Effect of Exercise Intensity Duration No Epoc

Sports Med 2003; 33 (14): 1037-1060REVIEW ARTICLE 0112-1642/03/0014-1037/$30.00/0 Adis Data Information BV 2003. All rights reserved.

Effect of Exercise Intensity, Durationand Mode on Post-ExerciseOxygen ConsumptionElisabet Brsheim and Roald Bahr

Norwegian University of Sport and Physical Education, Oslo, Norway

ContentsAbstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1037

1. Excess Post-Exercise Oxygen Consumption (EPOC) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10382. Early Studies on EPOC . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10393. Methodological Considerations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10394. Effect of Intensity and Duration of Aerobic Exercise on EPOC . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10405. Effect of Split Exercise Sessions on EPOC . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10476. Effect of Supramaximal Exercise on EPOC . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10487. Effect of Aerobic Exercise Mode on EPOC. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10488. Effect of Resistance Exercise on EPOC . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10499. Effect of Training Status on EPOC . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1051

10. Effect of Sex on EPOC . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105211. Possible Mechanisms for the Rapid EPOC Component . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105312. Possible Mechanisms for the Prolonged EPOC Component . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105313. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1056

In the recovery period after exercise there is an increase in oxygen uptakeAbstracttermed the excess post-exercise oxygen consumption (EPOC), consisting of arapid and a prolonged component. While some studies have shown that EPOCmay last for several hours after exercise, others have concluded that EPOC istransient and minimal. The conflicting results may be resolved if differences inexercise intensity and duration are considered, since this may affect the metabolicprocesses underlying EPOC. Accordingly, the absence of a sustained EPOC afterexercise seems to be a consistent finding in studies with low exercise intensityand/or duration. The magnitude of EPOC after aerobic exercise clearly dependson both the duration and intensity of exercise. A curvilinear relationship betweenthe magnitude of EPOC and the intensity of the exercise bout has been found,whereas the relationship between exercise duration and EPOC magnitude appearsto be more linear, especially at higher intensities.

Differences in exercise mode may potentially contribute to the discrepantfindings of EPOC magnitude and duration. Studies with sufficient exercisechallenges are needed to determine whether various aerobic exercise modes affectEPOC differently. The relationships between the intensity and duration of resis-tance exercise and the magnitude and duration of EPOC have not been deter-

1038 Brsheim & Bahr

mined, but a more prolonged and substantial EPOC has been found after hard-versus moderate-resistance exercise. Thus, the intensity of resistance exerciseseems to be of importance for EPOC.

Lastly, training status and sex may also potentially influence EPOC magni-tude, but this may be problematic to determine. Still, it appears that trainedindividuals have a more rapid return of post-exercise metabolism to resting levelsafter exercising at either the same relative or absolute work rate; however, studiesafter more strenuous exercise bouts are needed. It is not determined if there is asex effect on EPOC.

Finally, while some of the mechanisms underlying the more rapid EPOC arewell known (replenishment of oxygen stores, adenosine triphosphate/creatinephosphate resynthesis, lactate removal, and increased body temperature, circula-tion and ventilation), less is known about the mechanisms underlying the pro-longed EPOC component. A sustained increased circulation, ventilation and bodytemperature may contribute, but the cost of this is low. An increased rate oftriglyceride/fatty acid cycling and a shift from carbohydrate to fat as substratesource are of importance for the prolonged EPOC component after exhaustiveaerobic exercise. Little is known about the mechanisms underlying EPOC afterresistance exercise.

1. Excess Post-Exercise Oxygen neutral term excess post-exercise oxygen consump-Consumption (EPOC) tion (EPOC), which also includes the more pro-

longed increase in VO2 that may be observed forDuring exercise, there is an increase in oxygen hours after exercise.

uptake ( VO2) to support the increased energy need. EPOC consists of several components.[6,7] In thisAfter exercise, VO2 does not return to resting levelsreview, the term rapid component will be used toimmediately, but may be elevated above restingdescribe the sum of components that decays withinlevels for some period of time. Originally, the in-approximately 1 hour, whereas the prolonged com-creased VO2 after exercise was explained by theponent decays monoexponentially with a half-life inoxygen debt hypothesis. The theoretical basis forthe order of several hours (figure 1). Therefore,this was formulated by Hill et al.[1-4] They hypothe-processes active also beyond the first hour post-sised that the elevated VO2 after exercise was neces-

sary for the repayment of the oxygen deficit incurred exercise must be responsible for the prolongedafter the start of exercise, and ascribed the oxygen EPOC component.debt to the oxidative removal of lactate. Margaria et Training (i.e. repetitive bouts of exercise) mayal.[5] modified the concept, and suggested that the

also have a more chronic effect on resting metabolicoxygen debt consisted of a lactacid component

rate (RMR). In particular, this seems to be the casecaused by glycogen synthesis from lactate, and an

in trained compared with untrained individuals, es-alactacid component related to other factors. The

pecially when combined with high/sufficient energylactacid component was considered to be the slowerintake, resulting in a high energy flux or turnover.[8]component. However, the causality implied by theAt times it may be difficult to separate this effectterm oxygen debt is contrary to what is currentlyfrom the EPOC effect. In this review, we will onlyknown about the biochemical mechanisms underly-include studies of VO2 after an acute bout of exer-ing the increase in metabolism post-exercise. There-cise.fore, Gaesser and Brooks[6] introduced the causality

Adis Data Information BV 2003. All rights reserved. Sports Med 2003; 33 (14)

EPOC and Exercise Intensity and Duration 1039

Later, more controlled studies have been per-formed. Some studies have confirmed that there isan increase in VO2 after exercise that may last forseveral hours.[14-19] However, other studies haveconcluded that EPOC is transient and minimal afterexercise.[20-24] The conflicts in the results may beresolved if differences in exercise intensity and du-ration are taken into account, since this may beexpected to affect the metabolic processes underly-ing EPOC. Also, differences in exercise mode, train-ing status and sex may potentially contribute to thediscrepant findings.

3. Methodological Considerations

0 2 4 6 8 10 120

50

100

150

200

EPO

C (m

L/min)

Time after exercise (h)Fig. 1. Time plot of excess post-exercise oxygen consumption(EPOC) after exhaustive submaximal exercise (7180 minutes at6978% of maximal oxygen uptake; n = 12). The solid line showsthe prolonged EPOC component (reproduced from Bahr,[7] withpermission). There are several methodological issues that are

important to consider when studying EPOC. Accu-2. Early Studies on EPOC rate control over the pre-experimental conditions,

and an excellent reproducibility in the indirect calo-rimetry measures are prerequisites to be able toThe first report on an elevated RMR after physi-detect small, but potentially important differences.cal activity was published by Benedict and Carpen-Only few authors report the precision of the indirectter in 1910.[9] They observed a mean increase incalorimetry system used to measure VO2. TheRMR of 11.1% for their two study participantsDouglas bag method is generally considered to beduring sleep in a respiration calorimeter 713 hoursthe most accurate method of expired gas analysis,after severe exercise. Initially it was thought thatbut few authors, especially of newer studies, havepost-exercise elevation in VO2 contributed signifi-used this technique. Instead, automated systemscantly to the energy cost of exercise, and would behave been used, often with unknown validity andan important factor in daily energy expenditure.reliability.Herxheimer et al.[10] noted that the VO2 of five

untrained individuals did not return to baseline until Furthermore, the pre-experimental conditions3648 hours after exercise, and Edwards et al.[11] have not always been well controlled. The studyreported a 25% elevation in metabolism 15 hours participants should have a stable weight, and foodafter cessation of 2 hours of strenuous football. intake and exercise should be controlled. It is alsoAlso, Passmore and Johnson[12] found a 15% in- advisable for study participants to sleep overnight increase in RMR for 7 hours after a 16km walk at 6.4 the laboratory before a study to avoid exercise in thekm/hour in three males, and deVries and Gray[13] morning; however, an outpatient protocol may givefound a 10% increase in RMR for 6 hours after 1 no different values than an inpatient protocol whenhour of mixed aerobic exercise. However, in many the conditions are controlled and the study partici-cases, the intensity and duration of exercise was not pants are transported to the laboratory.[25,26] Also,quantified in these early studies, and they provided habituation of the study participants to testing proce-minimal information about the controls. Also, they dures is of utmost importance. The experimentaldid not account for other factors that may influence conditions both before and during measurementsRMR, such as time of day, prior uncontrolled exer- need to be strictly controlled. For female studycise, food, temperature, caffeine intake, habituation participants, it may also be necessary to control forand stress. menstrual cycle differences.


1040 Brsheim & Bahr

When reviewing even the newer EPOC literature, tilatory threshold,[21] and beyond 40 minutes after 4it is a problem that the methods for measuring 20 minutes of cycle ergometry at 3555% ofbaseline and EPOC duration are inconsistent among VO2max.[22] Only two males and two females tookinvestigations. In some studies, a separate control part in the last study. Brehm and Gutin[23] found astudy has been used to control for time effects, relationship between EPOC and the intensity ofwhereas others have used only one pre-exercise walking/running, but their intensity was still low,value as baseline. In many studies, only 30 minutes the highest being 11.3 km/hour in trained individu-of rest in the morning has been used and the VO2 als. After 3.2km running at this intensity, EPOCduring the final 10 minutes of this has been taken as amounted to only 71kJ (~3.5L oxygen). Elliot etbaseline for EPOC. This can lead to falsely high al.[24] also found a short lasting (

EPO

C and

Exercise Intensity and

Duration

1041

A

dis D

ata

Info

rma

tion

BV 2003. A

ll righ

ts rese

rved

.Sp

orts M

ed

2003; 33 (14)

Table I. Studies on the effect of aerobic exercise on post-exercise VO2a

Study Year Study Exercise mode, duration and intensity EPOC size and duration Commentscparticipantsb

Passmore & Johnson[12] 1960 3M TM: 16km, 6.4 km/h RMR 1418% for 6h, >7h Rest position not reported. NCdeVries & Gray[13] 1963 2M Cycling, bench step, run/walk: 45 min ~1.9L (57 kcal) NC

