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Scand J Med Sci Sports 2010: 20 (Suppl.

2): 1123 & 2010 John Wiley & Sons A/S


doi: 10.1111/j.1600-0838.2010.01193.x

Review

Speed endurance training is a powerful stimulus for physiological


adaptations and performance improvements of athletes
F. M. Iaia, J. Bangsbo
Department of Exercise and Sport Sciences, Section of Human Physiology, Copenhagen Muscle Research Centre, University of
Copenhagen, Copenhagen, Denmark
Corresponding author: Jens Bangsbo, Department of Exercise and Sport Sciences, Section of Human Physiology, Copenhagen
Muscle Research Centre, University of Copenhagen, August Krogh Building, Universitetsparken 13, DK2100 Copenhagen ,
Denmark. Tel: 145 35 32 16 22, Fax: 145 35 32 16 00, E-mail: jbangsbo@i.ku.dk
Accepted for publication 28 March 2010

The present article reviews the physiological and performance benet from performing speed endurance training. These
eects of speed endurance training consisting of exercise improvements dont appear . to depend on changes in max-
bouts at near maximal intensities in already trained subjects. imum oxygen uptake (VO2 max ), muscle substrate levels,
Despite a reduction in training volume, speed endurance glycolytic and oxidative enzymes activity, and membrane
training of endurance-trained athletes can maintain the transport proteins involved in pH regulation. Instead they
oxidative capacity and improve intense short-duration/re- appear to be related to a reduced energy expenditure during
peated high-intensity exercise performance lasting 30 s to submaximal exercise and a higher expression of muscle
4 min, as it occurs in a number of sports. When combined Na1,K1 pump a-subunits, which via a higher Na1,K1 pump
with a basic volume of training including some aerobic high- activity during exercise may delay fatigue development
intensity sessions, speed endurance training is also useful in during intense exercise. In conclusion, athletes from dis-
enhancing performance during longer events, e.g. 40 K ciplines involving periods of intense exercise can benet
cycling and 10 K running. Athletes in team sports involving from the inclusion of speed endurance sessions in their
intense exercise actions and endurance aspects can also training programs.

The literature contains a high number of studies which consists of maximal short-duration (210 s)
utilizing training at intensities higher. than the one exercise bouts followed by long recovery periods (50
eliciting maximum oxygen uptake (VO2 max ), which 100 s) (Reilly & Bangsbo, 1998). Speed endurance
have all indistinctively been referred to as sprint, training is used to describe all the other anaerobic
intermittent and interval training (Ross & Leveritt, intensities and has been divided into production and
2001; Laursen & Jenkins, 2002). It is often not maintenance training (Reilly & Bangsbo, 1998). In
recognized that these represent a broad variety of production training the exercise bouts last less
training protocols and a wide range of exercise than 40 s and are performed at near maximal
intensities.
. For example, a well-trained cyclist may intensity, while the recovery periods are compara-
reach VO2 max at a power output of 400 W and be able tively long (45 times the exercise duration) in order
to produce a peak power output (PPO) of  1200 W to perform maximally in the subsequent exercise
in a 10-s maximal
. test (Calbet et al., 2003). In this bouts. Maintenance training includes exercise
instance, VO2 max is obtained at  33% of PPO and bouts of 590 s with shorter rest periods in between
the remaining  67% represents exercise intensities the intervals ( 3 fold longer than the exercise time),
above the maximal aerobic power (which is dened resulting in a slightly lower intensity and a progres-
as supra maximal exercise). Dierences in the relative sive accumulation of fatigue as the training continues
exercise intensity and duration of recovery periods (Table 1).
should be taken into account when comparing scien- The present brief review deals with the eect of
tic studies using supramaximal exercise. To clarify speed endurance training on performance and phy-
the issue, a common terminology would be helpful. siological adaptations, but will mainly cover the
Anaerobic training is dened as training. where the eect of speed endurance training at near maximal
exercise is performed at intensities above VO2 max and intensities (470% of maximal intensity) in already
where the primary aim is to stimulate the anaerobic trained subjects. For the eects of exercise training at
energy production (Bangsbo, 1994). Part of the intensities around .or slightly higher than the one
anaerobic training is classied as speed training, corresponding to VO2 max , the reader is referred to

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Iaia & Bangsbo
Table 1. Types of anaerobic training

Type of anaerobic training Exercise intensity Duration of exercise (s) Duration of recovery No. of repetitions
(% of maximum speed)

Speed 100 210 50100 s 520


Speed endurance production 70100 1040 45 times exercise duration 312
Speed endurance maintenance 50100 590 13 times exercise duration 225

the articles by Laursen and Mujika in the present (a)


journal issue. A signicant number of studies have
examined the muscle adaptations with speed endur-
ance training in untrained subjects (Ross & Leveritt,
2001; Burgomaster et al., 2005; Gibala et al., 2006).
However, these are of limited interest in the discus-
sion of how to train athletes, but will be included
where they may add information to the data ob-
tained in athletes.

