The Influence of Frequency, Intensity, Volume and Mode of Strength Trainirg On Whole Muscle Cross-SectionaAl Rea in Humans

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Sports Med 2m7; 37 (3)i 22$264

0 1\2- | &2 7 I mi-O225t 5A4.95| O
@ 2@7 Adb Dqto Informqlbn
BV. Afl dghts reseNed
The Influence of Frequency,Intensity,

Volume and Mode of Strength
Trainirg on Whole Muscle
Cross-SectionalArea in Humans
Mathias Wernbom,rlesperAugustssonr,2and RolandThomel,2
1 Lundberg Laboratory for Human Muscle Function and Movement Analysis, Department of
Orthopaedics,SahlgrenskaUniversity Hospital, GteborgUniversity, Gteborg, Sweden
2 SportRehabPhysical Therapy and Sports Medicine Clinic, Gteborg,Sweden
Confenls
A bs trq c t.
...,.,226
l. Me th o d s
..........227
I.l L i l e ro tu re S e o rc h
.,.,,227
1.2 Clossificotion
. . .228
of MuscleActions,TroiningMethodsond Modolities
1.3 Quontificqtion
Frequency,
Intensity
ond Volume
.........229
of Exercise
1 .4 C l o s s i fi c o l l o fnT ro i n i nSgto tus
....,..230
.l.5
Areq (CSA).......230
Cqlculotionof Chongesond RoteofChongesin MuscleCross-Sectionql
2. R e s u l ts P ol :Qu
rt q d ri c e p s S tu d i es
....,...23.l
2.1 QuodricepsStudies:DynomicExternqlResislonce
. . . . 231
2 .1 .1 L e n g tho f T ro i n i nPe
g ri od,
A verogeIncreose
i n C S Aond C S AP erD oy.,.
,..,.232
2 .1 .2 R o te o fGo i n i n C S A:
MenversusW omen
......232
2 .1 .3 F re q u e n c y...
.,.,,....232
2 .1 ,4 In te n s i ty
..,,..,.232
..,,..,.232
2 .1 .5 V o l u me
2 .2 Qu o d ri c e p s Stu d i A
e sc :c o m modoti ng
R esi stonce......
...,.233
g ri od,A verogel ncreqsei nC S AC
ond
2 .2 .1 L e n g th o fT ro i n i nPe
S A P erD oy...
.....233
2 .2 ,2 F re q u e n c y...
.....,. ..23 3
2 .2 .3 V e l o c i ty
........234
2 .2 ,AIo td te
.....,...23A
2 .2 .5 V o tu m e
........2g4
2 ,2 ,6 l o tq l D u ro ti o n P e rSessi on
...,..235
2 .2 ,7 T i m e -T o rq u
Pro
e d u cPerS
l
essi on
......,235
2.3 QuodricepsStudies:lsometricResistonce.
. . . . 235
g e ri od,A verqge
Increosei nC S A qnd
2 .3 .1 L e n g tho f T ro i n i nP
C S A P erD oy,..
.....235
2 .3 .2 F re q u e n c y ...
.........235
2 .3 .3 In te n s i fy
,.....,.235
2 .3 .4 Vo l u m e
........235
2.4 QuodricepsStudies:CombinedStrengthond EnduronceTroining
. . .236
2 .4 .1 R o teo f Go i ni n C SA C
: ombi nedTroi niversus
ng P ureS trengthTroi ni ng ..,.....236
2 .4 .2 L e n g th o fT ro i n i n g Peri odondl ncreosei nC, S A
.,..,.,236
. , . ,236
2.5 QuodricepsStudies:All VoluntoryTroiningModes
......,236
2 .5 .1 L e n g th o fT ro i n i n g Peri odqndl ncreosei nC. S A
2 .6 Qu q d ri c e p s S tu d i Stre
C ountermeosure
D uri ngU nl oodi ng...........,.236
e s : n gthTroi ni ngoso
Rrvrew
Anncu
SportsMed 2m7;37(3)r22S264
0 I I 2-l42l07/@3-0225t544.95t0
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gV. All rights reserysd.
The Influence of Frequency,Intensity,

Volume and Mode of Strength
Trainirg on Whole Muscle
Cross-SectionalArea in Humans
Mathias Wernbom,rlesperAugustssonl,2and RolandThomel,2
1 Lundberg Laboratory for Human Muscle Function and Movement Analysis, Department of
Orthopaedics,SahlgrenskaUniversity Hospital, GteborgUniversity, Gteborg, Sweden
2 SportRehabPhysical Therapy and Sports Medicine Clinic, Gteborg,Sweden
Conlenls
A bs tro c t.
......226
l. Me th o d s
........,.227
l .l L i l e ro tu re S e o rc h
.....227
1.2 Clossificotion
of MuscleActions,TroiningMethodsond Modolities
. . .228
qnd Volume
1.3 Quontificqtion
Frequency,
Intensity
of Exercise
...,,....229
1 .4 C l o s s i fi c o tio fnT ro i n i nSgto tus
.......230
1.5 Cqlculqtionof ChongesondRoteofChongesinMuscleCros-Sectionol
Areo (CSA).......230
2. R e s u l ts P ol :Qu
rt o d ri c e p s S tu d i es
........231
2.I QuodricepsStudieslDynomicExlernolResislonce
. . . . 231
2 .1 .l L e n g th o fT ro i n i n g Peri od,A verqgeIncreosei nC S A qndC S A P erD oy... .....232
2.1.2 Roteof Goin in CSA:Men versusWomen
. . . . . .232
2 .1 .3 F re q u e n c y .,.
.........232
2 .1 ,4 In te n s i ty
,.,.....232
2 .1 .5 V o l u me
......,.232
2 .2 Qu o d ri c e p s S tu d i A
e sc:c o m modoti ng
R esi stonce......
.....233
g ri od,A verqgel ncreosei nC S AC
ond
2 .2 .1 L e n g rth oTf ro i n i nPe
S A P erD oy...
...,.233
2 .2 ,2 F re q u e n c y.,.
....... ..23 3
2 .2 .3 V e l o c i ty
........234
2 .2 .4 l o r u e
.........234
2 .2 .5 V o tu m e
........234
2 .2 .6 l o to l D u ro ti o n P e rSessi on
......235
cl
essi on
2 .2 ,7l i me -Io rq u e P ro d u PerS
.......235
2.3 QuodricepsStudies:lsometricResistonce.
. . . . 235
g e r i od,A veroge
IncreoseiC
n S A ondC S A P erD oy,..
.,...235
2 .3 .2 F re q u e n c y...
.........235
2 .3 .3 In te n s i ty
...,....235
2 .3 .4 Vo l u m e
....,...235
2.4 QuodricepsStudies:CombinedStrengthond EnduronceTroining
. , .236
2.4.1 Rqteof Goinin CSA:CombinedTroining
versus
PureStrength
Troining
...,,...236
2 .4 .2 L e n g l ho f T ro i n i n g Peri odond
Increosei nC S A .
.......236
2,5 QuodricepsSludies:All VoluntoryTroiningModes
. . . .236
..,....236
g e ri odond Increosei nC S.A
2 .6 Qu q d ri c e p s S tu d i Stre
Troi ni ng
os o C ountermeosure
D uri ngU nl oodi ng.......,..,..236
e s : n gth
Strength Training and Muscle Cross-SectionalArea
227
Strengthtraining has becomeincreasinglypopular in recent decades.Whereaspreviously strength

training had been used by a few selectedathletesto
improve their strengthand size, it is now an important componentin training for most sportsas well as
for injury preventionand rehabilitaliqn.tr-31
Quantification of the dose-response
relationshipsbetween
the training variables(e.g. intensity,frequencyand
volume) and the outcome (e.g. strength,power and
hypertrophy)is fundamentalfor the properprescription of resistancetraining.talSeveral meta-analysssta-8l
sr6 numerouslsviswstl'3'e-lllhavedealt with
various aspectsof optimising strength; however,
with the exceptionof a paperby Fry,ttzlfew systematic reviews have focused on the issue of how to
train specifically for muscle hypertrophy. The review of Frytr2ldiscussedthe role of training intensity on hypertrophy as measuredby increasesin muscle fibre area (MFA).
training periodsor training sessions.aimed

