It was determined that the MVIC, 60% MVIC and 10% MVIC conditions had deleterious effects on the subjects’ perception of time. More specifically, participants tended to underestimate the time intervals (estimated time was shorter than chronological time) across the different conditions compared to their chronological times (Figure 1). The higher intensity contraction conditions (MVIC and 60% MVIC) had more disturbance on time perception compared to the lower intensity 10% MVIC condition at the 30-second estimate. Lastly, the time estimates at 10-seconds were the most accurate when compared to estimates at 5-, 20-, and 30-seconds. There were no significant differences in time perception between the younger and older participants, even with the greater maximal and 60% submaximal contraction EMG activity of the younger cohort. The finding that the MVIC and 60% MVIC conditions yielded significant time underestimations compared to the control were in accord with the hypothesis. This time estimate disruption has been attributed to an intensity-dependent relationship between time perception and exercise, which has been found in other studies. Edwards and McCormick (2017) utilized cycling and had subjects estimate when 25%, 50%, 75%, and 100% of the trial was completed in different RPE conditions. They found that at the 75% and 100% intervals, time estimates for the RPE 20 condition (maximal exertion) was shortest when compared to RPE 11 (light intensity) and RPE 15 (moderate intensity). Subjects also completed a rowing task, where they found similar intensity-dependent results. Hanson and Lee (2020) investigated exercise intensity in individuals who self-selected their running pace. Results showed that participants significantly underestimated time when running at RPE 17 condition compared to RPE 11. Together with the results of the present study, these findings suggest that time is perceived to pass by more slowly when exercise intensity increases. This study also showed that the low-intensity, 10% MVIC contraction condition affected the participants’ time perception, a finding that contradicts the intensity-dependent results found by Edwards and McCormick (2017) and Hanson and Lee (2020). This may be attributed to the dual-task nature of this study. Participants were viewing a monitor, which displayed the force from their isometric contraction in real-time and asked to maintain a certain level of force. Maintaining the prescribed force while estimating the 5-, 10-, 20-, and 30-second time intervals may have impacted the participant’s ability to perceive time accurately. However, the distraction of viewing the monitor cannot be the primary factor underlying the underestimation of time, as the MVIC condition did not necessitate screen monitoring. Although the 60% MVIC condition was also a multi-task event with the distraction of viewing the monitor, maintaining a moderately intense contraction, and estimating time, time underestimation was not significantly different from the MVIC condition. Hence, while distractions (i.e., dual or multi-tasking) can affect time estimates, there was no additive adverse effect on the performance of moderate or high-intensity isometric contractions. The finding that exercise can lengthen an individual’s experience of time can be understood through the lens of the PAM (Gibbon et al. 1984, Grondin, 2010; Allman and Meck, 2012). With the MVIC and 60% MVC conditions, as participants isometrically contracted their knee extensors; muscle fatigue and discomfort may have been experienced due to the tension, partial blood occlusion, metabolite accumulation, and other factors. RPE values were higher with the MVIC and 60% MVIC illustrating the increased exertion at higher contraction intensities. This negative sensation acts as a form of physiological arousal (Edwards and Polman, 2013). Furthermore, the neuromuscular system will experience and contribute to heightened neural activity with increased motor unit recruitment and rate coding (firing frequency) (Behm 2004). Distractions (watching the computer monitor with 10% and 60% MVIC), increase sensory activity, and this overall increase in arousal may have an impact on their timing system. In the case of the PAM, arousal affects the mode switch in the clock stage. The mode switch is responsible for the storage of timing information, which is stored in an accumulator in a linear fashion. This timing information then passes through working and reference memory before a decision is made. Heightened arousal is thought to increase the rate at which the pacemaker processes information, resulting in extended perceptions of time intervals. Differences in attention, pacemaker speed, memory, and decision-making skills result in time perception differences (Allman and Meck, 2012). According to Dormal et al. (2017), exercise-induced arousal can produce this effect and generate distortion in time perception during exercise. Furthermore, attention was directed towards the monitor displaying their force production in the 10% and 60% MVIC conditions. According to the PAM, it is hypothesized that distraction away from the concept of time can cause the collection of pulses in the accumulator to begin at a later time (Droit-Volet and Gil, 2009, Hanson and Lee, 2020). With the distraction, the SB-FM suggests that oscillating neurons may synchronize and fire at a faster rate (Merchant et al., 2013). These physiological phenomena may affect the clock speed of the timing system, which is regulated by dopamine activity in the medial prefrontal cortex (Matell and Meck, 2004). Dopamine is directly involved with movement control as it modulates higher-order motor centers such as the basal ganglia with dopamine cells synapsing onto motoneurons’ dopamine receptors (Schwarz and Peever 2011). Both models suggest this state would cause the individual to estimate intervals of time to be longer than chronological time (retrospective timing) and to produce intervals of time that are shorter than chronological time (prospective timing). The greater time impairments with the higher intensity contractions (MVIC and 60% MVIC) suggest that heightened neuromuscular activity was more disruptive than the sensory distraction of watching the monitor during a low-intensity contraction (10% MVIC). Another difference is that distortions in time were found at all time point estimates in the present study. In contrast, Edwards and McCormick (2017) only found distortions when they had to estimate when 75% and 100% of the exercise bout time had elapsed (30-s of Wingate and 1200-s on a rowing ergometer). In the present study, participants were instructed to squeeze a hand trigger to estimate 5-, 10-, 20-, and 30-seconds. It is possible that having the participants engage in an additional motor task to squeeze the trigger interacted with the isometric knee extension contraction, causing an underestimation of time early in the time trial. It was anticipated that time variability would steadily increase as participants estimated the four consecutive times. As time progresses, you would naturally expect a greater variability as small errors made early may be amplified as the duration of the trial progresses. This effect was observed with 