Cell Metabolism

Crosstalk

Sprinting Toward Fitness

*Correspondence: [email protected]
http://dx.doi.org/10.1016/j.cmet.2017.04.030

Intense intermittent exercise, or interval training, is a powerful stimulus to induce many of the physiological
adaptations typically associated with traditional, moderate-intensity continuous training. While coaches and
athletes have recognized the value of interval training to enhance performance for over a century, recent sci-
entific interest has focused on the application of this training method for health promotion. Despite renewed
attention, the mechanistic basis for the physiological remodeling that occurs after interval training and the
role that the stochastic nature of this type of exercise plays in mediating adaptive responses remains to
be elucidated.

Interval Training: Learning From and-error observations of innovative ath- While coaches and athletes have
The Past letes and coaches. The technique was pio- appreciated the effectiveness of interval
There is renewed scientific inquiry along neered in Finland by coach Lauri Pikhala training since the early 20th century, the
with widespread public interest in the po- with champion runners including Hannes first scientific publications on the physio-
tential for intense intermittent exercise to Kolehmainen and Paavo Nurmi. Nurmi logical basis of interval training for human
induce physiological adaptations that are was the most dominant distance runner performance did not appear until the
similar or even superior to traditional in the world between 1920 and 1930, win- 1960s. Over subsequent decades, the
endurance exercise in both healthy individ- ning nine Olympic gold medals. His system potential health-related applications of
uals and people with lifestyle-induced car- of training focused on running a high num- this type of training were increasingly
diometabolic disease (Gibala et al., 2012; ber of repetitions (>20 efforts) at close to recognized. In 1974, physiologists Ed-
Weston et al., 2014). Recent systematic re- race pace with short (<60 s) rest intervals. ward Fox and Donald Matthews from
views and meta-analyses have concluded In the 1930s, a German physician and The Ohio State University declared that
that interval training, or alternating periods coach, Woldemar Gerschler, along with ‘‘interval training is the supreme way to
of relatively intense exercise and recovery, cardiologist Herbert Reindel, devised a condition a person,’’ with the principles
can be a time-efficient strategy to enhance system of training that involved work and they described being applicable to ‘‘the
cardiorespiratory fitness (CRF), as deter- recovery periods based on heart rate (HR) coach, the athlete, and the person who
mined by whole-body maximal oxygen targets. An athlete would run over a short desires to condition himself for health pur-
uptake (VO2max) (Batacan et al., 2017). distance fast enough to elicit a HR of poses’’ (Fox and Mathews, 1974). Similar
These reports are particularly relevant 180 beats/min, followed by a rest period to Gerschler and Reindel some 40 years
because exercise capacity is a strong pre- in which HR dropped to 120 beats/min earlier, Fox and Mathews (1974) empha-
dictor of mortality, with a 1-metabolic before they commenced the next effort. sized the importance of the recovery
equivalent (MET, 3.5 mL O2/kg/min) higher Gerschler and Reindel proposed that the period or ‘‘relief interval’’ for optimizing
CRF associated with a 13% lower risk of recovery interval was the most important cardiovascular conditioning. Other re-
dying from all causes and being compara- aspect of their approach because it was searchers subsequently recognized the
ble to a 5-mm Hg reduction in systolic during this phase that the heart adapted, potential to apply interval training to
blood pressure or 1 mmol/L lower fasting allowing it to grow larger and stronger less-healthy individuals. In the mid-
plasma glucose concentration (Kodama (Figure 1). Perhaps the most celebrated 1990s, Katarina Meyer conducted pio-
et al., 2009). Given that ‘‘lack of time’’ is a case of interval training is Sir Roger Bannis- neering work on heart failure patients,
common barrier to regular physical ter, the first person to run the mile in under deeming the method better suited for
activity, the identification of time-efficient four minutes. While a medical student at St such individuals, as ‘‘interval exercise al-
exercise strategies that confer health ben- Mary’s Hospital, London, Bannister trained lows greater stimuli which patients prob-
efits could favorably impact public health during his lunch hour using the 9 min jog to ably would not have tolerated if the
by reducing the economic burden associ- a local track to warm up, after which he same intensity had been applied using
ated with inactivity-related disorders. promptly ran 10 3 400 m in a little over a continuous method’’ (Meyer et al., 1996).
The notion that interval training is new or 60 s each, with 2 min recovery. He then
a groundbreaking scientific approach to ran back to work, leaving 15 min to eat Sprint-Interval Training: Punching
physical conditioning needs to be placed his lunch and (hopefully) shower. On May Above Its Weight
in historical context. The basic practice 6, 1954, Bannister’s training culminated in Interval training is infinitely variable but
dates back to the early 20th century and a world mile record of 3 min, 59.4 s, two can be broadly classified into two
has evolved largely as a result of the trial- seconds faster than the previous record. categories: high-intensity interval training