at stimulating maximum hypertrophyby high volume and
moderateloads are followed by periodsor sessions
aimedat increasingmaximum strengthby moderate
volume and heavy loads.The latter type of workout
presumably acts by optimising neural adaptations,trrlalthoughmarked hypertrophycan also occur if the volume is sufficient.tl2'15'l8l
Scanningmethodssuch as MRI and CT are regarded as the gold standardfor assessingwhole
musclesize.tle'201
To our knowledge,no systematic
review has been published that has analysedthe
impact of several important training variables
suchas frequency,intensityand volume on changes
in muscleareaor volume as measuredby scanning
methods.Such a review could provide evidencebased guidelines for the prescription of strength
training for increasingmuscle mass. Establishing
efficient models of strengthtraining for hypertrophy
in humanscould also be of value for the studyof the
Implicit in many articles in the literature are the
physiologicalmechanismsof the hypertrophyprofollowing assumptions:(i) training for strengthand
cess.Therefore,the purposeof the presentstudy was
training for hypertrophy is essentiallyone and the
relationshipsfor the develto identify dose-response
same thing; and (ii) the programmethat yields the
opment of muscle hypertrophy by calculating the
largestincreasesin strengthalso resultsin the larg- ratesand magnitudesof increasesin musclecrossest increasesin musclemass.Theseassumptionsare sectionalarea(CSA) or muscle volume inducedby
not necessarilytrue in all situations.For example, varying levels of frequency,intensity and volume,
studiesby Choi et al.tl3land Masudaet al.llalshowed as well as by different modesof strengthtraining.
smallerincreasesin one repetitionmaximum (lRM)
and isometric strength, but greater increases in
l. Melhods
quadriceps muscle area (as measuredby magnetic
resonanceimaging IMRII) and MFA after a typical
l.l LiterotureSeorch
moderate-loadbodybuilding regimen when compared with a high-intensity powerlifting proComputer searchesin the MEDLINE/PubMed,
gramme. Schmidtbleicherand Buehrletrslshowed SportDiscus@and CINAHL@ databaseswere pergreater increasesfor triceps brachii muscle area (as formed for articlesfrom 1970.when the first trainmeasuredby CT scanning)for a group that trained ing study using scanning techniquesto evaluate
with 3 sets of 12 repetitionsatTOVoof IRM when changes in anatomical muscle CSA was pubcompared with a group that trained with 7 sets of
lished.l2ll In addition. hand searchesof relevant
l-3 repetitions at 90-l0OVo of lRM, while the journals and books as well as the referencelists of
strengthincreasesfor the groups were similar. Thus, articles already obtained were performed. The data
it is apparentfrom these and other studies that the reviewedin this article was accumulatedas a result
training prescription for hypertrophy may differ
of literature searchesconducted for this and other
somewhat from the prescription for maximum projects over a period of several years. The last
strength.This observationhas been taken into ac- searchwas performed on 3 December 2006. As the
*6"t"
searchprogressed,it quickly became apparentthat
count in variousmodelsof periodisation,tl6'17l
@ 2m7 Adis Dqto Intormotion BV. All rights reserued.
Sports Med 2@7; 37 (3)
228
the quadriceps was by far the most studied muscle

group in humans undergoingstrengthtraining and
that the elbow flexors (biceps brachii and brachialis)
were the secondmost studied muscle group. Currently, these are probably the only muscle groups
that allow for meaningful evaluation of dose-responsetrends. Therefore, the present review is focusedon the human quadricepsand elbow flexors.
Wernbomet al.
1.2Clossificotionof Muscle Actions,Troining

Methods ond Modolities
For the purposesof this review, the terminology

of Knuttgenand Komit22lwas adoptedfor the classification of muscle actions. Accordingly, exercise
can be classifiedas eitherstatic(involving isometric
muscle actions) or dynamic (involving concentric
and/oreccentricmuscleactions).During an isometCriteria for inclusion were as follows:
ric muscleaction,the muscledevelopsforce but no
l. Studies must have examined the effects of
external movement occurs and the length of the
strengthtraining on anatomical muscle areaor mus- muscle-tendoncomplex does not change.During a
cle volume by a scanningmethod (i.e. MRI, CT or concentricaction, the muscle producesforce while
ultrasound[UL]).
shortening. An eccentric muscle action refers to a
2. Studiesmust have been conductedon healthy and situation where the muscle produces force while
lengthening.
uninjured participantsbetween 18 and 59 years of
In resistanceexercise,training methodsor moage.
dalities are often classified according to the type of
3. Sufficientdatato calculatechangesin musclearea
resistanceused. In his l98l review, arguably the
or volume must have been reported.
first systematic look at strength training, tr1ftstel
4. Sufficient information regarding the training varidivided the training modes into three main categoables (in particular: frequency, intensity and vol- ries: (i) isotonic; (ii) isokinetic; and (iii) isometric.
ume) and the type and mode of exerciseemployed Isometrictraining obviously involves isometricacmust have been reported to allow for replication of
tions.Isotonictrainingrefersto dynamicexercisein
which the muscle(s)exertsa constanttension.t22l
In
the study.
various
physiology,
textbooks
of
an
isotonic
muscle
Exclusion criteria were as follows:
action is often illustrated by an isolated muscle that
L The subjectsreceivedsupplements(e.g. creatine is shorteningor lengtheningagainst
a constantload,
monohydrate,amino acids, proteins) or anabolic thus developinga constantforce. This is not true in
hormones and/or growth factors that could poten- the intact muscle of a personperforming an exercise
tially have influenced the neuromuscularadapta- becauseofbiomechanicalfactorssuchaschangesin
tions to strengthtraining.Suchstudy groups(but not the lengthsof the lever armsof the muscleandof the
necessarilyother groups from the samestudy) were resistance.and also the accelerationsand deceleraexcludedfrom the current review.
tionsthat occurduring dynamicexercise.Thus,even
2. The subjectswere in negativeenergybalance(i.e. if the externalresistanceremainsconstant,the muscle will not develop a constantlevel of forae.l2z'231
on a weight-lossdiet).
For this reason,the term 'isotonic' is often replaced
3. The data from the study have been published
by the term 'dynamic constantexternalresistance'
before. However, sometimes it was necessaryto
(DCER).r2xIn DCER exercise,the absoluteload is
collect data from several different papers from the
constantthroughoutthe movement,as when lifting a
samestudy.
dumbell.Furtherexamplesof DCER would be sim4. Only part of the musclegroup (e.g.vastuslateral- ple cable pulley systems with no lever arms and
is) was scanned.Alternatively,only datafor the total machineswith circle shapedcams.The term 'variamuscle CSA of the limb (e.g. total thigh muscle ble resistance'is used when the resistancethrougharea) was reported without specifying CSAs for the out the rangeof motion is varied,for exampleby an
individual muscles(e.g. quadriceps).
irregularly shapedcamwheel or a lever nryl.tz3l15"
O 2007 Adis Dolo Informotion BV. All rlghts reseNed.
SportsMed 2@7;37 (3)
229
idea with variable resistanceis to closely match the

resistancewith the strengthof the subjectthroughout the rangeof motion.t24lHowever,becausemany
dynamic training studiesdid not specify if the resistance was constant or variable, no distinction between thesesubcategorieswas made in the present
review. The overall category for both these subcategories is termed dynamic external resistance
(DER).
In isokinetic training, muscle actions are performed at a constant angular velocity, which is
controlledby the machine.Unlike dynamic external
resistancetraining,thereis no setresistanceto overcome; however,sincethe velocity is controlled,any
force applied against the equipment results in an
the subequal and oppositereactionforce.t23lSince
ject can freely vary the level of effort in the entire
movement to accommodatefor pain or weaknessin
certainregionsof the rangeof motion, the isokinetic
Isokinetics
mode is very useful in rehabilitation.t24l
are sometimessortedinto a categorycalled 'accommodatingresistance'.[2s1
Also included in this category are devicesin which the resistanceis provided
by hydraulic cylinders, which function by limiting
the flow through an adjustable aperture. Although
not providing a strictly isokinetic movement, the
resistancesettingon thesemachinescan be adjusted
to limit the velocity within relatively narrow
ranges.l2slThese devices are sometimes also referred to as semi-isokinetic.t26lFor the purposesof
this review, the term 'accommodatingresistance'
was chosenfor the main category.Another form of
strengthtraining uses the inertia of a flywheel for
resistance.In this type ofergometer,the force exerted by the subject is transferred to a strap being
4
wound around the axle of a fixed flywheel.l271
concentric muscle action unwinds the strap and
overcomes the inertia of the flywheel, setting it
spinning on low friction bearings.The rotating flywheel soon causes the strap to start winding up
again; therefore, the machine returns the stored energy of the spinning flywheel via the strap and the
subject tries to resist the returning movement by
performing an eccentric muscle action. Caruso et
al.t28lhaveusedthe term 'isoinertial' to describethis
@ 2007 Adis Doto Infomotion BV. All rights reserued.
type of training, but Murphy and Wilsont2elhave