Physiological response to speed endurance training


Before discussing its eects, it is useful to present the
physiological and metabolic response to speed en-
durance training. In a study by Mohr et al. (2007), a
group of physical active
. subjects performed eight 30- (b)
s runs at  130% of VO2 max (158170 m) separated
by 90 s of rest. The heart rate uctuated with peaks
around 97% of the maximal heart rate (HRmax) and
mean heart rate of  84% of HRmax, reecting a
high cardiovascular stimulus. Blood lactate concen-
tration [Lac  ] increased progressively during the
session from 1.1 to 16.5 mM (Fig. 1a), with muscle
[Lac  ] rising to  45 mmol/kg d.w. Correspond-
ingly, muscle pH was lowered to 6.98 during the
training. Plasma K1 concentration varied through-
out the training session and peaked at 6.3 mmol/L
(Fig. 1b). Immediately after the training session
muscle creatine phosphate (PCr) and glycogen levels
were lowered to 41 and 350 mmol/kg d.w., respec- Fig. 1. Venous blood lactate (a) and plasma K1 concentra-
tively. In another experiment Bogdanis et al. (1996a) tions (b) during a speed endurance training session.
examined the metabolic response when repeating
two 30-s maximal cycling bouts separated by 4 min
of rest. Muscle pH decreased to 6.69 immediately
after the rst bout and increased only slightly (6.80) throughout recovery peaking at 170 mmol/L 3.5 min
prior to the second sprint, after which it dropped to after sprint 2 (Bogdanis et al., 1995; Bogdanis et al.,
6.61. Consistently, muscle [Lac  ] rose to 108 and 1996b). In addition, the calculated ATP turnover was
129 mmol/kg d.w. at the end of the rst and second markedly dierent when repeating 2 30-s maximal
bout, respectively. The muscle PCr levels at the sprints. PCr, energy release from glycolysis and
end of the rst and second bout were similar (12.6 aerobic metabolism accounted for 21, 50 and 29%,
and 8.8 mmol/kg d.w., respectively) and the PCr was respectively, of the total energy production during
resynthesized to 58.5 mmol/kg d.w. (79% of the the rst sprint, and changed to 20, 36 and 44% in the
resting value) before the second exercise bout. The second bout (Nevill et al., 1994). Consistently, when
muscle glycogen content decreased from 327 to 228 performing 4 repeated 30-s maximal cycling exercises
and 184 mmol/kg d.w. at the end of the rst and separated by 4 min of rest, Parolin et al. (1999)
the second bout, respectively. Plasma NH3 concen- observed a decreased substrate phosphorylation
tration was elevated from 29 to 94 mmol/L immedi- by phosphocreatine hydrolysis and glycolysis, and
ately after the rst 30-s sprint, and increased a concomitant shift toward greater reliance on

12
Speed endurance training in trained subjects
oxidative phosphorylation. These data taken to- 1000 m cycling, 100200 m swimming as well as team
gether with those by Bangsbo et al. (2001), Medbo sports.
& Tabata (1989) and Tabata et al. (1997) indicate Signicant increases in performance are also ob-
that speed endurance training taxes both the anae- served in events lasting 46 min with the studies
robic and aerobic energy pathways to a signicant involving shorter recovery times between the inter-
extent, with the rate of glycolysis progressively de- vals (e.g. 30-s exercise separated by 30-s rest periods)
creasing and the aerobic energy production increas- (Paton & Hopkins, 2005; Hamilton et al., 2006)
ing as the exercise is repeated. showing less pronounced improvements as compared
with those using longer resting periods (Shepley
et al., 1992; Esfarjani & Laursen, 2007). However,
Effect of speed endurance training on physical at present there is insucient data and research
performance in trained human subjects comparing speed endurance production vs. mainte-
nance training to provide clear evidence based re-
Short-term and repeated intense exercise performance commendations about the most appropriate exercise:
There is a general consensus that a period of speed rest ratio. The recommendations of the type of
endurance training in endurance-trained subjects protocol (intensity, exercise duration and rest) are
leads to marked performance improvements during dependent also upon the type of performance im-
supramaximal and repeated high-intensity exercise provements and adaptations an athlete is looking for.
lasting less than  10 min (Tables 2 and 3). Houston In general it appears that production training
& Thomson (1977) observed that after a 6-week (exercise rest ratio of  1:6) is benecial for improv-
program involving anaerobic hill running, ve en- ing the ability to perform maximal eorts repeatedly,
durance-trained men increased the mean distance whereas maintenance training (1:13) may be
covered in 2 maximal runs of 60 s and 90 s each by eective in increasing the ability to sustain exercise
 13%. Similarly, Daniels et al. (1978) showed a at high intensity.
 10% lower time over  800 m ( 130 s) when The eect of speed endurance training has been
trained runners performed part of their training as investigated also in football (soccer). In a series of
100600 m sprints. More recently, others reported studies, elite football players underwent 210 weeks
improved time to exhaustion (11.2%) (Bickham of intensied training including 13 weekly sessions
et al., 2006), predicted speed (4.4%) (Hamilton of various speed endurance training (Table 2) and
et al., 2006) and mean power (8.7%) (Paton & improved their repeated sprint ability (1.92.1%)
Hopkins, 2005) during various supramaximal exer- (Ferrari Bravo et al., 2008; Thomassen et al., 2010),
cises lasting 23 min. These changes are, however, maximal aerobic speed (8.1%) (Dupont et al., 2004)
lower than those found in other investigations (Table and Yo-Yo intermittent recovery (Yo-Yo IR) test
3). Part of the discrepancies observed could be due to (Bangsbo et al., 2008) performance (2228%) (Fer-
the dierent testing protocols (i.e. time trial vs time rari Bravo et al., 2008; Hill-Haas et al., 2009). The
to exhaustion) and measurements (i.e. time, distance, latter is in agreement with Iaia et al. (2008) who
speed, power) utilized. Nevertheless, it appears from observed a 19% increase in Yo-Yo IR level 2
Table 2 that the three studies showing the greatest performance when for a 4-week period trained run-
increases in performance used speed endurance pro- ners replaced their endurance training with 34
duction training involving 30-s near maximal inten- weekly sessions of speed endurance production train-
sity bouts with relatively long rest periods (41:4) ing. The potency and time-eciency of very high-
(Roberts et al., 1982; Iaia et al., 2008; Bangsbo intensity training is also evident from a recent study
et al., 2009). In addition, two out of the three had by Thomassen et al. (2010). Trained football players
also a concomitant signicant reduction in the improved their performance during a repeated sprint
amount of training. Thus, speed endurance produc- test, in association with an increased expression of
tion training with 30-s all-out exercise and  3 min some muscle proteins involved in ion homeostasis,
rest is a very potent stimulus and a time ecient after a 2-week reduction in training volume incorpor-
strategy for markedly improving performance during ating 5 aerobic high-intensity and 5 speed endurance
very intense short-term exercise, i.e. from 30 s to training sessions. Thus, speed endurance training is
3 min duration. Furthermore, these pronounced per- benecial for people participating in team sports
formance improvements were obtained under condi- involving intense activities by elevating their ability
tions of reduced weekly mileage, clearly indicating to recover faster after the intense phases of a match
that high training volumes may not be needed, and in and thereby potentially increasing the number of
some cases may even be detrimental for improve- high-intensity periods during a game. Less clear is
ments during supramaximal exercise performances. the eect of speed endurance training on sprint and
This information is relevant for athletes participating jump performance with studies reporting either im-
in sports such as 400, 800 and 1500 m track events, proved or unchanged performances (Table 3).

