Summary: | The physical limits of the human being have been the object of study for a considerable time. Human and exercise physiology, in combination with multiple other related disciplines, studied the function of the organs and their relationship during exercise. When studying the mechanisms causing the limits of the human body, most of the research has focused on the locomotor muscles, lungs and heart. Therefore, it is not surprising that the limit of the performance has predominantly been explained at a "peripheral" level. Many studies have successfully demonstrated how performance can be improved (or not) by manipulating a "peripheral" parameter. However, in most cases it is the brain that regulates and integrates these physiological functions, and much of the contemporary literature has ignored its potential role in exercise performance. This may be because moderating brain function is fraught with difficulty, and challenging to measure. However, with the recent introduction and development of new non-invasive devices, the knowledge regarding the behaviour of the central nervous system during exercise can be advanced. Transcranial direct current stimulation (tDCS) and repetitive transcranial magnetic stimulation (rTMS) are two such methods. These methods can transiently moderate the activity of a targeted brain area, potentially altering the regulation of a particular physiological (or psychological) system, and consequently eliciting a change in exercise performance. Despite the promising theory, there is little or no experimental data regarding the potential to moderate neurophysiological mechanisms through tDCS to improve exercise performance. Consequently, the experiments performed as part of this thesis investigated the capacity for tDCS to alter physical performance. The ability of tDCS as a targeted and selective intervention at the brain level provides the unique opportunity to reduce many methodological constraints that might limit or confound understanding regarding some of the key physiological mechanisms during exercise. Therefore, the primary aim of this thesis was to investigate how tDCS may moderate both central and peripheral neurophysiological mechanisms, and how this may effect various exercise tasks. The first study investigated the effect of a well-documented analgesic tDCS montage on exercise-induced muscle pain. This study demonstrated for the first time, that although anodal tDCS of the motor cortex (M1) reduces pain in a cold pressor task, it does not elicit any reduction in exercise-induced muscle pain and consequently has no effect on exercise performance. As reductions in exercise-induced pain have previously been documented to improve performance, probably the lack of effect was due to either the M1 having a limited processing role in exercise-induced pain, or that the cathodal stimulation of the prefrontal cortex negated any positive impact of anodal M1 stimulation. Given the lack of guidelines for tDCS electrode montage for exercise, the second study examined the effect of different electrode montages on isometric performance and the neuromuscular response of knee extensor muscle. Given that the anode increases excitability and the cathode decreases excitability, the placement of these has the potential to elicit significant effects on exercise performance. The results showed that exercise performance improved only when an extrachepalic tDCS montage was applied to the M1, but in the absence of changes to the measured neuromuscular parameters. These results suggest that tDCS can have a positive effect on single limb submaximal exercise, but not on maximal muscle contraction. The improvement in performance was probably the consequence of the reduction in perceived exertion for a given load. This is the first experiment showing an improvement in exercise performance on single joint exercise of the lower limbs following tDCS. The results suggest that the extrachepalic set-up is recommended for exercise studies in order to avoid any potential negative effect of the cathodal electrode. Previous studies investigating tDCS have shown its potential to alter autonomic activity, and in some circumstances reduce the cardiovascular response during exercise. Considering the emerging studies and applications of tDCS on exercise and the potential benefits of tDCS in the treatment of cardiovascular diseases, the third study monitored multiple cardiovascular variables following tDCS in a group of healthy volunteers. Using more advanced techniques and methods compared to previous research, including the post exercise ischemia technique and transthoracic bioimpedance, the results suggest that tDCS administration has no significant effect on the cardiovascular response in healthy individuals. The final study sought to apply the findings obtained in the study 2 to whole body exercise. The same extrachepalic set up was applied over both the motor cortices, with both anodal and cathodal stimulation conditions. The neuromuscular response and cycling performance was also monitored. Following anodal tDCS, time to exhaustion and motor cortex excitability of lower limbs increased. Interestingly, cathodal stimulation did not induce any change in cycling performance or neuromuscular response. This study demonstrated for the first time the ability of anodal tDCS to improve performance of a constant load cycling task, and highlights the inability of cathodal tDCS to decrease cortical activation during muscle contraction. Taken together, the experiments performed as part of this thesis provide new insights on how brain stimulation influences exercise performance, with notable findings regarding the role of M1 excitability and perception of effort. Furthermore, considering the lack of knowledge regarding the use of tDCS on exercise, these findings will help further understanding of how to apply tDCS in exercise science. This consequently improves the knowledge base regarding the effect of tDCS on exercise and provides both a methodological and theoretical foundation on which future research can be based.
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