(effective time 25 min)Knuttgen[28] 1970 5F, 7M Cycling: 4598%, 1555 min 5L, duration NA Bed rest. NC. EPOC magnitude

related to intensity and durationSegal & Brooks[29] 1979 11M Cycling: 55 and 95%, 2 min 4L, duration NA Seated. NC. EPOC magnitude

related to intensityHagberg et al.[20] 1980 18M Cycling: 50%, 65% and 80%, 5 and 20 5L, duration NA Moderate cycling (used as

min base-line). NC. EPOCmeasured for 15 min

Hermansen et al.[30] 1984 1M Cycling: 75%, 80 min 12h EPOC: 48L. At 24h: VO2 Bed rest. C5.9%

Bielinski et al.[14] 1985 10M (T) TM: 50%, 3h RMR 9% for 4.5h. At 18h: Seated/respiratory chamber. C.RMR 4.7% Food given 30 min post-

exercisePacy et al.[22] 1985 2F, 2M (T) Cycling: 3555% for 20 min 4 (40

1042B

rsheim &

Bahr

A

dis D

ata

Info

rma

tion

BV 2003. A

ll righ

ts rese

rved

.Sp

orts M

ed

2003; 33 (14)

Table I. ContdStudy Year Study Exercise mode, duration and intensity EPOC size and duration Commentsc

participantsbPoehlman et al.[35] 1989 6M Cycling: 50%, 90 min

EPO

C and

Exercise Intensity and

Duration

1043

A

dis D

ata

Info

rma

tion

BV 2003. A

ll righ

ts rese

rved

.Sp

orts M

ed

2003; 33 (14)

Table I. ContdStudy Year Study Exercise mode, duration and intensity EPOC size and duration Commentsc

participantsbSmith & McNaughton[51] 1993 8F, 8M Cycling: 40%, 50%, 70%, 30 min M: 16.3, 22.1, 28.1L for 31.2, Rest position not reported. NC.

42.1, 47.6 min. F: 12.1, 20.8, EPOC duration increased with24.3L for 26.9, 35.6, 39.1 min intensity

Neary et al.[52] 1993 7M (T) Swimming & cycling: 65% of mode- Swim: 5.3, 5.6, 5.6L; 8.1, 10.1, Seated. NC. Relationshipspecific, 30, 45, 60 min 9.4 min. Cycling: 8.2, 9.9, between core temperature and

10.0L; 18.3, 20.4, 22.9 min EPOCFrey et al.[53] 1993 13F (7UT + 6T) Cycling: 65% and 80%. 2445 min UT: 4.0, 5.9L, >1h. T: 4.7L, 50 Seated. NC

(until 300 kcal) min; 5.6L, 40 minBrsheim et al.[54] 1994 6M Cycling: 78%, 60 min 6.5h EPOC: 14.4L, 6.5h Bed rest. C. EPOC measured

for 6.5hThomas et al.[55] 1994 7M Cycling, jog and downhill jog: 60%, 60 Magnitude NA, 3h Seated. C. EPOC measured for

3hHarms et al.[59] 1995 16M TM: 70%. 20 min Magnitude unclear, 1h Seated. NCPhelain et al.[63] 1997 8F (T) Cycling: (a) 50%, 78 min; (b) 75%, 51 3h EPOC: (a) 4.8L, 3h measured for 3hLaforgia et al.[64] 1997 8M (T) TM: (a) 70%, 30 min; (b) 105%, 20 1 (a) 6.9L, 1h; (b) 15L, 8h Bed rest. C

min with 2 min break (equal work)Short & Sedlock[65] 1997 11F, 11M Cycling: (a) 70%, 30 min; (b) 1.5L (a) UT: 3.5L, 50 min; T: 3.2L, Seated. NC. EPOC duration

(10UT + 12T) oxygen/min, 30 min 40 min; (b) UT: 2.4L, 39 min; T: shorter in T1.5L, 21 min

Burleson et al.[66] 1998 15M TM: 45%, 27 min ~3.4L, 4.5h Bed rest. C. EPOC measured

for 4.5h

Continued next page

1044 Brsheim & Bahr

31.9L of oxygen after 20, 40 and 76 minutes at 70%of VO2max, respectively. After the longest bout,EPOC duration was at least 10 hours.

Similar comprehensive studies have been doneby Gore and Withers.[18,19] Their treatments rangedfrom a 20-minute walk at 30% of VO2max to an80-minute run at 70% of VO2max. The maximalEPOC was 14.6L oxygen (297kJ) after the longestand hardest bout. In accordance with our stud-ies,[16,43] they found an exponential relationship be-tween exercise intensity and the magnitude ofEPOC. They calculated that the intensity explainedfive times more of EPOC than either exercise dura-tion or total work completed.[18] While intensityaccounted for 45.5% of the variation in EPOC, theduration of exercise, and the interaction betweenintensity and duration accounted only for 8.9% and7.7%, respectively.

In our studies,[15,16,43] we found much higherEPOC values, and also more prolonged duration ofEPOC, compared with Gore and Withers.[18,19] In thestudy by Mhlum et al.,[15] the response was abouttwice the response of that observed in the study byGore and Withers.[18] Even though the measurementperiod was 4 hours shorter in the latter study, thiscan still not explain the difference. While we usedcycling in our experiments, Gore and Withers usedrunning, and the different exercise modes may be apossible explanation of the discrepancy in results.Another possible explanation, perhaps more likely,is the different training status of the study partici-pants. In the studies by Gore and Withers,[18,19] theparticipants were well trained, whereas in our stud-ies they were not.

In support of a significant EPOC after high-intensity exercise, a 37% increase in energy expen-diture was found as late as 1216 hours after inter-mittent exercise for a total of 71 minutes at 85% ofVO2max.[32] The most exhaustive bout for any of theEPOC studies was a 35km road run (intensity about70% of VO2max for about 160 minutes).[42] A totalEPOC of 32.4L of oxygen, lasting for a total of 8hours, was found. Furthermore, an EPOC lasting for3 hours was observed after only 30 minutes tread-mill walking at 70% of VO2max in young trained


Tabl

e I.

Cont

dSt

udy

Year

Stud

yEx

erci

se m

ode,

dur

atio

n an

d in

tens

ityEP

OC

size

and

dura

tion

Com

men

tsc

parti

cipan

tsb

Brs

heim

et a

l.[69]

1998

8MCy

cling

: 56%

, 90

min

4.5h

EPO

C: 8

.1L,

>4.

5hBe

d re

st. C

. EPO

C m

easu

red

for 4

.5h

Mat

suo

et a

l.[70]

1999

7FCy

cling

: 60%

, 60

min

6h E

POC:

8.4

L (fo

llicula

r);R

est p

ositio

n no

t rep

orte

d. C

.12

.3L

(lutea

l), du

ration

NA

EPO

C m

easu

red

for 6

h

Lee

et a

l.[71]

1999

10M

TM: (

a) 40

%, 40

min;

(b) 8

5%, 2

0 min

Mag

nitu

de N

A. (a

) 10 m

in; (b

)Su

pine

. NC

30 m

in

Fuku

ba e

t al.[7

2]20

005F

Cycli

ng: 7

0%, 6

0 m

in7h

EPO

C: 8

.6L

(follic

ular);

8.9L

Bed

rest

. C(lu

teal),

durat

ion N

A

aAl

l stu

dies

are

in y

oung

peo

ple.

Exe

rcise

is s

hown

in p

erce

ntag

e of

VO

2max

or

VO2p

eak.

EP

OC

size

is gi

ven

in li

tres

of o

xyge

n.

bIf

noth

ing

is m

entio

ned,

indi

vidua

ls we

re u

ntra

ined

/med

ium

trai

ned.

c In

divid

uals

po

sitio

n du

ring

mea

sure

men

ts o

f res

ting

VO2

is g

iven

first

.

C =

sepa

rate

rest

con

trol e

xper

imen

t; co

nt =

co

ntin

uous

; EPO

C =

exc

ess

pos

t-exe

rcise

oxy

gen

cons

umpt

ion;

F

= fe

mal

es; M

=

male

s; m

ax =

ma

xim

um; N

A =

dat

a no

t ava

ilabl

efo

r est

imat

e; N

C =

no r

est

con

trol e

xper

imen

t, bu

t pre

-exe

rcise

val

ues

used

as

base

line

inst

ead;

NS

= not s

igni

fican

t; R

MR

=

re

stin

g m

etab

olic

rate

; T =

tra

ined

; TM

=

tre

adm

ill;UT

=

untra

ined

; VO

2 =

oxy

gen

upta

ke;

VO2m

ax =

m

axi

mal

oxy

gen

upta

ke;

VO2p

eak

= pe

ak o

xyge

n up

take

; in

dica

tes

incr

ease

.


Exercise duration (min)0 20 40 60 80

0

10

10

20

30

40

Exercise intensity (%)0 20 40 60 80

EPO

C (L)

0

10

20

30

40

50

a b

Fig. 2. (a) Plot of excess post-exercise oxygen consumption (EPOC) magnitude versus exercise intensity (constant duration of 80 minutes).(b) Plot of EPOC magnitude versus exercise duration (constant intensity of 70% of maximal oxygen uptake). Different symbols are used forindividual study participants (reproduced from Bahr,[7] with permission).

women.[58] Also, a comparison of interval-type exer- protocols have supported the conclusion that exer-cise intensity is more important than total work forcise alternating between 30% and 90% of VO2max,EPOC magnitude.[51,60]with continuous exercise of equal duration (36 min-

utes) at 60% of VO2max (equal work) showed a There are also some studies of the effect oflonger EPOC (38 versus 17 minutes) and higher intensity on EPOC, where EPOC has been measuredEPOC magnitude (~3.6 versus ~1.9L oxygen) after for a specific pre-determined time period. Phelain etthe interval exercise.[50] al.[63] found a significantly higher 3-hour EPOC

after cycling at 75% versus 50% of VO2max (equalSedlock et al.[33] also investigated the effect ofwork, 500 kcal) in women. Brockman et al.[49] inves-exercise intensity and duration on EPOC, but used atigated 1-hour EPOC after intense intermittent run-more narrow spectrum of exercise conditions. In onening (7 2 minutes at 90% of VO2max, 2 minutes ofseries of experiments, caloric output was kept con-rest between intervals), continuous running for 10stant (300 kcal), whereas intensity was varied (50%minutes at 81% of VO2max, and continuous walkingversus 75% of VO2max). It took only 30 and 20for 2 hours at 24.5% of VO2max. EPOC was higherminutes, respectively, for the trained male studyafter running versus walking and highest after theparticipants to finish these work bouts. In anotherhighest intensity running. However, since EPOCseries, intensity was kept constant (50% of VO2max),was still increased after 1 hour, no definite conclu-whereas duration was varied (30 versus 60 minutes).sion can be drawn.Hence, no strenuous exercise was performed, and no

substantial EPOC duration or magnitude was found. In contrast to the findings of an intensity-relatedStill, it was concluded that the intensity of exercise EPOC, Chad and Wenger[31] did not find any rela-influenced both the magnitude and duration of tionship between exercise intensity and EPOC. TheyEPOC, whereas exercise duration only influenced had one group of individuals cycling for 15 and 30EPOC duration. It was also concluded that the dura- minutes at 70% of VO2max. Another group cycledtion of EPOC and the subsequent caloric output was for 38 minutes at 50% of VO2max and 30 minutes atnot necessarily related, since high-intensity exercise 70% of VO2max (equal total work). They found thatof short duration produced a higher EPOC (~6L EPOC was highest after the longest duration in bothoxygen) than lower intensity exercise of long dura- groups. In another study, Chad and Quigley[38] hadtion (~2.5L oxygen), even though no difference in trained and untrained women cycle for 30 minutes atEPOC duration (33 versus 28 minutes, respectively) 50% and 70% of VO2max, and found a higher EPOCwas found. Similar studies with modest exercise after the lowest intensity bout. The results of these