988 Cell Metabolism 25, May 2, 2017 ª 2017 Elsevier Inc.

Crosstalk

cols involved a brief warm-up and cool- that closely resemble resistance exercise,
down totaling 5 min, such that SIT consti- yet elicit adaptations associated with
tuted 1 min of intense exercise within traditional endurance training. Training
a 10 min time commitment per session, volume has been proposed to be a pri-
whereas MICT involved 50 min of contin- mary determinant of the exercise-induced
uous exercise per session. VO2peak was increase in mitochondrial content in hu-
increased by 19% in both groups after man skeletal muscle, but other evidence
training, with similar training-induced im- highlights the potential role of exercise in-
provements in insulin sensitivity, as deter- tensity in mediating responses (MacInnis
Figure 1. Early Interval Training Research mined by intravenous glucose tolerate and Gibala, 2016).
German physician and coach, Woldemar Gersch- tests. Skeletal muscle mitochondrial con- A core principle of all training protocols
ler, together with cardiologist Dr. Herbert Reindel,
carried out ‘‘controlled training’’ studies in athletes
tent, assessed by the maximal activity of is that any acute exercise signal needs to
based on the concept of intermittent exercise. citrate synthase, also increased to a exceed a certain ‘‘threshold stimulus’’ to
Heart rate was monitored during all workouts with similar extent after SIT and MICT. These induce a variety of physiological adapta-
specific target zones prescribed during exercise
results (Gillen et al., 2016) are a timely tions that ultimately result in long-term
and recovery. The investigators found a 20% in-
crease in heart volume and improved performance reminder of the potency of SIT to stimu- phenotypic changes. Exercise provokes
after 21 days of training. The subjects included late physiologically meaningful and clini- widespread changes in numerous tissues
Gordon Pirie of Great Britain, who won the silver cally relevant improvements in health- and organs that are caused by the
medal in the 5,000 m at the 1956 Olympics and
subsequently coached one of the authors (J.A.H.). related outcomes with minimal time increased metabolic activity of active skel-
commitment. The findings also highlight etal muscle. To meet this challenge, multi-
a fundamental question regarding the ple integrated inter-organ responses
(HIIT), which typically denotes submaxi- mechanisms underpinning such robust function to blunt the homeostatic threats
mal efforts eliciting R80% of maximal whole-body and tissue-specific adapta- caused by the increased muscle energy
heart rate, and sprint interval training tions after interval training in humans. turnover and whole-body oxygen demand
(SIT), which involves ‘‘all out’’ efforts or Namely, how do a few hard sprints in (Hawley et al., 2014). During MICT lasting
an intensity corresponding to R100% of such a short intervention period elicit 1 hr, O2 supply is plentiful and substrate
the power output or speed that is associ- such profound remodeling of physiolog- demand by the active muscles is largely
ated with an individual’s VO2max (Weston ical systems? met by the oxidation of carbohydrate-
et al., 2014). SIT is a particularly potent and fat-based fuels. There is a primary reli-
variation of interval training, as demon- The Signal for Adaptation: Is ance on type I, slow-twitch muscle fibers,
strated by the classic study by Tabata Interval Training Different? and the rate of change of cellular dynamics
et al. (1996). These workers employed a Exercise has traditionally been catego- and disturbances to whole-body homeo-
protocol comprising eight 20 s sprints on rized as either aerobic/endurance or stasis is negligible. In contrast, both HIIT
a cycle ergometer, at an intensity corre- strength/power, with these extremes and SIT evoke extensive perturbations to
sponding to 170% of VO2max with 10 s of placed at opposite ends of a continuum. both local (muscle) and systemic (cardio-