20-s and 30-s intervals demonstrating greater underestimations than 10-s. However, it is interesting that subjects also underestimated time more at 5-seconds compared to 10-seconds (Figure 1). No other significant differences between times were found in the analysis. Edwards and McCormick (2017) found no deficits at 50% of total time (which corresponds to the 10-second estimate). However, their study utilized another method to quantify time perception. Instead of measuring the variability for each time point (i.e., referring to a pre-test value), they compared their time estimates to chronological time and had no control group to compare estimates. Hanson and Lee (2020) utilized a similar protocol and found no differences between any of the time estimates. The finding that time estimates at 10-seconds were more accurate compared to 5-, 20-, and 30-seconds may be attributed to lifelong learning. Countdowns from 10-seconds are commonly used in our society, from rocket take-offs to space, the countdown to the New Year and the end of many time-restricted sports. Many films from the mid-twentieth century included 10-second countdowns before the movie began. As the subjects in this study were recreationally active, they may also be accustomed to 10-second intervals from sports and exercise, where it is common for a trainer to push athletes by saying, “only 10-seconds remaining”. It is speculated that this additional lifelong exposure of 10-second time intervals led subjects to estimate the 10-second time point most accurately. The results did not show any differences in time perception between the younger and older cohorts. It was expected that older adults would be less accurate and more variable in their timing compared to their younger counterparts (Wittmann and Lehnhoff, 2005). Coelho et al. (2004) suggested that with age, internal clock speeds up, such that older adults tend to underestimate time compared to younger adults. Nevertheless, their study was not exercise-related and the findings lost statistical significance when controlled for literacy. However, the more plausible explanation is that the internal clock becomes slower with age (Block et al., 1999; Bherer et al., 2007). This means people of advanced age tend to understimate and over-produce intervals relative to chronological time (i.e., a person with a slow internal clock may perceive a 5-second stimulus as lasting only 3 seconds, and when asked to produce a 3-second interval, instead produce a 5-second one). Furthermore, cognitive aging of the ventral tegmental area (VTA) has been shown to decrease dopamine levels, which could also explain why older adults seem to have a slower internal clock (Peterson et al., 2017). The SB-FM theory suggests that our internal timekeeping mechanism begins with a phasic release of dopamine from dopaminergic midbrain projections to the cortex and dorsal striatum at the onset of the “to-be-timed” interval (Matell and Meck, 2004). This causes groups of cortical neurons to reset, synchronize their firing, and begin oscillating at their respective periods in the dorsal striatum (Allman and Meck, 2012). These dopamine-reliant oscillating neurons are essentially the clock mechanism, and thus age-related declines could adversely affect senior’s time perception. Interestingly, a review by Turgeon et al. (2016) concluded that alternative neural strategies can mask age-related declines in time perception, allowing older adults to perform nearly or as well as younger adults until cognitive or physical demands push them past the threshold for compensation. It was proposed that seniors rely more on the cortico-cerebellar and hippocampal regions which are recruited to the timing system and are less affected by aging (Meck, 2005; Merchant et al., 2013; Lusk et al., 2016). Furthermore, older adults seem to use additional cognitive resources and external cues to increase their reliance on their internal timing networks, their reliance on feedback, and adaptive corrections to perform well at time perception tasks. When interrupting older adults’ ability to use these systems, the actual age-related deficits in time perception become more apparent (Turgeon and Wing, 2012). Such a threshold appears not to be reached in this study, meaning that the older adults were able to compensate for their potentially slower internal clock using the above circuitry. As the subjects were physically active individuals, they may have relatively well-developed and active cortico-cerebellar regions, allowing for efficient compensation of their proposed slower internal clock. Furthermore, the older adults recruited for this study were quite educated (50% of participants had university-level education) and were physically active across the lifespan. Being physically (Hillman et al., 2008) and mentally (Coelho et al. 2004, Valenzuela and Sachdev, 2006) active throughout the lifetime may have offered similar neuroprotective effects. It was expected for participants to experience significant deficits in timing during the MVIC and 60% MVIC contractions. In theory, these exercise intensities should be high enough to break the threshold of compensation for older adults. However, this was not observed in this study. Though the old adults were physically active, most were not accustomed to high-intensity anaerobic work. This would suggest that some participants, even following the familarization session may not have been performing true MVICs and were contracting at a lower intensity during the protocols. There was no significant difference in RPE between the MVIC and 60% MVIC suggesting the participants experienced similar exertion levels with both conditions. With a prolonged MVIC, EMG would be expected to decrease over time due to derecruitment of motor neurons and attenuated rate coding (Behm 2004). However, EMG activity increased by 11.5% and 19.6% in the young and elderly participants respectively during the 30-second MVIC protocol (Table 5). This increase in muscle activity rather than decrease suggests that the participants may have subconsciously paced themselves in preparation for a 30-second MVIC and the older group paced (contracted at a lower relative contraction intensity) to a greater extent than the younger group. As such, it might be possible that the older adults were unknowingly contracting just until they reached the threshold for compensation, allowing them to estimate the time intervals as accurately as their younger counterparts. Increases in body temperature have been suggested to distort temporal perception (Pieron (1923; 1945), with small circadian increases (i.e., afternoon) inducing time overestimation (Hoagland 1933) or warm water immersion (380C) leading to underestimation of time (van Maanen et al., 2019). Time underestimation was reported when core temperature increased with running in a warm, humid environment (Tamm et al. 2015). However, there was no significant increase in body temperature in the present study. There was also no significant differences in heart rate between the experimental conditions with only the control condition experiencing lower heart rates. Thus, neither heart rate nor tympanic temperature were sensitive indicators of changes in arousal. |