1046 Brsheim & Bahr

0

10

20

30

40

50

EPO

C (L)

02040

6080100

120140

Exercise duration (min) 0

20

40

60

80100

120

Exer

cise i

ntens

ity

(% of

VO2m

ax

or VO

2pea

k)

Fig. 3. Relationship between exercise intensity, exercise duration and excess post-exercise oxygen consumption (EPOC) magnitude. Plotshows mean EPOC values from studies that have used cycling exercise, and where EPOC values are reported or possible to esti-mate.[15-17,22,24,30,33,39,40,43-47,51-54,56,57,60,63,65,67-70,72] Note that recovery oxygen uptake was not measured until it had returned to resting controlvalues in all the included studies (for more details on studies, see table I). VO2max = maximal oxygen uptake; VO2peak = peak oxygenuptake.

two studies are somewhat puzzling when compared was increased from 30 minutes to 45 and 60 min-with other findings. The authors explain the results utes, respectively. After the most exhaustive bout,with a lower respiratory exchange ratio (R-value) EPOC lasted for 7.5 hours. Sedlock,[40] on the otherafter the low intensity bout. It should also be men- hand, did not find any effect of exercise duration ontioned that post-exercise VO2 was still elevated at EPOC, but the intensities used (40% versus 60% ofthe end of the experiments 3 hours post-exercise in VO2max) were low.the second study, whereas EPOC duration is unclear The interaction between exercise intensity andin the first study. Also, the experimental conditions duration is not completely understood, and it iswere less rigorously controlled (menstrual cycle difficult to separate the effect of each of these fac-does not appear to be controlled for in the female tors. However, the fact that exercise has to be of aparticipants, participants were sitting during rest and certain intensity for the linear relationship betweenwere allowed to write, watch television and read, the exercise duration and EPOC magnitude to becometime of the day for a repeated trial appears to be apparent, indicates that the interaction between themslightly different, and VO2 was recorded in short is synergistic rather than additive. This is illustratedperiods). in figure 3, which shows mean EPOC values from

Chad and Wenger[17] also investigated the effect studies that have used cycling exercise, and whereof duration (30, 45 and 60 minutes) on EPOC after EPOC magnitude is presented or can be estimat-cycling at 70% of VO2max. They found that EPOC ed.[15-17,22,24,30,33,39,40,43-47,51-54,56,57,60,63,65,67-70,72] Itincreased 2.3- and 5.3-fold when exercise duration should be noted that some of the studies included



probably did not capture the entire EPOC, since 70% of VO2max), the corresponding values can beestimated to approximately 700kJ after the exercisepost-exercise VO2 was still elevated compared withbout, and about 109 200kJ per year (training threeresting control values at the end of the measurementtimes per week), equivalent to about 2.9kg of fat.period (for further details on these studies see tableHowever, such strenuous exercise cannot be expec-I). Gore and Withers[19] analysed the interactionted to be undertaken by overweight or untrainedeffect between exercise intensity and duration onindividuals. Although overweight may be the resultEPOC after treadmill exercise ranging from 20 min-of a small positive energy balance over a long time,utes at 30% of VO2max to 80 minutes at 70% ofand EPOC may contribute to the opposite whenVO2max. They found that the interaction betweenstrenuous exercise is undertaken regularly, it mustexercise intensity and duration accounted for a muchbe concluded that EPOC is negligible in relation tosmaller part (7.7%) of the variation in EPOC, com-weight loss in the overweight or obese person. Itpared with exercise intensity alone (45.5%). Themust be noted that exercise per se has an importantstudies illustrated in figure 3 appear to support therole for weight regulation. Furthermore, EPOC mayhypothesis that exercise intensity has to be of ahave greater implications for elite athletes, whocertain size to achieve a significant EPOC. In addi-often have two bouts of exercise on the same day astion, they show that the highest EPOC values havea normal training routine. If VO2 is elevated from abeen found when both exercise intensity and dura-previous exercise bout, the mechanical efficiencytion are substantial.may be reduced during the following bout.[74]

It appears that the post-exercise increase in ener- In summary, the magnitude of EPOC after aero-gy expenditure per se after exercise bouts spanning bic exercise is clearly dependent on both the dura-from 20 minutes at 30% VO2max to 80 minutes at tion and intensity of exercise. There is a curvilinear70% VO2max may be negligible in relation to energy relationship between the magnitude of EPOC andbalance and weight loss. However, this may be a the intensity of the exercise bout. At least for cy-speculative conclusion, since there are no available cling, it appears that an intensity above 5060% ofdata from prolonged studies of the effect of EPOC

VO2max is required in order to induce an EPOCon energy balance. Also, it should be noted that lasting for several hours. The relationship betweenthere may be a high inter-individual variability in exercise duration and EPOC magnitude appears toEPOC in response to the same relative exercise be more linear, especially at higher intensities.stimulus, as can be seen from figure 2. Thus, it canbe hypothesised that there are high, medium and low 5. Effect of Split Exercise Sessionsresponders, corresponding to what has been found on EPOCfor other parameters in response to exercise train-ing.[73] However, there are no studies where EPOC A couple of studies have shown a higher EPOChas been measured repeatedly in the same individu- after split exercise sessions compared with a contin-als under the same conditions. Therefore, it is not uous bout. Kaminsky et al.[36] investigated womenknown if the observed variability is the result of during 50 minutes of continuous running versus twobiological variation related to differences in, for 25-minute sessions at 70% of VO2max. The com-example, body composition or training status, or bined EPOC after the two split sessions correspond-whether it is caused by measurement errors. Still, if ed to ~3.1L versus 1.4L of oxygen after the continu-we estimate that EPOC amounts to 50100kJ after ous session. Almuzaini et al.[67] compared EPOCmoderate exercise (1 hour, approximately 50% of after 30 minutes of continuous cycling versus twoVO2max), this would result in an extra energy loss of 15-minute sessions (separated by 6 hours) at 70% ofabout 11 700kJ per year if training is undertaken VO2max. In this study, the sum of EPOC after thethree times per week. This represents only about split sessions was 7.4L versus 5.3L of oxygen after311g of fat. After more strenuous exercise (1 hour, the continuous bout. Hence, total EPOC was ap-


1048 Brsheim & Bahr

proximately 120% and 40% greater after split ses- VO2max, very similar 1-hour EPOCs were found;sions compared with a continuous session in these 7.6L (submaximal exercise) and 7.8L (supramax-two studies. It should be noted that even though imal exercise), respectively. Of this, about 4.5LEPOC is higher after split sessions, the extra EPOC (submaximal) and 2.0L (supramaximal) could beis small in relation to the exercise energy expendi- attributed to the prolonged curve component, andture. Thus, prolonging the exercise bout by a few about 3.1L (submaximal) and 5.8L (supramaximal)minutes may make up for the increase in EPOC after to the rapid component. Hence, it appears that high-splitting the session. Furthermore, in split sessions, intensity short exercise affects mainly the rapidthere will be one oxygen deficit for each session. EPOC component, whereas more prolonged ex-This means that the relative difference in energy hausting exercise stimulates mechanisms also pre-expenditure during recovery after split versus con- sent beyond the first hour of recovery. However, thistinuous sessions will be smaller than the difference relationship remains to be fully elucidated.in EPOC. This was not taken into account in the twostudies mentioned.[36,67] 7. Effect of Aerobic Exercise Mode

on EPOC6. Effect of Supramaximal Exercise

A curvilinear relationship between exercise in-on EPOCtensity and EPOC, and a linear relationship between

Given the relationship between exercise intensity exercise duration (when intensity is higher than theand EPOC, it comes as no surprise that supramax- break-point) are both shown for cycle exercise[16,43]imal exercise stimulates EPOC. We found that brief and for treadmill exercise.[18,19] However, the abso-intermittent bouts of exhaustive supramaximal exer- lute EPOC magnitude for a certain exercise intensitycise (1, 2, or 3 2-minute bouts of cycling at 108% and duration may differ depending on exerciseof VO2max, separated by 3-minute rest periods) ele- mode.vated post-exercise energy expenditure for 4 Muscle damage is more likely to occur afterhours.[45] eccentric-type exercise than after concentric exer-