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Table 2. Effect of speed endurance training on physiological adaptation and performance in trained human subjects

14
Study Fitness status n Protocol Intensity Duration Changes in Performance Adaptations Physiological response to
and exercise mode (week) training volume changes exercise
.
Bangsbo Well-trained
. runners 12 812  30 s3 min rest, 9095% speed 69 #  30% (from  50 to "9%  8-min incremental "2.7% (NS) VO2 max $HR during 3 and 10 k
et al. (2009) (VO2 max 63 mL/kg/min 23  wk achieved over 30-s  35 km/wk) exhaustive test "68% Na1,K1 pump a2 "3% running economy at
3 km time 10.4 min run at full maximal " 6% distance 30-s "10% (NS) Na1,K1 12 km/h
Iaia & Bangsbo

10 km time 37.3 min) effort max test pump b1 #RER at 17 km/h1


"36% time to exhaustion $Na1,K1 pump a1 "peak blood [La  ] after
.
130% VO2 max ( 2 min) "24% (NS) NKCC1 incremental test
"6% (NS) time to exhaustion $NHE1, MCT1, MCT4 #[K1]v after EX1 and EX2
.
130% VO2 max ( 70 s, EX2) $CK,PFK, CS, HAD
"3.3% 3 km performance
"3.1% 10 km performance
.
Bickham Moderately endurance- 7 4  (48  40100 m 90100% maximal 6 Unchanged ( 50 km/wk "11% time to exhaustion at
.
#2% (NS) VO2 max
et al. (2006) trained
. runners sprints jog 35 fold ex aerobic moderate intensity) 110% VO2 max (2.3 min) " 45% MCT1
(VO2 max  58 mL kg/ time) 5-min jog between $MCT4
min) sets, 3  wk #11% ST fibers
"14% FT fibers
$body mass
.
Daniels et al. Trained
. runners 15 5 subjects run half of their Not specified 8 " From 2030 to " 10%  800 m $ VO2 max
(1978) (VO2 max  63 mL kg/ training as 100600 m sprint 5070 km/wk performance
min, 800 m time runs " 15%  3 km
 2 min) performance

