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Physiological Adaptations to Interval Training to Promote Endurance Gibala, Bostad and Mccarthy 181 Available online at www.sciencedirect.com ScienceDirect Physiological adaptations to interval training to promote endurance Martin J Gibala, William Bostad and Devin G McCarthy The physiological determinants of endurance exercise endurance include the ‘lactate threshold’, generally performance are complex but depend in part on the capacity of defined as the exercise intensity at which lactate starts the cardiorespiratory system to transport oxygenated blood to to accumulate in blood, and critical power, which is working skeletal muscle, and the oxidative capacity of skeletal derived from the relationship between power output muscle, which is largely determined by mitochondrial content. and the time for which it can be sustained [1,2]. Both Interval training — characterized by intermittent bouts of higher factors are related to skeletal muscle oxidative capacity, intensity exercise interspersed by periods of lower-intensity which in turn is primarily determined by mitochondrial exercise or complete rest for recovery within a given session — volume density, although other mechanisms are involved. enhances endurance exercise performance including in those who are already well trained, although the mechanisms may be Traditional endurance training, characterized by a high different as compared to less-trained individuals. This brief volume of moderate-intensity continuous training review considers the evidence for cardiac and skeletal muscle (MICT), improves endurance exercise performance in responses to interval training that are associated with an part by increasing maximal cardiac output and thus oxy- improved capacity for whole-body aerobic energy provision, gen delivery to working skeletal muscle. MICT also with a focus on recent findings. stimulates mitochondrial biogenesis in skeletal muscle, which serves to increase the overall capacity for the Address oxidative metabolism of substrates [3]. In more highly Department of Kinesiology, McMaster University, Hamilton, Ontario, L8S trained individuals who already possess well-developed 4K1, Canada cardiovascular and muscle oxidative capacities, endur- Corresponding author: Gibala, Martin J ([email protected]) ance exercise performance can be further enhanced through intensified training that typically incorporates intervals, although the mechanisms involved are likely Current Opinion in Physiology 2019, 10:180–184 different compared to less-trained individuals [4]. This review comes from a themed issue on Exercise physiology Edited by Harry B Rossiter and Brian Glancy Interval training is characterized by intermittent bouts of For a complete overview see the Issue and the Editorial higher intensity exercise interspersed by periods of lower- Available online 30th May 2019 intensity exercise or complete rest for recovery within https://doi.org/10.1016/j.cophys.2019.05.013 a given training session. Intensity can be gauged in numerous ways, including based on absolute or relative 2468-8673/ã 2019 Elsevier Ltd. All rights reserved. power output, speed, heart rate, or perceived exertion. The terminology used to characterize interval training can be complex, owing in part to the large number of variables that can be manipulated and which interact to determine the precise training impulse, especially when viewed through the lens of program prescription for high- Introduction level athletes [5]. A simpler classification scheme, and one Endurance refers to the capacity to perform prolonged often employed to broadly categorize studies examining exercise, and is typically characterized in terms of the physiological responses to interval training, distinguishes highest velocity or power that can be sustained for a given two main types: high-intensity interval training (HIIT), period of time, or over a given distance. Numerous generally defined as relatively intense but submaximal physiological factors interact to determine endurance efforts that elicit 80% of maximal heart rate, and sprint capacity, and the precise mechanisms that regulate both interval training (SIT), a more intense version in which training adaptation and the limits of human performance bouts are performed in an ‘all out’ manner or at an remain topics of considerable interest to researchers and absolute workload that elicits VO2max [6]. practitioners [1,2]. One key determinant of endurance is maximal oxygen uptake (VO2max), which largely reflects This brief review considers cardiac and skeletal muscle the capacity of the cardiovascular system to deliver oxy- responses to interval training that are associated with genated blood to working skeletal muscle, although the improved endurance exercise capacity. The focus is on pulmonary system and other factors may be