Laforgia et al.[64] measured EPOC after cise,[75-77] and it is possible that this influencessupramaximal (20 1 minute runs at 105% of EPOC. However, in studies where concentric andVO2max with 2-minute recovery periods between) eccentric exercise have been compared, no differ-versus submaximal running (30 minutes at 70% of ences have been found in EPOC[47,55] or RMR forVO2max). A significantly higher 9-hour EPOC was several days after exercise.[78] Sedlock[47] comparedobserved after the supramaximal exercise. Interest- 30 minutes of cycling and treadmill exercise at aningly, the EPOC values in this study were similar to intensity of 6065% of mode-specific peak oxygenthe values found after 80 minutes of running at 70% uptake ( VO2peak). No differences could be detectedVO2max in the study by Gore and Withers,[18,19] even between modes, but the exercise was moderate, andthough more than twice the total work was per- hence EPOC was small (1517 kcal; ~3.5L oxygen).formed in the latter study. EPOC has also been compared after 60 minutes of

either jogging, downhill jogging or cycling at 60%Few authors have tried to divide EPOC intoof mode-specific VO2max to induce different de-different components. This is unfortunate, since ex-grees of eccentric muscular activity.[55] No increaseercise intensity and duration may affect the shortin recovery energy expenditure was found after ec-and prolonged components to a different extent. Wecentric exercise, but it may be that the detectioncalculated the short EPOC component as the differ-level was not sufficient (power = 0.40 for mediumence between the observed 1-hour EPOC and theeffect size).prolonged curve component.[7] By comparing EPOC

after 7080 minutes of cycling at 6978% of To investigate the effect of relative metabolicVO2max with 3 2 minutes at 104117% of rate of the active musculature on EPOC, Sedlock[39]



compared the effect of 20 minutes of arm crank 14.5 hours after the resistance exercise comparedexercise versus cycling at 60% of mode-specific with a resting control experiment.VO2peak. VO2 during arm crank exercise was about Burleson et al.[66] compared weight training exer-72% of uptake during cycling in absolute terms. No cise (two circuit sets of eight exercises at 60% ofdifference in EPOC was found between modes, but 1RM for 812 repetitions) and treadmill exerciseagain the exercise was short and not very strenuous, (27 minutes at 45% of VO2max). The weight trainingand consequently EPOC lasted less than 25 minutes.

exercise was performed first, and the average VO2Short et al.[61] have investigated the effect ofwas used to determine the intensity for the treadmill

intensity and duration of upper body exercise aloneworkout. EPOC was found to be higher the first 30

on EPOC, and found a similar pattern as for theminutes after resistance exercise, but not at 60 andlower body. EPOC was measured after 15 and 30 90 minutes. Even though the VO2 volumes were

minutes of arm crank exercise at 35% of VO2peak,equated, the resistance exercise was considered to

and after 15 minutes at 70% of VO2peak. The intensi- be of higher intensity activity than the aerobic exer-ty was found to have a greater effect than duration,cise, and this may explain the higher EPOC afterbut durations were low and within a narrow spec-resistance exercise. Hence, it is difficult to comparetrum in this study. Since exercise duration and inten-resistance exercise to steady-state exercise, since it

sity did not vary over a range of values, it was notis not easy to precisely quantify the energy cost ofpossible to fully describe the relationships withresistance exercise with indirect calorimetry.EPOC.

The relationship between the intensity and dura-In summary, it is not clear whether varioustion of resistance exercise, and the magnitude andmodes of aerobic exercise affect EPOC differently.duration of EPOC has not been determined. Table IIFurther studies with a sufficient exercise challengeshows a review of studies on the effect of resistanceand adequate statistical power are needed.exercise on EPOC.

8. Effect of Resistance Exercise on EPOC Williamson and Kirwan[89] and Dolezal et al.[84]found that RMR remained elevated for 48 hours

EPOC has also been compared between aerobicafter an acute moderate- to high-intensity bout of

and resistance exercise. Elliot et al.[46] comparedresistance exercise. This was hypothesised to be due

aerobic cycling (40 minutes at 80% of maximal largely to protein turnover and tissue repair. Itheart rate), circuit weight training exercise (4 sets, 8should be noted that to avoid eccentric muscle work,

exercises, 15 repetitions at 50% of one repetitiononly the concentric phase was used in the study of

maximum [1RM]), and heavy resistance exercise (3 Williamson and Kirwan,[89] and still they found asets, 8 exercises, to exhaustion at 8090% of 1RM). prolonged effect on recovery metabolic rate. TheirThey found that heavy resistance exercise produced

study was done in 59- to 77-year-old men. Similarly,the biggest EPOC, but it is unclear how the workMelby et al.[81] found that VO2 was still increased byvolumes related to each other.9.4% and 4.7% as late as 15 hours after two hardStudies in which similar estimated exercise ener-resistance exercise work bouts (each 90 minutes ofgy cost[57] or similar exercising VO2[66] have beenweight lifting, six sets of ten exercises, 812 repeti-

used to equate continuous aerobic exercise and in-tions at 70% of 1RM).termittent resistance exercise, have indicated that

After more moderate resistance exercise (threeresistance exercise produces a greater EPOC res-sets of seven exercises, ten repetitions at 12RM),[79]ponse. A higher EPOC was found after hard resis-a smaller EPOC was found, but the VO2 was mea-tance exercise (50 sets of 812 repetitions at 70% ofsured for only 1 hour after exercise, and was still1RM, 2 minutes of rest between sets) compared withelevated at this time point. Binzen et al.[86] found anan equated work bout of aerobic cycling (50% ofEPOC shorter than 2 hours after three sets of tenVO2max for 60 minutes).[57] RMR was still elevated


1050B

rsheim &

Bahr

A

dis D

ata

Info

rma

tion

BV 2003. A

ll righ

ts rese

rved

.Sp

orts M

ed

2003; 33 (14)

Table II. Studies on the effect of resistance exercise on post-exercise VO2a

Study Year Study Exercise EPOC size and duration Commentsbparticipants

Melby et al.[79] 1992 6M 4 sets of upper and 3 sets of lower body. 710 1h EPOC: ~4L (20 kcal), 1h Supine. C. EPOC measured for 1h onlyreps. 12RM

Elliot et al.[46] 1992 5F, 4M 8 exercises, upper and lower body. (a) 4 sets, 15 (a) ~10.2L (48 kcal); (b) ~10.6L Supine. C. EPOC duration unclearreps, 50% of 1RM, 30 sec rest; (b) 3 sets, 38 (51 kcal). 30 minreps, 8090% of 1RM, 30 sec rest

Murphy & 1992 10M 6 exercises, upper and lower body. Circuit: 3 Circuit: 5L, 20 min. Standard: Rest position not reported. NC.Schwarzkopf[80] times, 812 reps, 50% of 1RM, 30 sec rest. 2.7L, 15 min Experiments in the afternoon. 5 min

Standard: 3 sets, reps to exhaustion, 80% of baseline measurement1RM, 120 sec rest

Melby et al.[81] 1993 (a) 6M; (b) (a) 6 sets of 10 exercises, upper and lower body, (a) 2h EPOC: 7L, 15h; (b) 2h Supine. (a) NC; (b) C. EPOC measured6M ~812 reps, 70% of 1RM, 90 min; (b) as (a), but EPOC: 7L, 15h for 2h, minus first 5 min post-exercise.

5 sets, and longer rest, 90 min Measured again at 15hOlds & 1993 7M 2 sets of 7 exercises, upper and lower body, 3.5 (a) 8.2L,


exercises with ten repetitions at 10RM (1 minute of multiple sets), weights, sets, repetitions, and lengthrest between sets) in resistance-trained women. of rest periods. These factors will all influence the

energy cost of the exercise, but this effect is difficultMurphy and Swartzkopf[80] compared two differ-to quantify precisely. Furthermore, resistance train-ent protocols of resistance exercise (three sets of sixing is similar to interval training and split sessions ofexercises, repetitions to exhaustion at 80% of 1RMaerobic exercise in that each session or set of resis-with 120-second rest periods versus three circuit setstance exercise will have an EPOC of its own duringof the six same exercises, 1012 repetitions at 50%the recovery period between exercises. To deter-of 1RM with 30-second rest periods). Work volumemine the total energy expenditure, this has to beof each session was similar, but intensity (weightincluded in the calculations.lifted per unit of time) was greater for the circuit

All this taken into consideration, it seems likelyexercise. The higher intensity session producedthat EPOC after resistance exercise is influenced byhigher EPOC than the standard set exercise (4.9the intensity of the exercise, since a more prolongedversus 2.7L oxygen), but the duration of EPOC wasand substantial EPOC has been found after hardonly 20 minutes.versus more moderate exercise. More research isOlds and Abernethy[82] compared high- and low-still needed to elucidate the effect of intensity andintensity resistance exercise with equated work vol-duration of resistance exercise on the magnitude and

ume, and found no difference in EPOC. However,duration of EPOC.their range of intensities was narrow (12 repetitions

at 75% of 1RM and 15 repetitions at 60% of 1RM),9. Effect of Training Status on EPOCand may not have been large enough to elicit a

treatment effect. Accordingly, VO2 returned to Individuals of different fitness levels have beenbaseline within 1 hour after exercise. Also, the age

used in EPOC studies, and this may also potentiallyrange was wide (2255 years) and there was a high

explain some of the differences observed in magni-inter-individual difference in EPOC (0.727L). tude and duration of EPOC. The effect of training

In a recent study, Thornton and Potteiger[87] in- status is not easy to study, since comparing groupsvestigated the effect of high- and low-intensity resis- of different fitness levels at the same absolute exer-tance exercise of similar work volume on EPOC. cise intensity level means that trained are working atThe exercise bouts consisted of two sets of nine a lower relative intensity, which has been shown toexercises. In the high-intensity bout, eight repeti- influence EPOC. Furthermore, if trained and un-tions at 85% of 8RM of each exercise were per- trained are compared at the same relative exerciseformed, whereas 15 repetitions at 45% of 8RM were intensity, when total work is equal, the untrainedperformed in the low-intensity bout. The rest period have to work for a longer duration, which may alsobetween sets was 2 minutes. The duration was 23 influence the results. In other words, there is nominutes for the high-intensity bout and 26 minutes study design available to provide a definite answerfor the low-intensity bout, whereas the actual exer- to whether training status affects EPOC.cise time was 6.9 minutes (high) and 8.3 minutes Brehm and Gutin[23] found similar values for(low), respectively. VO2 was measured between EPOC in runners versus non-exercisers following a020, 4560, and 105120 minutes post-exercise. 3.2km walk at 6.4 km/hour (i.e. same absolute inten-The magnitude of EPOC was higher during each sity), but the exercise intensity may have been toomeasurement period after high- versus low-intensity low to detect a difference. Sedlock[56] compared fitresistance exercise. and unfit males after cycling at 50% of VO2peak,

Understanding the effect of resistance exercise until 300 kcal were used (27 and 35 minutes toon EPOC is confounded by the considerable diversi- finish, respectively). No difference in EPOC dura-ty in the protocols commonly employed in this re- tion or magnitude was found, but again, the exercisesearch, for example, type (i.e. circuit training or challenge was low. Finally, in a study of treadmill


1052 Brsheim & Bahr

exercise at approximately the anaerobic threshold, it more, the magnitude of EPOC was not differentbetween groups after exercise at the same relativewas concluded that fitness level does not significant-intensity (3.2L oxygen in trained versus 3.5L oxy-ly alter magnitude and duration of EPOC, but bothgen in untrained). The trained group had higher VO2the separation of study participants into groups andat the end of the exercise bout, and if the data werethe exercise intensity were somewhat unclear in thatnormalised to percentage change afterwards, thestudy.[21]trained had a more rapid fall in post-exercise VO2.In contrast to these studies, Chad and Quigley[38]After exercise of similar absolute intensity, the un-found a higher 3-hour EPOC in trained female cy-trained group had higher EPOC (2.4L oxygen) ver-clists versus untrained women after 30 minutes cy-sus trained (1.5L oxygen). Longitudinal trainingcling at 50% or 70% of VO2max. The difference wasstudies also support the findings of a faster recoverymost apparent immediately after exercise, whichin VO2 in trained individuals.[90,91]was explained by higher absolute exercise intensity,

In summary, it appears that trained individualsand thereby a higher VO2 in trained versus un-have a more rapid return of post-exercise metabol-trained. VO2 was still increased 3 hours after exer-ism to resting levels when exercising at either thecise, and no definite EPOC magnitude could besame relative or same absolute work rate, but studiesdetermined.after more strenuous exercise bouts should be done.