usedthe exactsameterm to describeDCER training.
With isoinertial flywheel ergometers, as with
isokinetictraining,the resistanceis effort dependent
for each type of action. Therefore, this mode of
exercisewill be included in the 'accommodatine
resistance'category.
1.3 Quontificotion of ExerciseFrequency,
lntensity ond Volume
Regardingexerciseintensityin dynamicexternal
resistancetraining, we choseto expressintensityas
a function of IRM in the exercise(s)performedfor
the muscle group in question(e.g. SOVoof IRM in
the squat).In many of the studies,the authorsprovided exact percentagesor at leastestimatesof the
percentage.In cases where the intensity was expressedonly as a function of how many repetitions
the subjects were able to perform (e.g. 8RM), we
estimatedthe relative intensitybasedon dataon the
relationshipbetweenthe number of repetitionsperformed and the IRM for the same or similar exerFor the squat,we usedthe RM tablesof
cises.t2'30-331
Wathan,t32l
which have been shown to be accurate
For
for the squatin previouslyuntrainedsubjects.t33l
the leg press,kneeextensionand arm curl exercises,
we useddatafrom Hoegeret al.t3olThe intensityfor
isometrictrainingwas quantifiedas a functionof the
maximumforce achievedduring a maximal voluntary isometricaction (MVIA).
While the quantificationof exerciseintensity in
conventionalresistancetraining and isometrictraining is relatively straightforward,how to expressthe
intensityor load for isokinetic and other accommodating training modes is less obvious. The forcevelocity relationshipof skeletalmuscledictatesthat
as the velocity of the concentricmuscle action increases,the maximum possibleforce decreases.[34'351
Depending on the training status, the maximum
torquedevelopedby the quadricepsduring eccentric
muscleactionsis slightly higher (trainedsubjects)or
not significantlyhigher (sedentarysubjects)thanthe
maximum isometric torque.t3slIn caseswhere the
peaktorqueduring eccentricmuscleactionsexceeds
that of isometric muscle actions, the peak torque
Med 2m7;37(3)
Sports
z5u
increasesonly marginally with increasingvelocity

and appears to reach a plateau at relatively low
velocities.t35l
The amount of force developedby the
muscleis generallyregardedas an importantstimuli
for muscle hypertrophy.t36l
Since maximum effort
(and therefore,recruitment of the greatestnumber of
motor units possible)was used in most accommodating training studies, an estimate of the torque
developed in relation to the isometric maximum
could provide some insight into the level of force
necessaryto produce hypertrophy as a consequence
of accommodatingresistancetraining.Therefore,in
addition to listing the training velocity, level of
effort and type(s)of muscle action(s),we estimated
the relative level of torque normalisedto the maximum isometric torque basedon the peak torque data
for the quadriceps in untrained subjects from
Amiridis s1 al.t3slFor the elbow flexors, data from
Hortobagyiand Katcht37land Paddon-Jones
et al.t38l
were used.
Wernbomet al.
sessionsper muscle group per week, as opposedto

training days per week. For example, a training
group may have trainedthe quadricepstwice a day,
three days a week. The frequency then is reportedas
six times per week. We are aware of the possibility
that the physiologicalresponsesmay differ between
training once a day six times per week and twice a
day three times per week.
1.4Clossificotionof TroiningStotus
In the majority of the studies,the subjectswere

reported as either untrained/sedentaryor as physically active. Physically active subjects generally
performed some form of endurancetraining, but not
any systematicstrengthtraining.Sincemany studies
reportedvarying levelsof activity amongthe participants and becauseendurancetraining induceslittle
if any muscle hypertrophy,l3el
the categories'untrained' and 'physically active' were combinedfor
analyses.Deschenesand Kraemertaolmade the folTraining volume is a measureof the total amount lowing classificationof training statuswith referof work (oules) performed in a given time peri- ence to resistancetraining: untrained, moderately
od.t23lIn this review, training volume refers to a trained,trained,advancedand elite. They suggested
single session.In a less strict sense,volume can be that the window of adaptationfor strength becomes
estimatedby the sum of repetitionst23lor even by the progressively smaller as the subject progresses.
number of sets performe6.t+tWhile simply stating However, becauseof the lack of studiesinvolving
athletesofdifferent trainingstatus,datafrom studies
the number of sets may seem a crude measureof
volume, severalmeta-analysest4-6'81
have shown sig- with trained, advancedand elite athletesare disnificant differencesin strengthgains betweentrain- cussedtogetherin the presentreview. In caseswhere
the strengthtraining statusof the subjectswas uning with single and multiple setsin favour of multiple sets, particularly for trained subjects.Theoreti- certain,information was soughtregardingthe CSA
of the exercised muscles and comparisonswere
cally, a given exercisevolume can be distributedin
madewith datafrom previouslyuntrainedindividumany different ways and as a consequence
result in
subiectsand strength
different adaptations. Therefore, several estimates alsl4l45land resistance-trained
61fle1es.lal,as-+71
of volume (number of sets,total number of repetitions, total duration of work and total work) were
L5 Colculotion of Chonges ond Rote of
usedin this review. However, insteadof expressing
Chonges in Muscle Cross-Sectionol
the total amount of work performed in joules, work
Areo (CSA)
was calculatedin arbitraryunits (setsx repetitionsx
intensity;.tzsl
If severalexerciseswereperformedfor
In most of the studiesreviewedhere,the authors
the muscle group (e.g. leg pressesand knee exten- reportedthe changesin CSA or at leastthe pre- and
sions for the quadriceps),the volumes for each of
post-trainingvalues.Sometimes,figures were used
thesewere summedto yield the total volume for the insteadof numericaldata; in such casesthe graphs
musclegroup.Regardingthe training frequency,we were measuredif possible. The relative changes
have chosen to report frequency as the number of
were calculatedby simply dividing the post- with
@ 2007 Adis Doto Informotion BV, All rights reserved.
Sports Med 2007j 37 (3)
231
the pre-training values. To allow for comparisons

