the switching stimulus occurred at either the midpoint or reversal locations within the movement cycle, suggesting that the switch location may affect the intrinsic dynamics of the system. NEW & NOTEWORTHY The current study provides novel information regarding preparation and execution of intentional switching between in-phase and anti-phase bimanual coordination patterns. Using a startling acoustic stimulus, we provide strong evidence that the switching response is prepared before the switch stimulus, and switch execution is accelerated by the startling stimulus. In addition, the time required to switch between patterns and relative limb contribution is dependent upon where in the movement cycle the switch stimulus occurred.

Go-activation endures following the presentation of a stop-signal: evidence from startle

It has been proposed that, in a stop-signal task (SST), independent go- and stop-processes “race” to control behavior. If the go-process wins, an overt response is produced, whereas, if the stop-process wins, the response is withheld. One prediction that follows from this proposal is that, if the activation associated with one process is enhanced, it is more likely to win the race. We looked to determine whether these initiation and inhibition processes (and thus response outcomes) could be manipulated by using a startling acoustic stimulus (SAS), which has been shown to provide additional response activation. In the present study, participants were to respond to a visual go-stimulus; however, if a subsequent stop-signal appeared, they were to inhibit the response. The stop-signal was presented at a delay corresponding to a probability of responding of 0.4 (determined from a baseline block of trials). On stop-trials, a SAS was presented either simultaneously with the go-signal or stop-signal or 100, 150, or 200 ms following the stop-signal. Results showed that presenting a SAS during stop-trials led to an increase in probability of responding when presented with or following the stop-signal. The latency of SAS responses at the stop-signal + 150 ms and stop-signal + 200 ms probe times suggests that they would have been voluntarily inhibited but instead were involuntarily initiated by the SAS. Thus results demonstrate that go-activation endures even 200 ms following a stop-signal and remains accessible well after the response has been inhibited, providing evidence against a winner-take-all race between independent go- and stop-processes. NEW & NOTEWORTHY In this study, a startling acoustic stimulus (SAS) was used to determine whether response outcome could be manipulated in a stop-signal task. Results revealed that presenting a SAS during stop-signal trials led to an increase in probability of responding even when presented 200 ms following the stop-signal. The latency of SAS responses indicates that go-activation remains accessible and modifiable well after the response is voluntarily inhibited, providing evidence against an irrevocable commitment to inhibition.