Frey et al.[53] measured VO2 for 1 hour aftercycling at ~65% and ~80% of VO2max until 300 kcal 10. Effect of Sex on EPOChad been expended in trained and untrained women.No difference in EPOC magnitude was found after Sex is also a factor that can potentially influencecycling at the highest intensity, but EPOC was sig- EPOC, as well as have implications for study de-nificantly smaller in the untrained (4.0L oxygen) sign. Energy expenditure at rest or during exerciseversus the trained group (4.7L oxygen) after the may vary with menstrual phase,[92-95] and this haslowest intensity. EPOC decreased rapidly during the not always been taken into consideration when stud-first 10 minutes post-exercise in both groups. Fol- ying EPOC. Basal metabolic rate has been shown tolowing both intensities, the initial EPOC (first 10 be at its lowest level 1 week before ovulation.[92]minutes post-exercise) was greater in trained versus Webb[96] found an 816% increase in 24-hour ener-untrained, most likely because of higher VO2 during gy expenditure during the 14-day luteal phase fol-exercise in this group. While there still was a signif- lowing ovulation. Accordingly, Matsuo et al.[70]icant EPOC at the end of the experiment in the found a higher EPOC after 60 minutes cycling atuntrained group, post-exercise VO2 was elevated for 60% of VO2max in the luteal versus the follicularonly 50 and 40 minutes after low- and high-intensi- phase in seven healthy women. On the other hand,ty, respectively, in the trained group. Thus, EPOC Fukuba et al.[72] did not detect any significant effectduration was shorter in the trained. of menstrual cycle on EPOC in five young women

after 60 minutes cycling at 70% of VO2max.Short and Sedlock[65] compared EPOC in trained( VO2max: 53 mL/kg/min) versus untrained males When comparing EPOC between men and(37 mL/kg/min) after 30 minutes cycling on both women, the same question as for trained versussimilar absolute (1.5L oxygen/min) and relative untrained must be addressed: should EPOC be com-(70% of VO2peak) intensities. At a VO2 of 1.5 L/ pared after absolute or relative workloads? Berg[41]min, the trained group exercised at 45% of VO2peak, measured VO2 for 1 hour after 30 minutes of exer-while the untrained worked at 61% of VO2peak. The cise at 40% of VO2max in active men and women. Aresults showed that the trained group had a signifi- higher post-exercise energy expenditure was foundcantly shorter duration of EPOC whether compared among men. However, the calculations are some-at the same absolute or relative intensity, but in all what unclear, since the data are not compared withsituations EPOC lasted less than 1 hour. Further- control resting values, and hence it is the absolute



levels rather than the increase in energy expenditure ture may contribute; however, the cost of this is lowafter exercise that is compared. (

1054 Brsheim & Bahr

pared with rest where more of the carbohydrates nephrine and norepinephrine are potent stimulatorsmay be stored as fat. The energy cost of storing of the energy metabolism.[118,119] Their calorigenicdietary carbohydrates as fat requires 2324% of the effect seems to be mediated through the -adre-ingested energy, whereas storage as glycogen re- noceptors.[120,121] Secondly, the sympathoadrenalquires only 5.3%.[106] On the other hand, emerging system is activated during exercise with elevateddata[107,108] suggest that hepatic de novo lipogenesis concentrations of plasma catecholamines as a result.is quantitatively insignificant under most conditions During dynamic exercise, the plasma concentrationsof carbohydrate overfeeding. of catecholamines increase linearly with the exer-

cise duration and exponentially with the exerciseIt has been suggested that when food is given inintensity,[111,122] a similar relationship to that observ-the recovery period there is a synergistic interactioned between EPOC and exercise duration and intensi-of food and exercise on energy expenditure. Thety, respectively (figure 2). Thirdly, catecholaminesdifference in opinion as to whether this interactionare important regulators of TG/FA cycling and FAexists[109,110] may be caused by the large intra- andoxidation through stimulation of lipolysis via -inter-individual variations in the thermogenic effectadrenoceptors. Both processes are increased afterof a standard meal. We[44] could not detect anyexercise and may account for a significant part ofmajor interaction effects between food and previousEPOC.aerobic exercise on the VO2.

In early studies on the effect of -adrenoceptorSeveral hormones are potential stimulants of en-blockade on post-exercise VO2, antagonists wereergy expenditure, including insulin, cortisol, thyroidadministered in dogs before an exercise bout.[114,123]hormones, growth hormone, adrenocorticotropicThe results showed that short-term EPOC decreasedhormone (ACTH) and catecholamines. Plasma con-after administration of the non-selective -adre-centrations of growth hormone and ACTH may in-noceptor antagonist, propranolol. The results havecrease during exercise, but no sustained increase inbeen used as an indication of the importance of thesecretion has been found after exercise.[111,112]sympathoadrenal system for EPOC, but sinceMhlum et al.[15] found no changes in plasma insu-propranolol was administered before the start of thelin and free thyroxine during the recovery periodexercise, the physiological effects of the exerciseafter exhausting endurance exercise, and only a tran-bout were different between the control and thesient increase in the plasma concentration of cor-propranolol situation. We have shown that bothtisol. However, in this experiment, food was givenpropranolol and also the selective 1-adrenoceptorin the recovery period, which may have influencedantagonist, atenolol, reduced VO2 to a similar extentthe hormonal response. Although we found thatwhen given intravenously during rest post-exercisevenous plasma insulin concentration rapidly re-and during rest without previous exercise inturned to resting levels after aerobic exercise.[68,69]humans.[68] Hence, there was no effect of -adre-we observed a prolonged depression in arterial plas-noceptor blockade on EPOC after aerobic exercise,ma insulin concentrations after similar exercise.[113]and the results do not support the hypothesis that theThis may be important for the increase in fatprolonged EPOC component is caused by increasedmobilisation during the recovery period. We alsosympathoadrenal activity. A possible 3-adre-found an increased hormonal response to a repeatednoceptor effect on EPOC could not be excluded, butbout of endurance exercise for catecholamines, AC-the 3-adrenoceptor does not seem to be of impor-TH, cortisol and growth hormone.[112]tance for the sympathoadrenal-mediated thermogen-Many authors have suggested that an increasedesis,[121] and thus it seems unlikely that there is any

sympathoadrenal activity may be one of the mech- 3-adrenoceptor effect on EPOC.anisms underlying EPOC, and of the prolongedAn increased sensitivity to catecholamines in thecomponent in particular.[6,15,30,37,114-117] This hypo-

post-exercise period has also been proposed.[7,124]thesis was based on several findings. Firstly, epi-



The hypothesis was built on observations in vitro of expensive, and thus it seems reasonable to speculatethat the energy cost associated with an acceleratedan increased lipolytic response to catecholaminerate of protein synthesis in the post-exercise statestimulation in human gluteal adipocytes[125] and incan contribute to higher energy expenditure. Furthersuprailiac adipocytes[126] removed immediately afterstudies on the importance of increased protein turn-an exercise bout compared with resting samples.over and adaptive protein synthesis on EPOC afterHowever, we have shown[69] that isoprenaline (-different types of exercise are necessary. To ouradrenoceptor agonist) stimulated whole body VO2knowledge, EPOC and protein turnover have notto the same extent during rest with and withoutbeen measured in the same study.previous aerobic exercise. Hence, no increased sen-

sitivity to catecholamines was detected in the post- Both whole body (pulmonary) and muscle VO2exercise period. Also, the lipolytic effect of isopren- should be measured to determine the quantitativealine in abdominal adipose tissue was not increased contribution of muscle metabolism to EPOC. In aafter a prolonged moderate exercise bout, but in- study by Bangsbo et al.,[134] leg VO2 accounted forstead a desensitisation was seen.[113] only one-third of EPOC in the 60-minute recovery

period after exhaustive knee extensor exercise last-Another potential effect of the sympathoadrenaling 3 minutes. Only a minor fraction (26%) of thesystem on EPOC is by stimulation of various pro-excess leg VO2 could be attributed to the oxygencesses during the exercise bout, which are reversedrequirements for resynthesis of substrate. Hence, aslowly after the end of the exercise bout, even iflarge part of the increase in muscle VO2 after exer-there is no increased sympathoadrenal activity incise remained to be explained. Non-exercised mus-this period. During an exercise bout, catecholaminescles may also contribute to EPOC, as both in vitrostimulate both the heart rate, the contractility of theand in vivo studies have shown an increased VO2 inheart, glycogenolysis, gluconeogenesis, and lipoly-inactive muscles perfused with blood high in lac-sis in adipose tissue and in muscles.[127] The cat-tate.[135,136]echolamines also influence the release of other hor-

mones, e.g. insulin and renin. During exercise, The energy efficiency may change during exer-blood flow in some tissues is decreased through - cise,[97,137] and this may be the case also duringadrenoceptors, which may be of importance for recovery from exercise. One potential mechanismblood flow and VO2 in the tissues after exercise. for reduced energy efficiency is the activity of un-Hence, the influence of catecholamines on various coupling proteins (UCP). The expression of UCP3processes during exercise may in turn influence in various organs is consistent with a role in adaptiveEPOC. This is in agreement with the finding of a thermogenesis.[138-140] It remains to be determined ifreduced EPOC when propranolol was administered the cellular milieu during exercise stimulates thein dogs before exercise.[114,123] activity of UCP, and if changes in energy efficiency

during and after exercise may explain part of EPOC.An elevated rate of both protein breakdown andprotein synthesis has been demonstrated in the re- In summary, whereas several of the metaboliccovery period after exercise.[128] An increased whole processes believed to be responsible for the rapidbody protein synthesis has been demonstrated after EPOC component are well known (replenishment of3.75 hours of treadmill running at 50% of oxygen stores in blood and muscle, resynthesis ofVO2max,[129] and after 1 hour of cycling at 75% of ATP and creatine phosphate, lactate removal, andVO2max.[130] Also, an increased muscle protein syn- increased body temperature, circulation and ventila-thesis has been shown after prolonged exercise,[131] tion) the mechanisms underlying the prolongedwhereas studies of the breakdown of muscle protein component are less well understood. An increasedafter such exercise is lacking. Resistance exercise TG/FA cycling, and a shift from carbohydrate to fatalso stimulates protein synthesis in the post-exercise as substrate source, may explain a substantial part ofperiod.[132,133] Synthesis of protein is energetically the prolonged EPOC component after exhaustive