betweenstudiesofdifferent length,percentchanges
per day were calculatedby dividing the changein
area with the length of the training period in days.
Although some studiest43'481
have shown preferential
hypertrophyof individual musclesin a musclegroup
and of different levels in the same muscle,no consistent pattern has yet emerged. Furthermore, data
from other studiest44'4el
suggestthat changesin muscle CSA at the middle level and muscle volume after
a training period are of a similar magnitude. In
addition, studies by Aagaard and co-workersts0land
Tracy et al.t5llhave confirmed that quadricepsmuscle volume can be accuratelypredictedfrom a single
scan at the middle level. Therefore,no distinction
was made in this review between muscle volume
and CSA as indexesof muscle size. A further discussion on the subject is provided by Tracy and
colleagues.ts2l
There are severalreasonsfor focusing mainly on
changesin whole muscle CSA and not MFA. As
arguedby Narici and colleaguest43l
and D'Antona et
a1.,t471
care should be taken when drawing conclusions concerning the whole muscle mass from
changesoccurring at a single biopsy site. In the
study of Narici et al.,tarl16" changesin mean fibre
area(2Voincreasein MFA) were not representative
of either the vastuslateralisat the samelevel as the
biopsy site(7.sEoincreasein muscleCSA by MRI),
or of the quadricepsas a whole (=l6Vo increasein
muscle CSA). Apart from the obvious risk of not
detectinghypertrophybecauseof samplingmuscle
tissue from just a single site in a very large and
architecturallycomplex muscle group such as the
quadriceps,the lower limit for the increasein fibre
size that can be detected is l0vo. Furthermore, the
coefficient of variation betweenrepeatedbiopsiesis
quite large,=L5-2OVo(seeNarici s1al.ta3lfor discussion). Thus, hypertrophic changes that are detectable at the whole muscle level may go unnoticedif
only fibre areasare measured.Finally, as discussed
by McCall and colleagues,t5rl
on" should be cautious in ruling out hyperplasia(an increasein the
numberof muscle fibres in a muscle)as a possible
o 2007 Adis Dolo Informotion BV. All righls reseNed.
contributingmechanismfor whole musclehypertro-
phv.
This is of coursenot to say that changesin fibre
area are irrelevant as an index of muscle mass.
Ideally, a training study would include measurements at both the cellular level and at the whole
musclelevel. Also, it should be noted that the relative changesin mean fibre area are usually of a
larger magnitude than the changesin anatomical
muscle area.tal However, we feel that the focus on
changesin whole muscleCSA and volume,as measuredby scanningtechniques,isjustified sincethese
are arguably the most sensitiveand representative
measuresof whole muscle mass,even though they
may underestimatethe changesat the cellular level.
Regardingtraining issueswhere the evidenceat the
whole musclelevel is limited, we will discussrelevant studies,if any, which have measuredMFA
changes.
2. Results Port l: Quodriceps
Studies
2.1 Quodriceps Studies:Dynomic

ExternolResistonce
After applicationof the inclusion and exclusion
criteria,the literaturesearchresultedin 44 original
afticlestl3'4346's4-e21
investigatingquadricepsmuscle
CSA or volume before and after DER training.
Becausethere were often more than one training
group or more than one limb that receivedtraining,
these44 articlesyielded65 datapointsfor CSA. Five
ofthe articles(sevendatapoints)involved trainedto
elite strength athletes.These were too few to allow
for any meaningful analysisand will be discussed
briefly in sections4.1 and 4.6. Five studiesdealt
with pure concentricand/or eccentrictraining, with
four datapointsfor concentric training and three for
eccentric training. Four datapointsfrom four different studiesinvolved training wherethe subjectsfinished each set with several repetitionsin reserve.
Since stopping well short of muscular failure has
beenshownto yield modesthypertrophyin comparison with performingeachsetto muscularfailure,lEsl
even when the total volume is similar, thesestudy
groups were excluded. As a result, except where
Sports
Med 2007;37(3)
Wernbom
et al.
ZJL
otherwiseindicated,the analysisfor DER dealswith

training using combined concentric-eccentric
muscle actions (47 datapoints), with previously untrained subjects.
sions per week, the increasewas 0.ll%o per day

(range:0.04-0.267o).
2.1.4Intensry
The mean peak intensity (the highest value
reached during a session, averaged over the entire
2.1.1 Lenglhof lroining Peilod,Averdge lncrcq,se
in CSAond CSAPet Doy
period)wasT3Voof lRM. The averageintensity(the
The averagelength of the training period was 79 meanof all training sets)was 66Voof lRM. Inspecdays.The shorteststudy was 14 daysand the longest tion of figure 2 reveals a tendency for greater rates
lasted6 months.The averageincreasein quadriceps of increasefor intensities>60Voof 1RM when comCSA was 8.5Vo(range: l.l-17.37o). The average pared with intensitiesbelow this level; however,it
increaseper day in CSA for training with combined should be noted that only six datapointsinvolved
concantric-eccentric
muscle actions(47 datapoints) training with mean peak intensities<607oof lRM.
was0.l2%oper day (range:0.044.55Vo).If the study The study of Abe s1 al.tsalis not shown becauseof
of Abe et al.l841
is excludedas an 'outlier' becauseof
the unusuallyhigh training frequency(12 times per
the unusually high training frequency and rate of
week) and very high rate of increase.The peak
gain, the average increase was O.llVo per day intensityin this study was 20Voof lRM.
(range: 0.04-0.267o).For pure concentrictraining,
2.L5 Volume
the increasewas O.06Voper day and for pure eccenThe mean number of setswas 6.1 and the mean
tric training O.03Voper day.
number of total repetitions was 60. The results are
shownin figure 3. The study of Abe s161.t8+l
2.1.2 Rateot Goin in CSA:Men versusWomen
was not
included in the following analysis of volume. InIn six studies, groups of men and groups of
women followed exactly the sametraining program- spectionof the datapointsrevealedfour identifiable
mes.The averageincreasesin CSA per day in these 'clusters' in the range of total repetitions.Fourteen
studieswere 0.l3%ofor men and0.l4%ofor women. datapointswere found in the rangeof 2l-39 repetitions, 14 datapointsin the range of 40-60 repetiBecausethesedifferences were consideredas negligible and becauseseveralstudiescontainedgroups tions, I I datapointsin the 66-90 repetition range
and finally 6 datapointsin the range>100 repetitions
consistingof both men and women, data from all
studieswere pooledin the analysisfor DER training per session.The averagerate ofincreaseofCSA for
each cluster was as follows: 2I-39 repetitions=
(section2. I ).
O.l2% per day; 40-60 repetitions= O.l3Voper day;
2.1.3Frequency
66-90 repetitions= 0.087oper day; and >100 repetiThe mean training frequency was 2.8 times a tions = 0.l2%oper day. No studieswere found that
week. The most common frequency was three times
+ 0.6
a week (22 of 47 datapoints),followed by two times
<
a week (17 datapoints).Frequenciesbetween and
o.u
abovethese(2.3-4 times per week) were notedin a
0.4
few cases. No studies were found that iirvolved
(,
U.J
O
training at frequencies of more than two times per
.- v.z
o
week. A plot of frequency versus percentageinl. l'
<! nr
o u. l
creasein quadricepsCSA per day is shownin figure
o
1. The highestrate of increase(O.55Vo
per day) was
-U
0 1 2 3 4 5 6 7 8 I 1011121314
reported for the training studyts+lwith the highest
Numberof sessionsDerweek
frequency(12 times per week). For the frequencyof
Fig. 1. Frequencyof traihingvs percentageincreasein cross-sec.
two sessionsper week, the averageincreasewas tionalarea (CSA)per day of the quadricepsduringdynamic
exter.
0.11% per day (range: 0.034.21Vo); for three ses- nal resistancetraining(numberof study groups= 47).
f t'
O 2007 Adis Dolo Informolion BV. All righls reseNed.
Sports
Med 2007;37(3)
233
0.3
2.2.1Lenglhof frolnlng Peilod,Avercge lncrcose

ln CSAond 6A PetDdy
The averagelength of the training period was 52
0.2
days.
The shorteststudy was l3 daysand the longest
.
.
;
'
.3
'
.
6 o.rs
lasted84 days.The meantotal increasein CSA was
t.tt
.s 0.1
..
o
5,8Vo(range:2.5-l8.4%o)for all typesmuscleaction
t.t.3
o 3
o.os
For pure concentric training, the mean
combined.
$
lr'
s0
total CSA increasewas 6.l%o (range: 2.5*18.47o),
0 10 20 30 40 50 60 70 80 90 100 for pure eccentric training the mean increase was
(%of1RM)
Intensity
4.2Vo(range:2.5-6.2Vo)and for combinedconcenFig.2. Peakintensity
vs percentag
Increas
in crossof training
(CSA)
per
quadriceps
sectional
area
dayof the
duringdynamic tric-eccentric training, the corresponding figures
= 46).RM= were 6.0Vo(range: 4,1-7.47o).The mean CSA rn(number
resistanco
external
training
of studygroups
repetition
maximum.
crease per day for pure concentric training was
0,l3%o(range:0.054.44Vo),for pure eccentrictraininvolved <21 repetitionsper sessionon average.For
ing 0.067o (range: 0.044.09Vo) and for combined
the number of sets, the following groupings were concentric-eccentric training 0.l6Vo (range:
made: 3 sets (average CSA increase = O.O9Voper 0.064.21Vo).The averageCSA increasefor all the
day); 4 sets (0.1370per day); 5-6 sets (O.l3Voper accommodatingresistancemodes combined was
per day.
day);7-9 sets(0.097oper day); and >10 sets(0.147o 0.l3%o
<
f o.zs
per day). No studieswere found which involved <3

sets.When training volume was expressedin arbitrary units [sets x repetitions x intensity], no apparent relation was found between the volume and
increasesin CSA per day (data not shown).
2.2 Quodriceps Studies:
Accommodoting Resistonce
The literature search resulted in 17 original articlest2T'42'48'87'e3-1051
investigatingquadricepsmuscle
CSA or volume before and after an accommodating
resistance-trainingprogramme. These 17 articles
yielded 2l datapoints for CSA. Of these, 14
datapoints involved pure concentric training, three
involved pure eccentrictraining and four involved
training with combinedconcentric-eccentric
muscle
actions. Some of the studieste3'e5-e8l
included subjects with strength training experience. Based on
anthropometric and quadriceps muscle CSA data,
they were considered as 'moderately trained' and
are included in the analysisin sections2.2.1-2.2.7,
together with previously untrained and physically
active individuals.
o 2m7 AdisDoto Informcrtion