limiting in recent findings from studies mainly conducted in healthy some instances [1,2]. Other key determinants of but not highly trained individuals, which have established Current Opinion in Physiology 2019, 10:180–184 www.sciencedirect.com Physiological adaptations to interval training to promote endurance Gibala, Bostad and McCarthy 181 that interval training per se can induce physiological potential effect of SIT on cardiovascular adaptations, remodeling similar to or greater than MICT, often despite which could in turn explain the observed increases in a reduced total exercise volume or with reduced training VO2max. Stroke volume at submaximal exercise may not time commitment [6]. The interested reader is also be an accurate reflection of Qmax [25] however, and thus referred to other reviews that more comprehensively SIT-induced improvements in submaximal stroke vol- examine the mechanistic basis for adaptation [7 ,8 ], ume do not necessarily indicate that SIT is improving the application for performance in highly trained individ- Qmax. In the study by Astorino et al. [24], the effect if SIT uals [9,10] and the potential to enhance indices of cardi- per se on changes in Qmax cannot be isolated given the ometabolic health [11,12]. study design. O’Driscoll et al. [26] provided support for an effect of SIT on cardiac adaptations and demonstrated a Cardiac responses to interval training SIT-induced improvement in resting left ventricular end- A systematic review and meta-analysis by Milanovic diastolic function. Of note, there was no change in end- et al. [13 ] found that HIIT increases VO2max superior systolic function or ejection fraction [26], which suggests to MICT in healthy individuals when total exercise the potential SIT-induced improvement in Qmax could be volume or energy expenditure is matched. While the the result of improvements in diastolic function and/or improved VO2max after MICT is generally attributed to blood volume facilitating larger end-diastolic volumes increased maximal cardiac output (Qmax) [14], largely as opposed to cardiac contractility per se. However, mediated by haematological [15 ] and cardiac adaptations because Qmax was not measured in this study, the effect [16 ], the precise role of exercise intensity and volume in of these adaptations on Qmax cannot be established. mediating these adaptations is not well understood. A Given the limited number of studies, conflicting findings, recent study reported that a six week HIIT protocol and divergent methodologies employed, the effect of SIT involving four, 4-min efforts at 90% of maximal heart on Qmax and mechanisms underlying SIT-induced rate (HRmax) was more effective than work-matched improvements in VO2max, remain unclear. MICT at improving VO2max and Qmax in healthy individ- uals, although the HIIT-induced improvement in Qmax Emerging evidence is exploring the minimal volume of was not explained by an increase in blood volume [17 ]. brief intermittent exercise training that nonetheless Previously, Helgerud et al. [18] found that a similar HIIT induces physiological adaptation as reflected by an protocol conducted for eight weeks was more effective increased VO2max. Interestingly, reducing the volume than performing the same total work in a continuous of a SIT protocol by half did not diminish the increase manner at lower intensities for improving VO2max, and in VO2max [27]. Further, when a very low volume SIT this corresponded to an increase in stroke volume, sug- protocol involving two, 20-s bursts of ‘all out’ exercise was gesting a close link between the two. In contrast, another reduced to two, 10-s bouts, there was a blunted, yet recent study found that six weeks of HIIT improved measurable, increase in VO2max [28]. Spreading three, VO2max but did not induce cardiac structural or functional 20-s vigorous bouts of activity in the form of stair climbing adaptations [19]. Other studies have found that as little as throughout the day, separated by one to four hours, was six sessions of HIIT over two weeks can increase VO2max also recently shown to induce a modest improvement in the absence of measurable changes in cardiac and (5%) in VO2max [29]. As with MICT, there is also haematological factors, suggesting enhanced skeletal considerable inter-individual variability in the VO2max muscle oxygen extraction might contribute in this regard response to interval training [30], with one study showing [20]. The mechanisms explaining HIIT-induced changes the incidence of ‘non-response’ to either a SIT or MICT in VO2max including the magnitude and time course for protocol reduced by switching to the other training stim- potential cardiovascular responses requires further ulus [31]. A recent multi-center comparison of VO2max investigation. trainability between interval and traditional
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