1056 Brsheim & Bahr

submaximal exercise. A minor part may be ex- Acknowledgementsplained by a sustained elevation in circulation, ven-

No sources of funding were used to assist in the prepara-tilation and temperature. No increasedtion of this manuscript. The authors have no conflicts of

sympathoadrenal activity has been found after such interest that are directly relevant to the content of this manu-exercise. Little is known about the mechanisms un- script.derlying EPOC after resistance exercise.

References1. Hill AV, Lupton H. Muscular exercise, lactic acid, and the13. Conclusions

supply and utilization of oxygen. Q J Med 1923; 16: 135-712. Hill AV, Long CNH, Lupton H. Muscular exercise, lactic acid,

and the supply and utilisation of oxygen: parts I-III. Proc R SocMuch of the conflicting reports of EPOC magni- Lond (Biol) 1924; 96: 438-75tude and duration may be resolved if differences in 3. Hill AV, Long CNH, Lupton H. Muscular exercise, lactic acid,

and the supply and utilisation of oxygen: parts IV-VI. Proc Rexercise intensity and duration are taken into ac-Soc Lond (Biol) 1924; 97: 84-138

count. Thus, a small and short-lasting EPOC has 4. Hill AV, Long CNH, Lupton H. Muscular exercise, lactic acid,typically been observed in studies with low exercise and the supply and utilitsation of oxygen: parts VII-VIII. Proc

R Soc Lond (Biol) 1924; 97: 155-76intensity and/or low exercise duration, whereas a5. Margaria R, Edwards HT, Dill OB. The possible mechanisms of

more prolonged and substantial EPOC can be ob- contracting and paying the oxygen debt and the role of lacticacid in muscular contraction. Am J Physiol 1933; 106:served after more strenuous exercise. The relation-689-715

ship between exercise intensity and EPOC magni- 6. Gaesser GA, Brooks GA. Metabolic bases of excess post-tude is curvilinear, whereas at higher intensities exercise oxygen consumption: a review. Med Sci Sports Exerc

1984; 16 (1): 29-43there is a linear relationship between exercise dura-7. Bahr R. Excess postexercise oxygen consumption: magnitude,

tion and EPOC. For resistance exercise, the relation- mechanisms and practical implications. Acta Physiol Scand1992; 144 Suppl. 605: 1-70ship between exercise intensity, duration and EPOC

8. Bullough RC, Gillette CA, Harris MA, et al. Interaction of acuteis less clear, but a more prolonged and substantialchanges in exercise energy expenditure and energy intake on

EPOC has been found after hard versus moderate resting metabolic rate. Am J Clin Nutr 1995 Mar; 61 (3):473-81resistance exercise. Furthermore, the individual ef-

9. Benedict FG, Carpenter TM. The metabolism and energy trans-fects of exercise mode, training status and sex on formations of healthy man during rest. Washington, DC: TheCarnegie Institute, 1910EPOC are unclear, partly because of difficulties in

10. Herxheimer H, Wissing E, Wolff E. Spatwirkungen er-designing appropriate protocols to investigate theseschopfender Muskelarbeit auf den Sauerstoffverbrauch. Z

effects. Still, it appears that trained individuals have Gesamte Exp Med 1926; 51: 916-2811. Edwards HT, Thorndike A, Dill DB. The energy requirementsa more rapid return of VO2 to resting levels after

in strenuous muscular exercise. N Engl J Med 1935; 213:exercising at either the same relative or the same 532-5absolute work rate, but studies after more strenuous 12. Passmore R, Johnson RE. Some metabolic changes following

prolonged moderate exercise. Metabolism 1960; 9: 452-5exercise bouts should be done. Finally, while some13. deVries HA, Gray DE. After effects of exercise upon resting

of the mechanisms underlying the more rapid EPOC metabolic rate. Res Q 1963; 34 (3): 314-2114. Bielinski R, Schutz Y, Jequier E. Energy metabolism during theafter strenuous aerobic exercise are well known, the

postexercise recovery in man. Am J Clin Nutr 1985 Jul; 42 (1):mechanisms underlying the prolonged EPOC com- 69-82ponent are more unclear. The sum of the increased 15. Mhlum S, Grandmontagne M, Newsholme EA, et al. Magni-

tude and duration of excess postexercise oxygen consumptionrate of TG/FA cycling and the shift from carbohy-in healthy young subjects. Metabolism 1986; 35 (5): 425-9drate to fat as substrate source has been suggested to 16. Bahr R, Ingnes I, Vaage O, et al. Effect of duration of exerciseon excess postexercise O2 consumption. J Appl Physiol 1987account for more than half of the prolonged EPOCFeb; 62 (2): 485-90component, but there is still a significant part that

17. Chad KE, Wenger HA. The effect of exercise duration on theremains to be explained. It is hypothesised that exercise and post-exercise oxygen consumption. Can J Sport

Sci 1988; 13: 204-7increased protein turnover and/or a change in energy18. Gore CJ, Withers RT. Effect of exercise intensity and duration

efficiency in the recovery period may explain some on postexercise metabolism. J Appl Physiol 1990; 68 (6):2362-8of this.



19. Gore CJ, Withers RT. The effect of exercise intensity and 39. Sedlock DA. Postexercise energy expenditure following upperduration on the oxygen deficit and excess post-exercise oxygen body exercise. Res Q Exerc Sport 1991 Jun; 62 (2): 213-6consumption. Eur J Appl Physiol 1990; 60: 169-74 40. Sedlock DA. Effect of exercise intensity on postexercise energy

20. Hagberg JM, Mullin JP, Nagle FJ. Effect of work intensity and expenditure in women. Br J Sports Med 1991 Mar; 25 (1):duration on recovery O2. J Appl Physiol 1980; 48: 540-4 38-40

21. Freedman-Akabas S, Colt E, Kissileff HR, et al. Lack of 41. Berg KE. Comparison of energy expenditure in men and womensustained increase in VO2 following exercise in fit and unfit at rest and during exercise recovery. J Sports Med Phys Fitnesssubjects. Am J Clin Nutr 1985; 41: 545-9 1991; 31 (3): 351-6

22. Pacy PJ, Barton N, Webster JD, et al. The energy cost of aerobic 42. Withers RT, Gore CJ, Mackay MH, et al. Some aspect ofexercise in fed and fasted normal subjects. Am J Clin Nutr metabolism following a 35 km road run. Eur J Appl Physiol1985 Nov; 42 (5): 764-8 1991; 63: 436-43

23. Brehm BA, Gutin B. Recovery energy expenditure for steady 43. Bahr R, Sejersted OM. Effect of intensity of exercise on excessstate exercise in runners and nonexercisers. Med Sci Sports postexercise O2 consumption. Metabolism 1991 Aug; 40 (8):Exerc 1986; 18: 205-10 836-41

24. Elliot DL, Goldberg L, Kuehl KS. Does aerobic conditioning 44. Bahr R, Sejersted OM. Effect of feeding and fasting on excesscause a sustained increase in the metabolic rate? Am J Med Sci postexercise oxygen consumption. J Appl Physiol 1991 Dec;1988; 296: 249-51 71 (6): 2088-93

25. Turley KR, McBride PJ, Wilmore JH. Resting metabolic rate 45. Bahr R, Grnnerd O, Sejersted OM. Effect of supramaximalmeasured after subjects spent the night at home vs at a clinic. exercise on excess postexercise O2 consumption. Med SciAm J Clin Nutr 1993 Aug; 58 (2): 141-4 Sports Exerc 1992 Jan; 24 (1): 66-71

26. Bullough RC, Melby CL. Effect of inpatient versus outpatient 46. Elliot DL, Goldberg L, Kuehl KS. Effect of resistance trainingmeasurement protocol on resting metabolic rate and respira- on excess post-exercise oxygen consumption. J Appl Sport Scitory exchange ratio. Ann Nutr Metab 1993; 37 (1): 24-32 Res 1992; 6 (2): 77-81

27. Maresh CM, Abraham A, De Souza MJ, et al. Oxygen con- 47. Sedlock DA. Post-exercise energy expenditure after cycle er-sumption following exercise of moderate intensity and dura- gometer and treadmill exercise. J Appl Sport Sci Res 1992; 6tion. Eur J Appl Physiol 1992; 65: 421-6 (1): 19-23