BV.All rightsreserued.
2.2.2Frequency
The mean frequency for concentric training was
3.4 times a week. Three sessions(nine datapoints)
per week yielded an average increase in CSA of
0.l3vo per day and 3.5-4 sessionsper week (four
datapoints)yielded an averageincreasein CSA of
O.l2Vo per day. Five sessions(one datapoint)per
week yielded an averageincreasein CSA of 0.22Vo
per day. The largestrate of CSA increase(O.44Voper
day) was noted in a studytlo4lusing three sessions
per week.No concentrictraining studieswere found
with training frequenciesbelow three sessionsper
week or abovefive sessionsper week.The frequen-
s
(t
0.3
0.25
o.2
0.15
.;
0.1
o
o
0.05
E
o
a'
. .. {
at'
tar
l'
0
0
20
oo
'
a
al
l'
40
100 120 140
60
80
Total number of repetitionsper session
160
Fig. 3. Total number of repetitionsvs percentageincreasein crosssectionalarea (CSA) per day of the quadricepsduring dynamic
erlernal resistancetraining (number ot study groups = 45).
Med 2@7;37(3)
Sports
234
3
-
Wernbomet al.
0.s
0.45
S o.o
0.3s
oo 0.3
u, 0.25
0.2
?
' o.ts
o.t
fi
0.05
When the percentage values for eccentric torque

were applied and all accommodatingmodes and
types of muscleactionswere combined,no relation
was found between the maximum torque developed
during trainingand the rate of increasein quadriceps
CSA (seefigure 5).
Eo
2.2.5 Volume
0123456
For pure concentrictraining,the largestincreases
perweek
Numbr
ofsessions
incross-sec- occurredwhen the total number was between50 and
Fig.4. Frequency
vs percentage
increase
of training
perdayofthequadriceps
tionalarea(CSA)
during
accommodating60 muscleactions(six datapoints,0.l9%o
increasein
= 21).
(number
and/or
training
ofstudygroups
concentric
eccentric
CSA per day l0.l3%oincreasein CSA per day withand Rafeeitrsll).
out the studiesof Akima et al.trO3]
cy for pure eccentric training was three times per Two datapointswere found in the interval of 3G40
week in all studies(threedatapoints).For the com- muscleactions(0.06Voincreasein CSA per day) and
bined concentric-eccentrictraining regimens, the six datapointsbetween120 and 480 muscleactions
frequencies were two (one datapoint), 2.3 (two
(0.l0Vo increasein CSA per day). If studieswith
datapoints) and three (one datapoint) times per pure eccentric and combined concentric-eccentric
week. The frequency plot for all categories commuscle actions are included in the analysis, the
bined is shown in figure 4. No relationshipbetween increasesstill tend to reach their maximum in the
the frequencyand the rate ofCSA increaseis appar- rangebetween50 and 60 muscleactionsper session
ent from figure 4.
(ten datapoints,0.187o increasein CSA per day).
See figure 6.
2.2.3 Velocity
Regardingthe number of setsfor pure concentric
For pure concentrictraining, the most common
training,
one datapoint was found for 3 sets,one for
velocities were 60o/s (seven datapoints,0.l3Vo in4
sets,
six
for 5-6 sets,none for 7-9 sets and six for
creasein CSA per day) and 120ols(four datapoints,
)10
sets.
The
highestincreaseoccurredat 5-6 sets
0.16%oincreasein CSA per day). Two studiesused
(O.l9Vo
in CSA per day [O.l37oincreasein
increase
180'/s (0.147oincreasein CSA per day) and one
per
CSA
day
minus Akima et al.tro3l and
study used 90"/s (0.05Voincreasein CSA per day).
in CSA for >10 setswas
Rafeeitroall).
The
increase
The velocities for eccentric training and combined
per
and
increase
for 3-4 sets was
O.lOVo
day
the
concentric-eccentrictraining were in the range bepure
with
and combined
O.06Vo.If
studies
eccentric
tween 45 and 90o/s.
2.2.4forque
Datafrom Amiridis e1al.t3slon untrainedsubjects
suggeststhat the isokinetic concentricpeak torque
of the quadricepsexpressedas a percentageof maximum isometric torque is =59Vo,69Vo,77Voand 887o
at the velocitiesof 180o/s,l2}ols,90o/s and 60o/s,
respectively. When these percentage values were
applied to the concentric training studies,no relation
was found between the level of torque developed
and the rate of increasein CSA. The eccentric peak
torque of the quadricepsin relation to the maximum
isometrictorque is =lO4Vo,lO6Voand l04Vo at the
velocities of 30o/s, 60o/s and 90"/s, respectively.
@ 2m7 Adis Dolo Informotion BV. All righls reseNed.
;x
-
u.c
Fno
o
q 0.3
o
? o.'
fr o.r
g
go o
0
20
40
60
80
Torque(% ol MVIA)
100
120
Flg. 5. Torque vs percentage incraase in cross-sectional area

(CSA) per day of the quadricepsduring accommodatingconcentric
and/or eccentrictraining(numberof study groups= 21). MvlA =
maximal voluntary isometric action.
SportsMed 2m7;37(3)
a
;<
0.5
tween 25 and 50 units (0.1470increasein CSA per

day), six between50 and 100 units (0.l4%oincrease
in CSA per day) and five in the interval between100
and 265 units (0. l07o increasein CSA per day).
oo
o
oo
o3
?o,
!.
I o t :ro.
o
co
0
50
100 150 200 250 300 350 400 450 500

Numberof muscleaclionsper session
Fig. 6, Numberof muscleactionsvs percentageincreasein crossseclionalarea (CSA) per day of the quadricepsduringaccommodatingconcentricand/oreccenlnctraining(numberof studygroups
= 211.
concentric-eccentric
muscle actionsare included in
the analysis,the results are as follows: 3 sets (two
datapoints)= O.08Voincreasein CSA per day; 4 sets
(six datapoints)= g.12Eoincreasein CSA; 5-6 sets
(sevendatapoints)= g.17qoincreasein CSA (O.|lVo
without the studies of Akima et al.tr03l and
Rafeeitr0al);
and >10 sets (six datapoints)= O.lOVo
increasein CSA per day.
2.3 Quodriceps Studies:lsometric Resistonce