Dupont Professional football 22  1215  (40 m30 s rest), Maximal 10 Not specifically reported but "8.1% max aerobic speed $body fat
et al. (2004) players running 1  wk. apparently maintained #3.5% 40 m sprint time $body mass
 2  (1215  15 s15 s 120% max aerobic ("50% games won) $HR
rest), 1  wk speed
. .
Esfarjani & Moderately trained 6 12  30 s4.5 min jog, 130% VO2 max 10 Unchanged ( 36 km/wk), "7.3% 3000 m time trial
.
"6.2% VO2 max "4.7% (NS) speed at
Laursen (2007) runners
. 2  wk 2  60-min "7.8% speed at VO2 max lactate threshold
(VO2 max  52 mL/kg/ treadmill. run "32% time
.
min, 3000 m time at 75% VO2 max to exhaustion at VO2 max
 11 min) ( 6 min)
.
Ferrari Sub-elite football
. players 13 3  (6  20120 m Maximal 7 Maintained "28.1% Yo-Yo IR1 "5% VO2 max
Bravo et al., running (VO2 max sprint 20 s rest) "2.1% repeated sprint ability "3% VT2
(2008)  56 mL/kg/min) 3 min rest, 2  wk #0.6% (NS) 10 m sprint time
$squat jump height, power
$CMJ height, power
.
Hamilton Trained runners (VO2 max 10 3  (20 explosive 1-leg Maximal 7 Pre training  30 km "4.4% predicted 800 m speed $ body mass "4.0% speed
et al. (2006) 66 mL/kg/min, 5 km time jumps)13  (5  30 s 65% peak running #  17% (3.63 h/wk) "4.1% predicted 1500 m $thigh cross-sectional at 4 mM [La  ]
o20 min) 30 s rest), 13  wk speed speed area
"2.7% peak incremental
speed
$2.2% 5 km time trial
.
Hill-Haas Junior elite
. football 9  18 to 20  3060 s 9095% HRmax 7 Maintained "22% Yo-Yo IR1 "2% (NS) VO2 max
et al. (2009) players (VO2 max 6090 s rest, 1  wk $repeated sprint ability
 60 mL/kg/min)  7  34 m35 s Maximal "2.5% (NS)  10-min
rest1 210  590 s incremental exhaustive test
1590 s rest, 1  wk OR "3.1% (NS) multistage fitness
 3035  1020 m, Maximal test
1040 s rest, 1  wk #0.9% (NS) 5 m sprint time
#1.5% (NS) 20 m sprint time
.
Houston and Endurance-trained
. 5 3  (60 s2 min jog) 3.3% Maximal 6 Similar ( 35 km/wk). $Stair-sprinting power test "3% (NS) VO2 max "14% peak blood [La  ]
Thomson (1977) runners (VO2 max (inclination) No aerobic exercise except "16.7% time to exhaustion "3.3% (NS) FT fiber
 59 mL/kg/min) 15  (6 s24 s rest) 44%  3 km warm-up steep (20%) #4% (NS) LDH
12  (90 s3 min jog) 3.3% treadmill run ( 50-s) #14.8% ATP at rest
115 leg press max rep., "13.7% distance 60-s full "4.5% (NS) PCr at rest
4  wk maximal effort (3.3% #13.7% cholesterol
grade) $body fat
"13.4% distance 90-s full $body mass
maximal effort (3.3%
grade)
.
Iaia et al. (2008) Moderately endurance-
. 8 812  30 s3 min rest, 9095% speed 4 # 64% "5%  7-min incremental #2% (NS) VO2 max #[K1]v in and after EX1
trained runners (VO2 max 34  wk achieved over 30-s (from  43 to  15 km/wk) exhaustive test "29% Na1,K1 pump a1 #glycogen degr. in EX2
 55 mL/kg/min, 10 km run at full-maximal (aerobic exercise only for "7% distance 30-s max test "17% (NS) Na1,K1 "H1 removal after EX1
time  40 min) effort warm-up and cool down "26% time to exhaustion
. pump a2 $PCr, lact glycogen,
activities) 130% VO2 max ( 100 s, EX1) $Na1,K1pump b1 [H1] at rest and
"19% time to exhaustion "30% NHE1 during exercise
.
130% VO2 max ( 75 s, EX2) "14% (NS) NKCC1
"19% Yo-Yo IR2 $b in vitro, MCT1, MCT4,
$CK, PFK
$body mass
.
Iaia et al. (2009) Moderately endurance-
. 9 812  30 s3 min rest, 9095% speed 4 #  65% (from  43 $10 km time trial #2% (NS) VO2 max "67% running economy
trained runners (VO2 max 34  wk achieved over 30-s to  15 km/wk) $CS, HAD #energy expenditure,
 55 mL/kg/min, 10 km run at full maximal (aerobic exercise only for "41% (NS) UCP3 HR and ventilation
time  40 min) effort warm-up and cool down #7% (NS) ST fibers during submaximal
activities) #6% (NS) FTa fibers running
"44% FTx fibers $blood [La  ] during
"7% (NS) No. capillaries submaximal running
.
Laursen et al. Highly
. trained cyclists 10 12  30 s4.5 min rec., 175% PP 4 Maintained regular low-intensity "3% PP "3% VO2 max
(2002) (VO2 max  64 mL/kg/ 2  wk training "4.4% 40 km time trial
min) $time to exhaustion at PP
( 4 min)
.
Laursen et al. Highly
. trained cyclists 10 12  30 s4.5 min rec., 175% PP 4 Maintained regular low-intensity "4% (NS) VO2 max "VO2-peak and HR during
(2005) (VO2 max  64 mL/kg/ 2  wk training $plasma protein 40 km time trial
min) content $RER and blood [La  ]
$hematocrit during 40km time trial
$hemoglobin
$plasma volume
"17% VT1
"9% VT2
"75% anaerobic capacity

Paton and Well-trained cyclists 9 3  (20 single-leg Maximal 45 Replaced 20% of their total "6.7% PP $body mass #3.2% energy
Hopkins (2005) jumps)13  (5  30 s weekly volume "8.7% 1 km mean power "8.4% expenditure
30 s rest), 2 min rest, 4 km mean power
23  wk

Roberts et al. Regularly active running 4 8  200 m 2 min rest, 90% of max speed 5 Not reported "25% 40-s exhaustive run "  110% PFK
(1982) 34  wk over 200 m (16 km/h, 15% grade) "  50% Phos
"  50% GAPDH
"  50% LDH
"  15% MDH
"17% (NS) SDH
Speed endurance training in trained subjects

15
Table 2. (continued)

16
Study Fitness status n Protocol Intensity Duration Changes in Performance Adaptations Physiological response to
and exercise mode (week) training volume changes exercise

. .
Shepley Highly trained
. distance 9 15  (500 m 67 min 115120% VO2 max 1 From 80 to 7.5 km/wk "22% treadmill run to $VO2 max $post-exercise plasma
et al. (1992) runners (VO2 max walk), 5  wk (warm up excluded) exhaustion
. at
"18% CS [La  ]
Iaia & Bangsbo

 67 mL/kg/min)  115% VO2 max ( 4 min) "15% rest glycogen


"13% (NS) peak torque "  5% blood volume
"  16% red cell volume
$hematocrit

Stepto Endurance-trained
. 4 12  30 s4.5 min rec., 175% PP 3 Not reported " 0.5%(NS) PP $HR, carbohydrate
et al. (1999b) cyclists (VO2 max 2  wk " 3.8% (NS) oxidation and plasma
 60 mL/kg/min) 25 kJ sprint power [La  ]
" 2.3% 40 km
( 55 min) time trial
. .
Tabata et al. Physical education 7 78  (20 s10 s rest), 170% VO2 max 6 Not reported " 15% VO2 max
(1996) students from various
. 5  wk
sport cycling (VO2 max
 48 mL/kg/min)

Thomassen Elite
. football players 7  8  2 min 1 min rest,  90% HRmax 2 #  30% "1.9% repeated sprint ability "14.5% Na1,K1 pump a2
et al. (2010) (VO2 max  55 mL/kg/ 23  wk "6.1% (NS) Yo-Yo IR1 "27.3% resting
min)  1012  2530 s 9095% speed $20 m sprint time phosphorylation status
2.53 min rest, 2  wk achieved over 30-s FXYD1
run at full maximal "18% (NS) Na1,K1
effort pump a1
$Na1,K1 pump b1
"13% (NS) MCT1,
MCT4
$NHE1, NKCC1