28. Knuttgen HG. Oxygen debt after submaximal physical exercise. 48. Donelly K, McNaughton L. The effects of two levels of caffeineJ Appl Physiol 1970; Nov 29 (5): 651-7 ingestion on excess postexercise oxygen consumption in un-29. Segal SS, Brooks GA. Effects of glycogen depletion and work trained women. Eur J Appl Physiol Occup Physiol 1992; 65

load on postexercise O2 consumption and blood lactate. J Appl (5): 459-63Physiol 1979; 47: 514-21 49. Brockman L, Berg K, Latin R. Oxygen uptake during recovery30. Hermansen L, Grandmontagne M, Mhlum S, et al. Post- from intense intermittent running and prolonged walking. Jexercise elevation of resting oxygen uptake: possible mechan- Sports Med Phys Fitness 1993 Dec; 33 (4): 330-6isms and physiological significance. In: Marconnet P, 50. Kaminsky LA, Whaley MH. Effect of interval-type exercise onPoortmans J, Hermansen L, editors. Medicine and sport

excess postexercise oxygen consumption (EPOC) in obese andscience. Vol. 17. Basel: Karger, 1984: 119-29normal-weight women. Med Exerc Nutr Health 1993; 2:31. Chad KE, Wenger HA. The effects of duration and intensity on 106-11

the exercise and post-exercise metabolic rate. Aust J Sci Med51. Smith J, McNaughton L. The effects of intensity of exercise onSport 1985; 17: 14-8

excess postexercise oxygen consumption and energy expendi-32. Devlin JT, Horton ES. Potentiation of the thermic effect of ture in moderately trained men and women. Eur J Appl Physiolinsulin by exercise: differences between lean, obese, and 1993; 67: 420-5noninsulin-dependent diabetic men. Am J Clin Nutr 1986; 43:

52. Neary JP, Docherty D, Wenger HA. Post-exercise metabolic884-90rate is influenced by elevated core temperature. Aust J Sci Med33. Sedlock D, Fissinger JA, Melby CL. Effect of exercise intensity Sport 1993; 25 (2): 43-7

and duration on postexercise energy expenditure. Med Sci53. Frey GC, Byrnes WC, Mazzeo RS. Factors influencing excessSports Exerc 1989; 21: 662-6

postexercise oxygen consumption in trained and untrained34. Chad K, Quigley B. The effects of substrate utilization, manipu-women. Metabolism 1993; 42 (7): 822-8lated by caffeine, on post-exercise oxygen consumption in

54. Brsheim E, Bahr R, Hansson P, et al. Effect of -adrenoceptoruntrained female subjects. Eur J Appl Physiol 1989; 59: 48-54blockade on post-exercise oxygen consumption. Metabolism35. Poehlman ET, LaChance P, Tremblay A, et al. The effect of1994; 43 (5): 565-71prior exercise and caffeine ingestion on metabolic rate and

55. Thomas TR, Londeree BR, Lawson DA. Prolonged recoveryhormones in young adult males. Can J Physiol Pharmacol 1989from eccentric versus concentric exercise. Can J Appl PhysiolJan; 67 (1): 10-61994; 19 (4): 441-5036. Kaminsky LA, Padjen S, LaHam-Saeger J. Effect of split exer-

56. Sedlock DA. Fitness level and postexercise energy expenditure.cise sessions on excess post-exercise oxygen consumption. BrJ Sports Med Phys Fitness 1994 Dec; 34 (4): 336-42J Sports Med 1990; 24 (2): 95-8

57. Gilette CA, Bullough RC, Melby C. Postexercise energy expen-37. Bahr R, Hansson P, Sejersted OM. Triglyceride/fatty acid cy-diture in response to acute aerobic or resistive exercise. Int Jcling is increased after exercise. Metabolism 1990; 39 (9):Sport Nutr 1994; 4: 347-60993-9

38. Chad KE, Quigley BM. Exercise intensity: effect on post- 58. Quinn TJ, Vroman NB, Kertzer R. Postexercise oxygen con-exercise O2 uptake in trained and untrained women. J Appl sumption in trained females: effect of exercise duration. MedPhysiol 1991; 70: 1713-9 Sci Sports Exerc 1994 Jul; 26 (7): 908-13


1058 Brsheim & Bahr

59. Harms CA, Cordain L, Stager JM, et al. Body fat mass affects 77. Friden J, Sjostrom M, Ekblom B. Myofibrillar damage follow-postexercise metabolism in males of similar lean body mass. ing intense eccentric exercise in man. Int J Sports Med 1983; 4:Med Exerc Nutr Health 1995; 4: 33-9 170-6

78. Kolkhorst FW, Londeree BR, Thomas TR. Effect of consecutive60. Dawson B, Straton S, Randall N. Oxygen consumption duringexercise days of jogging or cycling on the resting metabolicrecovery from prolonged submaximal cycling below the anaer-rate and nitrogen balance. J Sports Med Phys Fitness 1994; 34:obic threshold. J Sports Med Phys Fitness 1996 Jun; 36 (2):343-5077-84

79. Melby CL, Tincknell T, Schmidt WD. Energy expenditure61. Short KR, Wiest JM, Sedlock DA. The effect of upper bodyfollowing a bout of non-steady state resistance exercise. Jexercise intensity and duration on post-exercise oxygen con-Sports Med Phys Fitness 1992; 32: 128-35sumption. Int J Sports Med 1996 Nov; 17 (8): 559-63

80. Murphy E, Schwarzkopf R. Effects of standard set and circuit62. Trost S, Wilcox A, Gillis D. The effect of substrate utilization,weight training on excess post-exercise oxygen consumption. Jmanipulated by nicotinic acid, on excess postexercise oxygenAppl Sport Sci Res 1992; 6 (2): 88-91consumption. Int J Sports Med 1997; 18 (2): 83-8

81. Melby C, Scholl C, Edwards G, et al. Effect of acute resistance63. Phelain JF, Reinke E, Harris MA, et al. Postexercise energyexercise on postexercise energy expenditure and resting meta-expenditure and substrate oxidation in young women resultingbolic rate. J Appl Physiol 1993 Oct; 75 (4): 1847-53from exercise bouts of different intensity. J Am Coll Nutr 1997

82. Olds TS, Abernethy PJ. Postexercise oxygen consumption fol-Apr; 16 (2): 140-6lowing heavy and light resistance exercise. J Strength Cond64. Laforgia J, Withers RT, Shipp NJ, et al. Comparison of energyRes 1993; 7: 147-52expenditure elevations after submaximal and supramaximal

83. Haltom RW, Kraemer RR, Sloan RA, et al. Circuit weightrunning. J Appl Physiol 1997; 82 (2): 661-6training and its effects on excess postexercise oxygen con-65. Short KR, Sedlock DA. Excess postexercise oxygen consump-sumption. Med Sci Sports Exerc 1999 Nov; 31 (11): 1613-8tion and recovery rate in trained and untrained subjects. J Appl

84. Dolezal BA, Potteiger JA, Jacobsen DJ, et al. Muscle damagePhysiol 1997; 83 (1): 153-9and resting metabolic rate after acute resistance exercise with66. Burleson Jr MA, OBryant HS, Stone MH, et al. Effect ofan eccentric overload. Med Sci Sports Exerc 2000 Jul; 32 (7):

weight training exercise and treadmill exercise on post-exer- 1202-7cise oxygen consumption. Med Sci Sports Exerc 1998 Apr; 30

85. Osterberg KL, Melby CL. Effect of acute resistance exercise on(4): 518-22postexercise oxygen consumption and resting metabolic rate in67. Almuzaini KS, Potteiger JA, Green SB. Effects of split exercise young women. Int J Sport Nutr Exerc Metab 2000 Mar; 10 (1):

sessions on excess postexercise oxygen consumption and rest- 71-81ing metabolic rate. Can J Appl Physiol 1998 Oct; 23 (5): 86. Binzen CA, Swan PD, Manore MM. Postexercise oxygen con-433-43sumption and substrate use after resistance exercise in women.68. Brsheim E, Bahr R, Hstmark AT, et al. Effect of -adre- Med Sci Sports Exerc 2001 Jun; 33 (6): 932-8

noceptor blockade on post-exercise oxygen consumption and 87. Thornton MK, Potteiger JA. Effects of resistance exercise boutstriglyceride/fatty acid cycling. Metabolism 1998; 47 (4):of different intensities but equal work on EPOC. Med Sci439-48 Sports Exerc 2002 Apr; 34 (4): 715-22

69. Brsheim E, Bahr R, Knardahl S. Effect of -adrenoceptor 88. Schuenke MD, Mikat RP, McBride JM. Effect of an acutestimulation on oxygen consumption and triglyceride/fatty acid period of resistance exercise on excess post-exercise oxygencycling after exercise. Acta Physiol Scand 1998; 164: 157-66

consumption: implications for body mass management. Eur J70. Matsuo T, Saitoh S, Suzuki M. Effects of the menstrual cycle on Appl Physiol 2002 Mar; 86 (5): 411-7

excess postexercise oxygen consumption in healthy young 89. Williamson DL, Kirwan JP. A single bout of concentric resis-women. Metabolism 1999 Mar; 48 (3): 275-7 tance exercise increases basal metabolic rate 48 hours after

71. Lee YS, Ha MS, Lee YJ. The effects of various intensities and exercise in healthy 59-77-year-old men. J Gerontol A Biol Scidurations of exercise with and without glucose in milk inges- Med Sci 1997 Nov; 52 (6): M352-5tion on postexercise oxygen consumption. J Sports Med Phys 90. Hagberg JM, Hickson RC, Ehsani AA, et al. Faster adjustmentFitness 1999 Dec; 39 (4): 341-7 to and recovery from submaximal exercise in the trained state.