The literature searchresulted in six original articlest55'lOGll0l
investigatingquadricepsmuscle CSA
or volume before and after an isometric resistance
training programme, yielding nine datapointsfor
CSA.
2.3.1Lengthof froining Peilod,Avercge lncrease
in CSAond CSAPer Doy
The averagelength of the training period was 84
days.The shorteststudy was 56 daysand the longest
lasted98 days.The meanincreasein total CSA after
the training period was 8.9Vo(range: 4.8-l4.6Vo)
and the averagerate of CSA increasewas 0. I I7o per
day (0.06-0.26Vo).
2.3.2Frequency
Three sessionsper week (four datapoints)resulted in an increasein CSA of 0.127oper day and four
2.2.6 fotol Durolion Per Session
For pure concentric training, eight datapoints sessionsper week (five datapoints)resulted in an
increasein CSA of 0.ll%o per day. The largestrate
were distributed between 37.5 and 75 secondsof
total duration of muscle work. The averageincrease of gain (0.26Voincreasein CSA per day) was reportin CSA for thesedatapointswas0.76Voper day. The ed in a studytlootusing three sessionsper week.
other six datapointswere distributed between 170
2.3.3lnlensity
and 300 seconds.The averageincreasein CSA for
The most common intensity was 70Voof MVIA
these datapoints was 0.09Vo per day. For the pure (seven datapoints),the intensity in the other
two
eccentric and combined concentric-eccentrictrain- caseswas 80Voand 1007o,respectively.The largest
ing groups taken together, the sevendatapoints(av- rate of CSA increase(O.26Vo
per day) was found in
erage of 0.l2%o increasein CSA per day) were the studyttoot
that used the highesttraining intensity
distributedbetween40 and 84 seconds.
(lO0Voof MVIA).
2.2.7Time-IorqueProduclPerSesslbn
The time-torqueproduct per sessionwas calculated by multiplying the total durationwith the estimatedpeak torque(with maximum isometrictorque
assigneda value of 1) and is reportedherein arbitrary units. No relation was found betweenthe timetorque product and the rate of gain in CSA, regardless of whethereccentricand combinedconcentriceccentrictraining was includedin the analysisor not
(data not shown). For all types of muscle actions
combined, ten datapointswere in the interval beO 2007 Adis Dqlo Informotion BV. All righls reseNed.
2.3.4Volume
The total number of repetitions rangedbetween4
and 150.The time each repetitionwas held ranged
betweenI and 30 seconds,while the total duration
of musclework per sessionwas between80 and 150
seconds.No relationwas found betweenthe number
of repetitions and the increaseper day in CSA.
Similarly, when volume was expressedas the total
durationper sessio4and as the product of intensity
and total duration, no apparent relation between
volume and rate of increasein CSA was observed.
Sports
Med 2m7;37(3)
Wernbom et al.
althoughthe highestrate of increasein CSA (0.267o increasein quadricepsCSA (347o)was notedin one
per day) was observedfor the trainingstudytl06l
with
of the longer studies,tll2lafrcr20 weeks of training
(seefigure 7). There was also an apparenttendency
the largest product of intensity and duration.
for the rate of increase in CSA to decreasewith
2.4 Quodriceps Studies:Combined Strength
increasinglengthsof the training period (figure 8). If
ond Enduronce Troining
one regards the studies of Abe et al.tsal and
Rafeeitloalas
outliersbecauseoftheir unusuallyhigh
The literature search resulted in seven original
ratesof increase,the slope becomesless steep.
investigating quadriceps
altislestTT'78'86'e6'lll-ll3l
muscle CSA or volume before and after a combined
2.6 Quodriceps Studies:Strength Troiningos
training programme,yielding ten datapoints.Some
o Countermeosure During Unlooding
of the studies included groups that performed only
The literature search resulted in eight original
strength training, these have been included in sections 2.1 and 2.2. Becauseof the limited data and afticlestll4-l2llinvestigatingquadricepsmuscleCSA
different modes of endurancetraining (rowing, runor volume before and after a resistance training
prograrnme as a countermeasureduring otherwise
ning and cycling), as well as the different modesof
strengthtraining, no analysis was performed regard- unloadedconditions (bed rest or limb suspension).
ing frequency,intensity and volume.
Theseeight papersyielded nine datapointsfor CSA
for the training groups and nine datapointsfor the
2.4.1Poteol Goin in CSA:Combined frclnlng
control groups, which performed no training. One
vercusPurc Strenglh Trcining
(one datapoint) used isometric training as a
In four of seven studies,tTT'78'e6'llll
comparisons study
countermeasure,
one study (one datapoint)used viwere made between pure strength training and comwith isometric training, four studbration
combined
bined training regarding increasesin quadriceps
(five datapoints)used dynamic external resisies
CSA. If theseare summarised,the resulting average
tance with coupled concentric-eccentricmuscle acincreases are as follows: pure strength training =
(two datapoints) used
O.09Voincrease in CSA per day; and combined tions and two studies
accommodating
resistance
with
coupled concentrictraining = O.ll%o increasein CSA per day. If the
eccentric
muscle
actions.
combined training groups of the other three studiesl86'll2'1131
are included, the averagerate of CSA
2.6.1UnloadingversusUnloadingand
ExerclseCountetmeoswe
increasefor combined training becomes0.l2Vo per
day.
The averagelength of the training and unloading
period was 49 days (range: 20-ll9 days). The mean
2.4.2 Lengthof fraining Periodand lncraose
decreasein quadriceps CSA for unloading was
in CSA
The shortest study was 70 days and the longest
lasted 168 days. The mean total increasein CSA
after the training period was l1.l%o. The largest
increasein CSA was 347o(0.24Voper day) and the
smallestwas 3.9Vo(0.O5Voper day).
2.5 Quqdriceps Studies:All Voluntory
TroiningModes
2.5.1Lengthol lrclning Periodand lncrcose
in CSA
Longer trainingperiodsgenerallytendedto result
in largerincreasesin CSA. Accordingly,the greatest
o 2007 Adls Dolo Informolion BV. All rights reserved.
40
Qa.
6go
o^_z a
g
920
'i tu
$to
cv
o
o
20
40 60 80 100 120 140 160 180 200

Length of traning period (days)
Fig.7. Traningperiod vs tbtal percentageincreasein cross-sectional area (CSA) ot the quadriceps during all types of voluntary
strengthtraining(numberof study groups= 91).
SDortsMed 2m7;37 (3)
g o'u
timulation to evoke muscle actions.In the first of

these,teelpreviously untrained subjects performed
3-5 sets of l0 unilateral combined concentricand
E o.a
eccentric actions in an isokinetic dynamometerat a
9? 0.3
velocity of 75'ls, for two sessionsper week for 9
c
t
6 0.2
weeks.The increasein quadricepsCSA was l0.l7o
a
o.r
(0.16%oper day). In the secondstudy,t1221

from the
o
;rl ll3
.l '
0
same research group, recreationally resistance20 40 60 80 100 120 '140 160 180 200
o
trained subjects performed an identical protocol to
Length of laining period (days)
that in the first study,for two sessionsper weekfor 8
period
percentage
vs
Flg. 8. Training
change in cross-sectional
weeks.Thesesubjectscontinuedtheir normal resisarea (CSA) per day of the quadricepsduring all types of volunlary
strengthtraining(numberof study groups= 91).
tance-trainingregimensduring the study, including
exercisesfor the quadricepsof both sides.The in-ll.lVo and the mean rate of decreasewas 43070
crease in CSA was 9.87o (0.l8%o per day). The
per day. The degreeof loss appearedto be related to
control limb that performed only regular resistance
the length of the unloading period, as the largest exercise showed no increasein quadricepsCSA.
decrementsin CSA were found in the studieswith
Another group performed exactly the same prothe longestunloading period. However, the rate of
grarnmebut also receivedcreatinesupplementation.
loss in CSA seemedto decreasewith the length of
This group increasedthe CSA with l2.l%o (O.22Vo
the unloading period, so that the longer studies per day) but this was not significantly greaterthan
showedlower averagerates of change.For the un- the other (placebo) group. The control limb that
loading plus strengthtraining countermeasure,
the performedregular resistanceexerciseonly showeda
CSA increasedby an averageof l.3%o(0.03Voper slight increase(SVo)in quadricepsCSA. In the third
day). If the isometric training and the vibration study,ttz3l
electromyostimulationwas usedto evoke
training groups are excluded,the increasein CSA 40 isometric muscle actions per session,four sesbecomes2.7Vo(0.O9Voper day). The largest total sionsa week for 8 weeks.The increasein CSA was
CSA increasewas 7.l%o (0.22V0per day) and the 6Vo(O.ll%oper day).
largestrate of gain w as 0.3OVoper day (6.0Voin total
3. Resulls Porl2: Elbow Flexor Studies
CSA). The greatesttotal CSA decrementwas-3.8Vo
(4.l9vo per day) for the isometriccountermeasure.
The greatest total CSA decrease for the dynamic
3..| Elbow FlexorStudies:Dynomic
groupswas-l.9Vo (4.O2Voper day). Becauseof the
ExternolResistonce
limited data and differences in both training modes
and unloading models, no analysiswas performed
The literature searchresulted in 16 original artiregarding frequency, intensity and volume. The cles[25'53'124-1371
investigating elbow flexor (biceps
lowest training frequencywas every third day (2.3 and brachialis)muscle CSA or volume before and
times per week) and the highestwas twice a day (14 after a DER training programme with specific exertimes per week).
cisesfor the elbow flexors. Thesepapersyielded 36
datapoints for CSA. Three of the papers (seven
datapoints)included highly-trainedsubjects,while
2.7 Quodriceps
one paper included recreationally-trainedsubjects
Studies:Electromyostimulotion
(onedatapoint),who had previouslytrainedwithout
any structuredprogrammesor specificgoals.Based
The literature search resulted in three original
nllislsstee'122'1231
investigating quadriceps muscle on strength and CSA data, the latter group was
CSA in healthy uninjured subjectsbefore and after a regardedas 'moderately trained' and included in the
resistancetraining programme using electromyos- analysisof'untrained' and 'physicallyactive' sub6
u .c
" i. l',
i','.
o 2m7 AdisDoto InformotionBV.All rightsreserued.
SportsMed 2m7j 37 (3)
238
jects in sections 3.1.1-3.1.4. Four groups (four