Thorstensson Moderately
. trained 4 2040  (5 s sprint Maximal 8 Not reported #1% (NS) 25 m sprint time "36% CK
et al. (1975) running (VO2 max 2555 s rest), "12% MVC "20% MK
 54 mL/kg/min, 10 km 34  wk "9% Jump "30% ATPase
time  40 min) "4% (NS) Margaria test $resting ATP,
. CP
"19% endurance time "3% (NS) VO2 max
at 50% #5% (NS) FTb fibers
MVC ( 50 s) "2% body mass
"3% thigh girth

ATPase, adenosine triphosphatase; CMJ, counter movement jump; CK, creatine kinase; CS, citrate synthase; FXYD1, Na1,K1 pump accessory and regulatory protein phospholemman; FT, fast twitch; GAPDH,
glyceraldehyde-3-phosphate dehydrogenase; HAD, 3-hydroxyacyl-CoA dehydrogenase; HR, heart rate; LDH, lactate dehydrogenase; MCT, monocarboxylate transporters; MDH, malate dehydrogenase; MK, myokinase;
MVC, maximal voluntary contraction; NHE1, Na1/H1 exchanger isoform 1; NKCC1, Na1-K12Cl  1 protein co-transporters; PCr, creatine phosphate; PFK, phosphofructokinase; Phos, glycogen phosphorylase; PP,
.
peak aerobic power; RER, respiratory equivalent ratio; SDH, succinate dehydrogenase; ST, slow twitch; UCP3, uncoupling protein 3; VO2 max , maximum oxygen uptake; VT, ventilatory threshold; Yo-Yo IR, Yo-Yo
intermittent recovery test; ", increased; #, decreased; $, unchanged; NS, non-significant; wk, week.
Speed endurance training in trained subjects
Medium- to long-term exercise performance 1996; Laursen et al., 2002; Ferrari Bravo et al., 2008).
In the study by Iaia et al. (2009) when already trained However, in two studies the participants were not
runners switched from regular endurance to speed highly trained (Tabata et al., 1996) and the total
endurance training, the 10 K performance was un- weekly volume was maintained (Laursen et al., 2002).
altered despite a 65% reduction in training distance. Furthermore, in some investigations brief rest peri-
Apparently, speed endurance training per se is su- ods were used between repetitions, which may have
cient to maintain endurance performance of already allowed the cardiovascular stimulus to be maintained
trained subjects. This is in accordance with Houmard at a high level for longer durations (Tabata et al.,
et al. (1990) who did not nd any changes in a 5 K 1996; Ferrari Bravo
. et al., 2008). Nevertheless, the
race after a 3-week period in which well-conditioned importance of VO2 max at an elite level
. is questionable
runners maintained the intensity while reducing their as often no relationship between VO2 max and perfor-
weekly training volume by 70% and frequency by mance are observed in athletes (Svedenhag & Sjodin,
17%. Bangsbo et al. (2009) even reported an im- 1985; Coyle, 1995) and in the majority of the studies
proved performance in 3 and 10 K runs after a 69- the improved short- and long-term. exercise perfor-
week period with speed endurance sessions where the mances were not associated with VO2 max enhance-
total amount of training was reduced by 30%. In ments (Table 2).
addition, other studies on trained cyclists observed a Speed endurance training has been shown to reduce
lower time (45%) to complete a simulated-labora- energy expenditure during exercise (Table 3), as is the
tory 40 K cycling trial (Stepto et al., 1999a; Laursen case after plyometric and sprint training (Paavolai-
et al., 2002), clearly indicating that when combining a nen et al., 1999; Turner et al., 2003; Saunders et al.,
basic volume of training including some aerobic 2006). In addition, improved performances were
high-intensity exercise, together with only 2 weekly associated with a reduced mean net rate of glycogen
sessions of speed endurance training, trained subjects degradation during supramaximal exercise (Iaia et
can improve endurance performance. It is interesting al., 2009) and an elevated fat oxidation during
to note that in the study by Laursen et al. (2002) the intense submaximal exercise (Bangsbo et al., 2009).
endurance performance was improved after 4 but not Thus, in trained individuals speed endurance training
after 2 weeks of speed endurance training, which is in appears to be a potent stimulus to decrease the net
contrast with the nding of untrained subjects where rate of muscle glycogenolysis during submaximal
pronounced changes were found already after 2 exercise as also observed for untrained subjects
weeks (Burgomaster et al., 2005). Apparently, (Harmer et al., 2000; Burgomaster et al., 2006).
trained individuals adapt more slowly compared
with untrained people. Nonetheless, the improve-
ments observed in endurance events are of a lower Muscle adaptations to speed endurance training in
magnitude than those reported during intense short- trained human subjects
duration exercises (Table 3). The ndings in the
While the alterations in performance occurring
abovementioned studies taken together with those
after a period of speed endurance training in
by Daniels et al. (1978), who observed greater 3 K
trained subjects have been extensively investigated,
improvements when combining speed endurance
only a few studies have focused on examining the
training with augmented training volume (Table 3),
changes in muscle variables with this type of training
suggest that for maximizing performance improve-
(Table 2).
ments, distance athletes should also regularly carry
out other types of training, e.g. aerobic high-inten-
sity, besides speed endurance training. Fiber type proportion
To what extent muscle bers can shift from slow
twitch (ST) to fast twitch (FT) bers in humans and
Effect of speed endurance training on VO2 max and
energy expenditure in trained human subjects vice versa is still a matter of controversy. Never-
theless, the proportion of ST muscle bers has been
Most of the studies on well-trained individuals eval- shown to decrease and the relative number of FT has
uating the eect of speed endurance production been demonstrated to increase or remain unaltered
training with an intensity higher than. 90% of the with anaerobic training (Table 3). In addition, a
maximum have reported unchanged VO2 max values. signicant increase in FTx bers was observed
This suggests that short maximal or. near maximal when speed endurance training was accompanied
eorts are not sucient to enhance VO2 max . On the by a severe reduction in training volume (Iaia et al.,
other hand, studies dealing with speed endurance 2009). Thus, the few studies being performed do not
maintenance training
. where the average intensity was lead to a rm conclusion, but they appear to suggest
lower, showed VO2 max improvements (Tabata et al., that speed endurance training of endurance trained