recovery. When this protocol was per- Concomitant with the vastly different vascular, respiratory, neural, and hor-
formed five times per week for 6 weeks, functional and phenotypic outcomes monal) homeostasis. SIT in particular
VO2max was increased by a similar magni- induced by these exercise modes, the requires substantially higher absolute
tude to a protocol involving 5 hr/week of molecular pathways associated with power outputs compared to MICT, neces-
moderate-intensity cycling. The potency these divergent training protocols are sitating the recruitment of type II, fast-
of SIT to elicit adaptations comparable distinct (Hawley et al., 2014). In brief, twitch fibers. This in turn requires exten-
to traditional endurance training despite traditional endurance training elicits sive use of non-oxidative substrate meta-
large differences in training volume and changes that increase mitochondrial pro- bolism to meet muscle energy demands,
time commitment was recently demon- teins and the respiratory capacity of the which are fueled exclusively by intramus-
strated by Gillen and colleagues (2016). trained myofibers. These adaptations, in cular substrates (high-energy phosphates
These workers had two groups of seden- turn, underpin the altered patterns of sub- and glycogen) with little or no contribution
tary young men perform either SIT or strate oxidation during submaximal exer- from fat-based fuels. The greater absolute
moderate-intensity continuous training cise (from carbohydrate- to fat-based energy demand and altered fiber recruit-
(MICT) three times a week for 12 weeks. fuels) that result in less lactate production ment drives the higher absolute oxygen
The SIT workout comprised 3 3 20 s at any given submaximal power output flux and total fuel requirement of interval
‘‘all-out’’ sprints on a cycle ergometer at or speed. In contrast, strength and compared to low- to moderate-intensity
a power output of 500 W (a work-rate resistance-based training stimulates the continuous exercise. Accordingly, in
approximately 2- to 3-fold the power myofibrillar proteins responsible for mus- contrast to MICT, the rate of change of
output reached by these subjects at the cle hypertrophy, culminating in increases cellular dynamics and disturbances to
end of a VO2max test), with 2 min of low-in- in maximal contractile force output whole-body homeostasis induced by
tensity cycling (50 W) recovery between without substantial changes in fuel use intermittent exercise, and SIT in particular,
sprints. MICT consisted of 45 min of during exercise. A paradoxical character- is extensive.
continuous cycling at 110 W (moderate istic of interval training and SIT in partic- The ‘‘stop-start’’ nature of intermittent
intensity, 50% of VO2max). Both proto- ular is the brief, intense repeated efforts exercise and the associated intracellular

Cell Metabolism 25, May 2, 2017 989

Crosstalk

‘‘spikes’’ in various signaling pathways is in fine-tuning signaling responses after neering Research Council (227858-2010). J.A.H.
is supported by the Novo Nordisk Foundation
one potential mechanism to explain skel- SIT. Additional studies are needed, both
(NNF140C0011493) and an Australian Catholic
etal muscle responses to interval training, in terms of the early time course of molec- University Research Funding Program Grant
including superior adaptation after HIIT ular events that occur in human muscle in (2016000353).
compared to MICT despite matched response to repeated bouts of SIT and
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ACKNOWLEDGMENTS
the increased reactive O2 species (2014). High-intensity interval training in patients
with lifestyle-induced cardiometabolic disease: a
levels, acidosis, and altered redox state, Work by M.J.G. has been supported by an oper- systematic review and meta-analysis. Br. J. Sports
including NAD/NADH, may also play roles ating grant from the Natural Sciences and Engi- Med. 48, 1227–1234.

990 Cell Metabolism 25, May 2, 2017