datapoints) from three other studies were excluded
because they either performed only non-specific
exercisesfor the bicepsand brachialismuscles(e.g.
latissimuspulldown, dumbell row) or they did not
perform their elbow flexor exerciseswith nearmaximal effort or near muscular failure in any set. One
group from another study was excluded becausethe
eccentric phasewas performed in a plyometric manner with the resistance building up momentum
before the subject started to resist it, making it very
difficult to estimate the actual intensity. As a result,
the analysisin sections3.1.1-3.1.4for the variables
of frequency,intensity and volume is basedon 24
datapoints for training with previously untrained to
moderately trained subjects.All studies involved
muscle
training with combinedconcentric-eccentric
actions, but in one study the eccentric phasewas
overloadedwith =1807oof lRM.
3.1.1Lenglhof froining Period,Average lncrcase
in CSAond CSAPerDoy
The averagelength of the training period was 9l
days.The shorteststudy was 30 daysand the longest
lasted6 months. The averageincreasein flexor CSA
was 15.87oand the averageincreaseper day in CSA
was O.20Vo.The highest increasein elbow flexor
CSA (337o)was noted in the longeststudy, after 6
months of training,tl3Olalthough an almost equal
increase (32.64o) was observed in another study
after ll weeks.tl25l
3.L2 Frequency
The results are shown in figure 9. The mean
training frequencywas 2.9 times a week. The most
common frequency was three times a week(l'l of 24
datapoints),followed by two times a week (six
datapoints). The highest frequency was four times
per week. No studieswere found that involved training at frequenciesof less than two times per week.
The highest rate of CSA increase (0.597oper day)
was noted for a training studytl28lwith a frequency
of four times per week. For the frequency of three
sessionsper week, the average CSA increase was
O.l8Voper day and for two sessionsper week, the
CSA increasewas 0. l87o per day.
@ 2007 Adis Doto Informolion BV. All righls reserved.
Wernbom
et al.
3.1.3lnlensity
The mean peak intensity (the highest value
reachedduring a session,averagedover the entire
period)wasl2Vo,which was the sameasthe average
intensity (the mean of all training sets), since no
study reported using different loads during one and
the samesession.The highestmeanintensityreported was 1807oof lRM, in a studytr26lthat used
eccentric overload, in addition to performing the
concentricphasewith a lower resistance.This was
also the only study that used such overload; the
others used the same resistance in both types of
muscle actions.The lowest resistancereportedin a
study was 107o. When the intensity was plotted
against the rate of increase, a tendency was found
for the rate to increasewith increasingintensity.The
highestratesof increasetendedto occur aroundT5Vo
of IRM (seefigure l0).
3.1.4Volume
The resultsfor the number of repetitionsversus
CSA per day are shown in figure 11. The mean
numberof setswas 5.4 and the meannumberof total
repetitionswas47. Threeclustersof datapointswere
identified as follows: (i) 7-38 repetitions (ten
datapoints);(1i) 42-66 repetitions (nine datapoints);
(iii) and 74-120 repetitions(five datapoints).The
maximum rate of CSA increase was found in the
interval between42 and 66 repetitions(0.267oper
day). For 7-38 repetitions,the CSA increasewas
O.lSVoper day, and for 74-l2O repetitions,the rate
was 0.1870per day. For total sets,the rate of CSA
increaseappearedto peak between4 and 6 sets(nine
g 0.,
0.6
b 0.s
< 0.4
U'
? o.'
0.2
E
G
E
0.1
o
-0
01234
Numberof sessionsper week
Flg. 9. Frequencyof training vs percentageincrease in cross-sectional area (CSA) per day of the elbow flexors during dynamic
external resislancetraining (number of study groups = 24).
Med 2@7;37(3)
Sports
g 0't
3,3 Elbow FlexorStudies:lsometric Resistonce
0.6
b 0.s

articlest2l'|24'l3el
investigatingelbow flexor muscle
o
.
CSA before and after an isometric resistancetrain?
ing programme, with a total of three datapointsfor
0.2
3
G
E 0.1
CSA. The mean CSA increaseper day was O.l4Vo.
The largesttotal increasein CSA (23.0Vo)was noted
0
o
20
2
0 4
40
0 6
80
0 1
100
00 1
120
20 1
140
4 0 160
160 180
180 200
60
0 8
for the longeststudy,tztlwhich lasted 100 days.The
(%of l RM)
Inlensity
frequencywas six times a week for two sludlest2l'l2al
Flg. 10.Peakintensityof trainingvs percentage
increasein cross- (O.23Voaverage CSA increase per day and O.06Vo
sectionalarea(CSA)per day of the elbowllexorsduringdynamic
per day, respectively) and three times per week for
externalresistance
training(numberot studygroups= 24). RM =
repetition
maximum.
the third studylt3el(OJ3Voper day). The intensityin
the three sludisst2l'l24'l3el
ws l00%o,67VoandSOVo
of
MVIA,
respectively.
The
number of total repetidatapoints, 0.247o increase in CSA per day). For
was
tions
3-24.
The
total
duration
of contractile
3-3.5 sets (ten datapoints), the CSA increase was
activity was 30 secondsin two studiesl2l'1241
and 96
0.177o per day and for >9 sets (five datapoints) the
seconds
in
the
third
study.tl3el
CSA increase was 0.187o per day.
0.4
<
g)
4. Discussion
3.2 Elbow FlexorStudies:

Accommodoting Resistonce
investigating elbow flexor muscle
311isls5t25'48'1381
CSA before and after an accommodatingresistance
training programme, yielding four datapoints for
CSA. Of these,two datapointsinvolved pure concentric training and two involved pure eccentric
training. The longest study was 140 days and the
shortest was 56 days. The mean CSA increasesper
day were O.l6Vo and O.l27o for concentricand eccentric training, respectively. Both the largest increasein total CSA (16.3Vo,O.l27oper day) and the
per day, ll%o
highestrate of CSA increase(O.2OVo
total increase in CSA) was noted for concentric
training. Becauseof the lack of data involving pure
concentric training, combined concentric-eccentric
and pure eccentrictraining, no formal analysisconcerning the effects of frequency,intensity and volume was performed. The frequency was three times
a week for all studies. The number of sets and
repetitionsper set was 4.6 and 10, respectively.The
average total duration of contractile activity was
84.8 seconds(range: 13.9-146.2seconds),
@ 2m7 Adis Doto Infomolion BV. All rights reserued.
4,1 Frequency
For quadricepstraining with dynamic external
resistance,the largest rate of gain in CSA (0.557o
per day) was noted in the studyt8alwith the greatest
training frequency(12 sessionsper week). However, it should be noted that (i) this study lasted for
only 2 weeks; (ii) the intensity was 2OVoof IRM;
and (iii) the training was performed in combination
with partial vascular occlusion. Therefore, the results from this study should be viewed with caution
when considering the application of extremely high
0.7
o
E
o
CL
0.5
0.6
0.4
0.3
o.2
E
o 0.1
c
0
0
20
40
60
80
100
120
Numberot repetitionsper session
140
Flg. 11, Total number of rpetitions vs percentage increase in