17
Iaia & Bangsbo
Table 3. Summary of the effects (expressed as percentage change) of speed endurance training on physiological adaptation and performance in trained
human subjects

Significantly increased Maintained Significantly


(% change) (% change) decreased (% change)

Anthropometric characteristics
Body mass 2 0.1, 1,  1.6, 0.3, 0.2, 0.2
Body fat  6,  0.7
Thigh girth 3 1.4
Exercise performances
30 s 7, 7
4060 s 25, 16.7, 13.7, 19
1.52.5 min 27, 11, 13.4,  10, 4.4, 8.7, 36
z
46 min 22, 32, 4.1, 8.4
820 min 9*, 3.3,  15, 8.1*, 7.3, 2.7*, 2.2, 2.5*, 3.1*,  0.5*
5*, 3*, 6.7*
3560 min 4.4,  2.3, 3.1 0.2
Repeated intense exercise to exhaustion 19, 28.1, 22 6.1
Repeated sprint ability 2.1, 1.9 0.2
Sprint time  0.6,  0.9,  1.5, 0,  1  3.5
Jump test 9 0
Pulmonary
. variables
VO2 max 6.2, 3, 5, 15w 3,  0,  2,  2, 3, 0.1,
4, 2.7, 2
Energy expenditure  7,  3.2,  3
Blood variables
Blood/plasma volume 5 1.7
Cholesterol  13.7
Hematocrit 3,  0.2
Muscle substrates
ATP 6  14.8
PCr 4.5,  12,  14
Glycogen 15 3
Muscle anaerobic enzymes
ATPase 30
CK 36 0.3, 10.4
GAPDH 50w
LDH 50w 4
MK 20
PFK 110w 1, 10.5
Phos 50w
Muscle oxidative enzymes
CS 18  5,  3.5
HAD 2,  11
w
MDH 15
SDH 17w
Muscle membrane transport protein
MCT1 45  3,  2, 13.3
MCT4 3,  8,  10,  13
Na1,K1 pump a1 29   10,  18
Na1,K1 pump a2 68, 14.5 17
Na1,K1 pump b1 1, 10,  4
NHE1 30 1,   4
NKCC1 14, 24, 0
Muscle histochemical variables
Capillarization 7
Fiber type I 7  11
Fiber type IIa 14 (FT)  6, 3.3 (FT)
Fiber type IIx 44 5
Muscle buffering capacity 3

*Incremental test trial.


w
Physically regularly active.
z
Percentage change not reported.
Data taken from studies reported in Table 2.
ATPase, adenosine triphosphatase; CK, creatine kinase; CS, citrate synthase; Fr, Fast Twitch; GAPDH, glyceraldehyde-3-phosphate dehydrogenase; HAD,
3-hydroxyacyl-CoA dehydrogenase; LDH, lactate dehydrogenase; MCT, monocarboxylate transporters; MDH, malate dehydrogenase; MK, myokinase;
NHE1, Na1/H1 exchanger isoform 1; NKCC1, Na1-K12Cl  1 protein co-transporters; PCr, creatine phosphate; PFK, phosphofructokinase; Phos,
.
glycogen phosphorylase; SDH, succinate dehydrogenase; VO2 max , maximum oxygen uptake.