72. Fukuba Y, Yano Y, Murakami H, et al. The effect of dietary J Appl Physiol 1980 Feb; 48 (2): 218-24restriction and menstrual cycle on excess post-exercise oxygen 91. Girandola RN, Katch FI. Effects of physical conditioning onconsumption (EPOC) in young women. Clin Physiol 2000 changes in exercise and recovery O2 uptake and efficiencyMar; 20 (2): 165-9 during constant-load ergometer exercise. Med Sci Sports 1973;

73. Bouchard C, Rankinen T. Individual differences in response to 5 (4): 242-7regular physical activity. Med Sci Sports Exerc 2001 Jun; 33 (6 92. Solomon SJ, Kurzer MS, Calloway DH. Menstrual cycle andSuppl.): S446-51 basal metabolic rate in women. Am J Clin Nutr 1982 Oct; 36

74. Bahr R, Opstad PK, Medb JI, et al. Strenuous prolonged (4): 611-6exercise elevates resting metabolic rate and causes reduced 93. Meijer GA, Westerterp KR, Saris WH, et al. Sleeping metabolicmechanical efficiency. Acta Physiol Scand 1991 Apr; 141 (4): rate in relation to body composition and the menstrual cycle.555-63 Am J Clin Nutr 1992 Mar; 55 (3): 637-40

75. Newham DJ, McPhail G, Mills KR, et al. Ultrastructural 94. Bisdee JT, James WP, Shaw MA. Changes in energy expendi-changes after concentric and eccentric contractions of human ture during the menstrual cycle. Br J Nutr 1989 Mar; 61 (2):muscle. J Neurol Sci 1983 Sep; 61 (1): 109-22 187-99

76. Friden J, Lieber RL. Eccentric exercise-induced injuries to 95. Hessemer V, Bruck K. Influence of menstrual cycle on thermo-contractile and cytoskeletal muscle fibre components. Acta regulatory, metabolic, and heart rate responses to exercise atPhysiol Scand 2001 Mar; 171 (3): 321-6 night. J Appl Physiol 1985 Dec; 59 (6): 1911-7



96. Webb P. 24-hour energy expenditure and the menstrual cycle. 117. Sagnol M, Claustre J, Pequignot JM, et al. Catecholamines andAm J Clin Nutr 1986 Nov; 44 (5): 614-9 fuels after an ultralong run: persistent changes after 24-h

recovery. Int J Sports Med 1989 Jun; 10 (3): 202-697. Bangsbo J, Gollnick PD, Graham TE, et al. Anaerobic energy118. Cori CF, Buchwald KW. Effect of continuous injection ofproduction and O2 deficit-debt relationship during exhaustive

epinephrine on the carbohydrate metabolism, basal metabol-exercise in humans. J Physiol 1990; 422: 539-59ism and vascular system of normal man. Am J Physiol 1930;98. Wolfe RR, Klein S, Carraro F, et al. Role of triglyceride-fatty95: 71-8acid cycle in controlling fat metabolism in humans during and

119. Webber J, Macdonald IA. Metabolic actions of catecholaminesafter exercise. Am J Physiol 1990; 258 (21): E382-9in man. Baillieres Clin Endocrinol Metab 1993; 7 (2): 393-41399. Miyoshi H, Schulman GI, Peters EJ, et al. Hormonal control of

120. Blaak EE, Saris WHM, van Baak MA. Adrenoceptor subtypessubstrate cycling in humans. J Clin Invest 1988; 81: 1545-55mediating catecholamine-induced thermogenesis in man. Int J100. Wolfe RR, Peters EJ. Lipolytic response to glucose infusion in Obes 1993; 17 (3): S78-81human subjects. Am J Physiol 1987; 252 (15): E218-23

121. Blaak EE, van Baak MA, Kempen KPG, et al. Role of - and -101. Chaliss RAJ, Arch JRS, Newsholme EA. The rate of substrateadrenoceptors in sympathetically mediated thermogenesis. Am

cycling between fructose 6-phosphate and fructose J Physiol 1993; 264 (27): E11-71,6-bisphosphate in skeletal muscle. Biochem J 1984; 221: 122. Kjr M, Secher NH, Galbo H. Physical stress and catecho-153-61 lamine release. Baillieres Clin Endocrinol Metab 1987; 1:102. Wolfe RR, Herndon DN, Jahoor F, et al. Effect of severe burn 279-89

injury on substrate cycling by glucose and fatty acids. N Engl J 123. Cain SM. Exercise O2 debts of dogs at ground level and atMed 1987; 317: 403-8altitude with and without -block. J Appl Physiol 1971; 30:

103. Stein TP, Rumpler WV, Leskiw MJ, et al. Effect of reduced 838-43dietary intake on energy expenditure, protein turnover, and 124. Bahr R, Hstmark AT, Newsholme EA, et al. Effect of exerciseglucose cycling in man. Metabolism 1991; 40 (5): 478-83 on recovery changes in plasma levels of FFA, glycerol, glu-

104. Weber JM, Klein SE, Wolfe RR. Role of the glucose cycle in cose and catecholamines. Acta Physiol Scand 1991 Sep; 143control of net glucose flux in exercising humans. J Appl (1): 105-15Physiol 1990; 68: 1815-9 125. Wahrenberg H, Engfeldt P, Bolinder J, et al. Acute adaptation

105. Wolfe RR, Klein S, Herndon DN, et al. Substrate cycling in in adrenergic control of lipolysis during physical exercise inthermogenesis and amplification of net substrate flux in human humans. Am J Physiol 1987; 253 (16): E383-90volunteers and burned patients. J Trauma 1990; 30 (12): 6-9 126. Savard R, Despres JP, Marcotte M, et al. Acute effects of

endurance exercise on human adipose tissue metabolism. Met-106. Flatt JP. The biochemistry of energy expenditure. In: Bray GA,abolism 1987; 36 (5): 480-5editor. Recent advances in obesity research II. London: New-

man, 1978: 211-28 127. Landsberg L, Young JB. Catecholamines and the adrenal me-dulla. In: Wilson JD, Foster DW, editors. Textbook of endocri-107. Hellerstein MK, Schwarz JM, Neese RA. Regulation of hepaticnology. 8th ed. Philadelphia (PA): WB Saunders Company,de novo lipogenesis in humans. Annu Rev Nutr 1996; 16:1992: 621-81523-57

128. Viru A. Postexercise recovery period: carbohydrate and protein108. McDevitt RM, Bott SJ, Harding M, et al. De novo lipogenesismetabolism. Scand J Med Sci Sports 1996; 6: 2-14during controlled overfeeding with sucrose or glucose in lean

129. Rennie MJ, Edwards RHT, Krywawych S, et al. Effect ofand obese women. Am J Clin Nutr 2001 Dec; 74 (6): 737-46exercise on protein turnover in man. Clin Sci 1981; 61: 627-39109. Richard D, Rivest S. The role of exercise in thermogenesis and

130. Devlin JT, Brodsky I, Scrimgeour A, et al. Amino acid metabol-energy balance. Can J Physiol Pharmacol 1989; 67 (4): 402-9ism after intense exercise. Am J Physiol 1990; 258: E249-55110. Poehlman ET, Horton ES. The impact of food intake and 131. Carraro F, Stuart CA, Hartl WH, et al. Effects of exercise and

exercise on energy expenditure. Nutr Rev 1989; 47: 129-37recovery on muscle protein synthesis in human subjects. Am J

111. Galbo H. Hormonal and metabolic adaptation to exercise. Stutt- Physiol 1990; 259: E470-6gart: Georg Thieme Verlag, 1983 132. Biolo G, Maggi SP, Williams BD, et al. Increased rates of

112. Rnsen O, Haug E, Pedersen BK, et al. Increased neuroendo- muscle protein turnover and amino acid transport after resis-crine response to a repeated bout of endurance exercise. Med tance exercise in humans. Am J Physiol 1995; 268 (31):Sci Sports Exerc 2001 Apr; 33 (4): 568-75 E514-20

113. Brsheim E, Lonnroth P, Knardahl S, et al. No difference in the 133. Phillips SM, Tipton KD, Aarsland A, et al. Mixed musclelipolytic response to -adrenoceptor stimulation in situ but a protein synthesis and breakdown after resistance exercise indelayed increase in adipose tissue blood flow in moderately humans. Am J Physiol 1997 Jul; 273 (1 Pt 1): E99-107obese compared with lean men in the postexercise period. 134. Bangsbo J, Gollnick PD, Graham TE, et al. Substrates forMetabolism 2000 May; 49 (5): 579-87 muscle glycogen synthesis in recovery from intense exercise in

114. Barnard RJ, Foss ML. Oxygen debt: effect of beta-adrenergic man. J Physiol 1991; 434: 423-40blockade on the lactacid and alactacid components. J Appl 135. Chin ER, Lindinger MI, Heigenhauser GJF. Lactate metabolismPhysiol 1969; 27: 813-6 in inactive skeletal muscle during lactacidosis. Am J Physiol

115. Chapler CK, Stainsby WN, Gladden LB. Effect of changes in 1991; 261: R98-105blood flow, norepinephrine and pH on oxygen uptake by 136. Bangsbo J, Graham T, Johansen L, et al. Muscle lactateresting skeletal muscle. Can J Physiol Pharmacol 1980; 58: metabolism in recovery from intense exhaustive exercise: im-93-6 pact of light exercise. J Appl Physiol 1994; 77 (4): 1890-5

116. Gladden LB, Stainsby WN, MacIntosh BR. Norepinephrine 137. Scott CB. Re-interpreting anaerobic metabolism: an argumentincreases canine skeletal muscle VO2 during recovery. Med for the application of both anaerobic glycolysis and excessSci Sports Exerc 1982; 14 (6): 471-6 post-exercise oxygen consumption (EPOC) as independent


1060 Brsheim & Bahr

sources of energy expenditure. Eur J Appl Physiol 1998; 77:200-5

Correspondence and offprints: Dr Elisabet Brsheim, De-138. Vidal-Puig A, Solanes G, Grujic D, et al. UCP3: an uncouplingpartment of Surgery, Metabolism Unit, Shriners Hospitalsprotein homologue expressed preferentially and abundantly in

skeletal muscle and brown adipose tissue. Biochem Biophys for Children/UTMB, 815 Market Street, Galveston, TXRes Commun 1997; 235: 79-82 77550, USA.

139. Fleury C, Neverova M, Collins S, et al. Uncoupling protein-2: a E-mail: [email protected] gene linked to obesity and hyperinsulinemia. Nat Genet1997; 15 (3): 269-72

140. Boss O, Samec S, Paoloni-Giacobino A, et al. Uncouplingprotein-3: a new member of the mitochondrial carrier familywith tissue-specific expression. FEBS Lett 1997; 408: 39-42


Contents 1037Abstract 10371. Excess Post-Exercise Oxygen Consumption (EPOC) 10382. Early Studies on EPOC 10393. Methodological Considerations 10394. Effect of Intensity and Duration of Aerobic Exercise on EPOC 10405. Effect of Split Exercise Sessions on EPOC 10476. Effect of Supramaximal Exercise on EPOC 10487. Effect of Aerobic Exercise Mode on EPOC 10488. Effect of Resistance Exercise on EPOC 10499. Effect of Training Status on EPOC 105110. Effect of Sex on EPOC 105211. Possible Mechanisms for the Rapid EPOC Component 105312. Possible Mechanisms for the Prolonged EPOC Component 105313. Conclusions 1056Acknowledgements 1056References 1056Correspondence 1060Email 1060

Effect of Exercise Intensity Duration No Epoc

Documents

Transcript of Effect of Exercise Intensity Duration No Epoc