18
Speed endurance training in trained subjects
.
subjects leads to a higher relative number of FT eliciting  130% VO2 max , suggesting that a high rate
bers. of PCr breakdown is an important stimulus for CK
adaptation (Mohr et al., 2007). In agreement, a
considerable number of studies on untrained people
Muscle metabolites, enzymes and capillarization have shown improvements in anaerobic enzymes
One study has shown a reduced amount of muscle activity after periods of training including very short
resting ATP concentration with speed endurance maximal/near maximal exercise bouts (530 s) inter-
training in trained subjects (Houston & Thomson, spersed with relatively long resting periods (45 s to
1977) and another an unaltered level (Thorstensson 20 min) (Costill et al., 1979; Jacobs et al., 1987;
et al., 1975). The dierence may be related to the Linossier et al., 1993; Hellsten-Westing et al., 1993;
extensive speed endurance training protocol used in Linossier et al., 1997; MacDougall et al., 1998; Parra
the former study which required subjects to run to et al., 2000; Rodas et al., 2000). On the other hand, in
exhaustion. This is in contrast with Thorstensson a study (Barnett et al., 2004) utilizing a training
et al. (1975) who employed shorter exercise bouts and protocol (30-s sprints with 34 min of rest) similar
longer recovery periods. In accordance, sprint train- to the one used by Iaia et al. (2008) and Bangsbo et
ing of untrained people has been shown to decrease al. (2009), the activity of phosphofructokinase (PFK)
the resting levels of skeletal muscle adenine nucleo- remained unaltered. Thus, the higher tness level
tides, if the training was frequent and very intense together with longer exercise periods (i.e. 30s) may
(Hellsten-Westing et al., 1993). However, it does not explain why no changes in the activity of CK and
appear to have a major negative impact on work PFK were observed in the latter two studies (Iaia
capacity as all the studies have shown increases in et al., 2008; Bangsbo et al., 2009). Nevertheless,
exercise performance. The muscle concentration of pronounced alterations in short-term performance
PCr does not appear to be aected by speed endur- were observed despite unaltered levels of enzymes
ance training. One study has shown increased levels related to anaerobic energy production (Iaia et al.,
of muscle glycogen, whereas in another investigation 2008; Bangsbo et al., 2009), suggesting that the
they were maintained (Table 3). Importantly, it is not changes in the activity of these enzymes are not
easy to examine the eect of training on muscle crucial for work capacity improvements during
glycogen in athletes as the control level may be exercise lasting 30 s to 2 min.
inuenced by the frequent training sessions. Never- The activity of oxidative enzymes in trained sub-
theless, it is doubtful whether a change in muscle jects does not appear to be elevated with speed
glycogen plays a role in the performance improve- endurance training (Table 3), which is clearly in
ment during a single high-intensity exercise bout as contrast to a number of studies of untrained subjects
muscle glycogen does not appear to be a limiting (Rodas et al., 2000; Parra et al., 2000; Burgomaster et
factor in short-term intense exercise (Bangsbo et al., al., 2005; Burgomaster et al., 2006; Gibala et al.,
1992a). On the other hand, elevated glycogen levels 2006; Burgomaster et al., 2008). This dierence may
in the working bers are benecial in sports with be related to the lower level of enzyme activity in
repeated intense exercise such as soccer and rugby untrained as compared with already trained indivi-
(Bangsbo et al., 1992b; Balsom et al., 1999). duals and to the fact that untrained subjects do
The eect of speed endurance training on the respond to almost all types of training stimuli by
activity of enzymes related to the anaerobic metabo- increasing a large number of muscle proteins. It is
lism of trained individuals is not clear. Roberts et al. interesting to note, however, that in a study with a
(1982) found that the maximal activity of glycolytic 65% reduction in training volume, the levels of
enzymes and creatine kinase (CK) was increased by citrate synthase (CS) and 3-hydroxyacyl-CoA dehy-
speed endurance training (Table 3). However, the drogenase (HAD) were maintained with regular
participants in this study were regularly active but speed endurance training, indicating that this type
not well trained and studies enrolling trained sub- of training also in trained humans stimulates the
jects have not been able to show changes (Iaia et al., mitochondrial oxidative proteins (Iaia et al., 2009).
2008; Bangsbo et al., 2009). It is traditionally be- In accordance, Shepley et al. (1992), investigating
lieved that in order to enhance the activity of meta- highly trained runners during a week of reduced
bolic enzymes it is fundamental to utilize an exercise training combined with speed endurance training,
mode that produces high ux through the relevant found an 18% increase in the CS level in association
pathways in the contracting muscles. It was recently with improved performance (Table 2). Thus, it ap-
reported that CK activity was higher when a group pears that speed endurance training is a powerful
of untrained subjects performed a sprint-training stimulus to mitochondrial protein synthesis and
program consisting of 6-s maximal runs separated whether it leads to net synthesis depends on the
by 1 min of recovery, whereas it remained unchanged training status as well as on the frequency and
for another group performing 30-s runs at a speed amount of training.

19
Iaia & Bangsbo
There is limited information about the eect of have already had an elevated protein content before
speed endurance training on muscle capillarization. the training, as endurance training has been demon-
Iaia et al. (2009) observed a non-signicant 7% strated to increase the Na1,K1 pump subunits
increase in the number of capillaries per ber despite (Green et al., 2004). The augmented Na1,K1 pump
a 65% reduction in the total amount of training. The a subunits may result in a high number of functional
lack of signicance may have been due to the fact pumps and play an important role for the increases in
that the two best trained subjects showed a decrease work capacity during intense short-term perfor-
in the number of capillaries. These ndings may mance. Indeed, a higher Na1,K1 pump maximum
suggest that speed endurance training is a stimulus activity reduces the net loss of K1 from the contract-
for muscle angiogenesis in trained subjects. However, ing muscles preserving the cell excitability and force
a basic volume of training may also be necessary in production (Sejersted & Sjogaard, 2000). This is
order to increase the capillarization. Further studies supported by the nding that in both studies the
are needed to examine this hypothesis. elevated expression of the Na1,K1 pump a subunits
observed after the speed endurance training period
was associated with a decreased accumulation of
Muscle ion transport proteins venous plasma K1 and an increased performance
Among the wide range of potential candidates in- during repeated supramaximal exercise.
volved in the fatigue processes, sarcolemmal depo- In trained people, isoforms of the muscle mono-
larization due to extracellular K1 accumulation has carboxylate transporters (MCT) have been shown to
been suggested to be of primary importance for be unchanged with speed endurance training (Iaia
fatigue development during high-intensity exercise et al., 2008; Bangsbo et al., 2009; Thomassen et al.,
(Sejersted & Sjogaard, 2000). This hypothesis is 2010), which is in contrast to data from untrained
based on observations that during dynamic exercise subjects (Pilegaard et al., 1999; Juel et al., 2004;
the contracting muscles lose K1 which progressively Bickham et al., 2006; Burgomaster et al., 2007;
accumulates in the extracellular space leading to a Mohr et al., 2007). One study has reported sprint
depolarization of the membrane potential (Em) (Juel, training-induced changes on MCT1 proteins in en-
1988; Cairns et al., 1995; Cairns et al., 1997). A lower durance-trained subjects (Bickham et al., 2006). In
Em has been shown in in vitro studies (Sjogaard et al., contrast to studies reporting no changes, in the latter
1985; Juel, 1988; Cairns et al., 1997; Sejersted & investigation the subjects maintained a high volume
Sjogaard, 2000) to cause membrane inexcitability of training (  50 vs  15 and  32 km/week,
and simultaneous tetanic force reduction probably respectively), which may be part of the explanation
via a slow inactivation of the voltage-gated Na1 for the dierences observed.
channels (Ru et al., 1988; Ru, 1999). The evidence In the study by Iaia et al. (2008) the in vitro
of K1 being of importance for fatigue development measured muscle buering capacity was not chan-
in humans is related to the ndings that muscle ged. Consistently, muscle and blood lactate concen-
interstitial K1 during intense exercise is elevated to trations as well as muscle H1 accumulation were
levels 410 mM (Nordsborg et al., 2003; Nielsen et found to be unaltered during repeated intense ex-
al., 2004), and that exercise training leads to reduced haustive exercise (Iaia et al., 2008). Apparently,
interstitial K1 accumulation (Nielsen et al., 2004) in improved short-term performance can occur without
association with improved exercise performance and changes in MCT and buering capacity. In accor-
a change in the amount of Na1,K1 pumps (Clausen, dance, Messonnier et al. (2007) observed that sub-
2003). jects with elevated performance levels compared with
Speed endurance training of trained subjects ele- untrained individuals are less dependent on the
vates the expression of either the Na1,K1 pump a1 muscle content of proteins involved in buering
or a2 subunits (Iaia et al., 2008; Bangsbo et al., 2009; mechanisms and lactate/H1 transport. As such, it
Thomassen et al., 2010) as well as its accessory and appears therefore reasonable to assume that in al-
regulatory protein phospholemman (FXYD1) (Tho- ready trained individuals mechanisms other than
massen et al., 2010), whereas no signicant increases changes of pH regulatory systems may be predomi-
have been observed in the b1 subunit (Iaia et al., nant in delaying fatigue development and enhancing
2008; Bangsbo et al., 2009; Thomassen et al., 2010). work capacity during intense exercise. On the other
In the rst two studies, the changes reported in the a1 hand, the expression of the Na1/H1 exchanger iso-
and a2 subunits were marked (29% and 68%, form 1 (NHE1) was enhanced after a speed endur-
respectively) and are comparable to those observed ance training period (Iaia et al., 2008). This could
in untrained and diabetic subjects after a period of have facilitated the Na1 uptake inside the muscle cell
speed endurance and strength training (Dela et al., and thus possibly resulted in a higher stimulation of
2004; Nielsen et al., 2004). These adaptations oc- the Na1,K1 pump (Ewart & Klip, 1995) causing a
curred despite the fact that the trained subjects may hyperpolarization of the Em.

20
Speed endurance training in trained subjects
Conclusion Practical implications
In many sports the time available for training is For a practical purpose, speed endurance training
limited during the season. Coaches and athletes are may be divided into two dierent forms: production
usually concerned that a reduced amount of training, and maintenance. In production training the ex-
i.e. number and duration of sessions, may be detri- ercise intensity is almost maximal and there is re-
mental for performance and sometimes athletes train search evidence that this is a very eective way to
more than required. The studies presented in this elicit adaptations in several muscle variables as well
review provide strong evidence that, although its as to improve performance during very intense short-
brevity, speed endurance training leads to perfor- duration and repeated high-intensity exercises. In
mance improvements during several high-intensity order to maintain a high-intensity throughout the
short-duration exercises in already trained subjects. session, the duration of exercise periods should be
Furthermore, despite marked reduction in training relatively short but suciently longer (410 s) to
volume, the muscle oxidative potential, capillariza- stimulate the systems which limit performance. To
tion and aerobic performance were shown to be perform maximally in a subsequent exercise bout, the
unchanged indicating that exercise at near maximal resting periods should therefore be long enough for
intensity is a powerful stimulus for maintaining muscle to approximate to their pre-exercise state.
physiological adaptation and performance gained Thus, exercise periods lasting 1040 s at near max-
from previous endurance training. On the other imal speed separated by rest periods of 15 min may
hand, when combined with a basic amount of aerobic be optimal for production training.
training, speed endurance training has even been In speed endurance maintenance training the re-
reported to improve endurance performance. These covery times between the intervals are shorter (one to
changes
. do not appear to be mediated by elevated threefolds the exercise time) and the duration of the
VO2 max , higher glycolytic and oxidative enzymes exercise periods may be longer (590 s) compared
activity, and improved pH regulation. It may instead with speed endurance production training, resulting
be related to a lower energy expenditure during in an overall lower exercise intensity. This exercise
exercise and changes in the Na1,K1 pump a1 or a2 mode leads to a gradual accumulation of fatigue as
subunits, which via a reduced contraction-induced the training progresses and it has been shown to be
net loss of K1 from the working muscles may useful to improve the ability to sustain intense
contribute to preserve cell excitability and maintain exercise.
force development. However, there may be a number
of other factors contributing to the elevated perfor- Key words: high-intensity intermittent, enzymes, pH,
mance. It should be emphasized that none of the membrane transport proteins, Na1,K1 pump.
studies presented have been performed on elite ath-
letes. Regardless, it appears reasonable to suggest
that athletes in sports involving intense exercise may Acknowledgements
benet from periodically reducing the amount of
workload and undertaking speed endurance training The studies presented by the authors were supported by the
on a regular basis. Nevertheless, further studies are Danish Ministry of Culture (Kulturministeriets Udvalg for
Idrtsforskning), the Danish Natural Science Research Coun-
needed to examine the eect of reducing the amount cil (272-05-0407), and Team Denmark.
of training and performing speed endurance training Conicts of interest: The authors have no potential conicts
over a longer period (several months). of interest.

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