cross-sectionalarea (CSA) per day ol the elbow flexors during
dynamiceliternalresistancetraining (numberol study groups= 24).
SportsMed 2m7;37 (3)
Wernbom
et al.
frequenciesin more conventionaltraining. Furthermore, it is interesting to note that there was no

difference in the mean rates of increasein CSA
between two and three sessionsper week for DER
quadricepstraining (O.llVo vs 0.ll%o per day, respectively).For accommodatingresistancetraining
and isometric quadricepstraining, it is difficult to
evaluateany trend becauseof the limited numberof
studies and the smaller range of frequencies.The
highestrate of CSA gain in the isometriccategory
(0.267o per day) was achieved in a studytro6lin
which the subjects trained three times per week;
however, this was also the study with the highest
training intensity (l00Vo of MVIA) and the highest
product of intensity and total duration.The highest
rate of CSA increasefor accommodatingresistance
training (O.44Voper day) was noted in a studytto+t
that used a frequency of three sessionsper week,
followed by a studytlorlthat useda frequencyof five
times per week (0.22Voincreasein CSA per day).
High rates of CSA increase(O.l74.2l%o per day)
were also observedin threes1udis5t27'87'1051
in which
performed
per
training was
two to three times
week.
For the unloadingplus resistanceexercisecountermeasurestudies, the highest rate of CSA gain
(O.3OVo
per day) was notedfor the studytrrslwith the
greatesttraining frequency(14 sessionsper week),
although a high rate of CSA increase(0.22Voper
day) was also noted in a studyttl8lwith a frequency
of every third day. However, caution is warranted
when comparingthe resultsof thesestudiesbecause
of possible differences in the effects of the different
models of unloading and,/orthe exerciseregimens.
For example, Tesch and co-workerstll8lemployed
limb suspensionfor unloading and knee extensions
for the exercise countermeasureand showed a
marked hypertrophic responsein the quadricepsof
the unloadingplus exerciselimb. In a differentstudy
by Alkner and Tesch,tllelwhen using bedrestfor
unloadingand horizontal squats,but with the exact
samedosageof training as in their other study,tll8l
the exercise resulted in no hypertrophy but still
managedto completely counteract the atrophy that
was evidentin the pure bedrestgroup. Also notable
is that in one bedrest51udy.tll+l
which usedisometric
@ 2007 Adis Doio Informoiion BV. All righis reserued.
training at l0o7o of MVIA as a countermeasure,

trainingoncea day was insufficientto preventall the
atrophy. In yet another bedrest study,tltzltraining
once a day using a leg press,with coupledconcentric-eccentricactionsat 90Voof I RM, preservedthe
muscle volume but failed to induce significant hyperuophy. Further research is obviously neededto
addressthe effects of different exercise modes and
regimens (e.g. different frequenciesand distributions of the total training volume), as well as the
impact of the unloadingmodel employed.
For elbow flexor training with dynamic external
resistance,the greatest rate of increase in CSA
(O.59Vo
per day; 17.1Vototal increasein CSA) was
observedin a studytl2slwith a frequency of four
times a week. The second, third, fourth highest
increasein CSA ratesnoted were0.42Vo,0.38Vo
and
O.32Voper day, respectively. These three studiestl25'126'1321
used a frequency of three times per
week. However, with the exception of these three
studies, the average values suggest that there is
relatively little difference between training the elbow flexorstwo or threetimes per week,in termsof
per day for both
the rate of increasein CSA (0.18%o
frequencies).However,the highestincreasein CSA
that was noted for two times per week was O.24Vo
per day, compared with 0.42Vo per day for three
times per week. As noted in section3.2, all accommodating resistancetraining studies involving the
elbow flexors used a frequency of three times a
week. Regarding frequency for isometric elbow
flexor training, the highest rate of CSA increase
(0.23Voper day) was noted for a studyt2rlwith a
frequencyof six times per week, but this was also
the study that had the highest intensity (I00Vo of
MVIA). Remarkably,this regimenconsistedof only
three isometric actions per day, each lasting l0
seconds,thus resulting in a total of 30 secondsof
maximal isometricactivity each day.
While a higher than normal training frequency
(four or more times per week) can result in rapid
hypertrophy in the initial stage, it should also be
noted that several of these sludisstsa'lls'1281
lasted
only between 2 and 4 weeks. It is uncertain if
trainingat thesehigh frequencieswould continueto
SportsMed 2m7j 37 (3)
yield very high ratesofincreasesin CSA or perhaps

result in diminishingreturnsor even overtraining
after a longerperiod.Inspectionofthe datafrom the
study of Abe et al.tsal suggests that the rate of
increasein CSA was considerablyslowerduring the
second week compared with the first week, dropping from a maximum of =l%o per day to =0.257o
per day. Using a frequencyof threetimesa weekand
a protocol of 4 sets of 7 maximal effort concentriceccentricactions,Seynness13l.tlOsl
showeda rate of
increaseof only =0.72Voper day during the first l0
days,but =0.25Voper dayduring the last 25 daysof a
training period that lasted35 days. Thus, there is a
possibility that training with a frequencyof two to
three times per week may start to producea high rate
of increaseafter the first two weeks; becauseof this
and the paucity ofdata regardinglong-termtraining
with high frequencies,it is not possibleto say which
frequencyis more optimal in the long run. Nevertheless,a high frequency(four or more timesper week),
in combination with relatively non-damaginglowto-moderatevolume training,may be a good way to
'kick-start' the hypertrophy process. For longerterm training, the studies included in this review
show that frequenciesbetweentwo and four times a
week can result in CSA gains for periodsof up to 6
months.For previously untrainedsubjects,no study
was found with a training frequencyof lessthantwo
sessionsper week.
Regarding the impact of training frequency in
more advanced trainers, we found nine studies[45,46,s6,61,?6,7e,133,t34,r37]
using ScanningmethodstO
monitor changesin CSA in the quadricepsand/or
elbow flexors as a result of training in strengthtrained subjects and strength"/powerathletes.In two
preliminary reports,tTe'I34]
different frequenciesof
training were directly compared. Vikne and coworkerstTel
investigatedthe effects of squattraining
with an eccentric overload on quadricepsCSA in
subjectsthat trained one, two and three times a week
for 12 weeks. All groups performed 5 sets of 4
repetitionsin each sessionusing a squat machine,
which was loadedto 50Voof IRM in the concentric
phase and to llo-1357o of lRM in the eccentric
phase.The groups that trained two and three times
o 2007 Adis Dofo Informotion BV. All righls reserued.
ai1
per week gainedsignificantlymore in lRM strength

and quadricepsCSA (5.99% and6.75Vo,respectively) than the group that trained once a week (3.17o
increasein CSA). There was no differencebetween
training two and three times per week. Wirth and
investigatedthe effects of frequency
colleaguestl3al
on elbow flexor CSA in subjectsthat trainedonce,
twice and threetimes a week for 8 weeks.All groups
performed several types of arm curls for a total of 5
setsof 8-12 repetitionsin each session.The groups
that trained two and three times per week gained
significantly more in elbow flexor CSA (6.6% and
T.4Vo,respectively)
than the group that trainedonce
a week (3.9Vo).
The resultsof Vikne e1sl.tTeland Wirth s1al.tl3+l
are remarkablysimilar despiteusing different muscle groupsand training modes.In both reports,two
and threesessionsper week yieldedalmosttwice the
increasein muscle CSAwhen comparedwith one
session,with no apparentfurther advantagefor three
versustwo sessions.This seemslogical in view of
the typical pattern of changes in muscle protein
synthesisafter a resistancetraining session,with
peak synthesis rates observed between 3 and 24
hours, and with elevated rates sometimeslasting
between48 and 72 hoursafter exercise.t140-1431
However,it cannotbe excludedthat larger volumesand/
or different modesof training would yield different
results.Furthermore,the total weekly volumes were
not matchedbetween the groups in the studiesof
Vikne et al.t?eland Wirth et al.tr3ajAlso, it shouldbe
notedthat most studieson muscle protein synthesis
in humansafter resistanceexercisehave only studied a time spanof up to 48 hours post-exercise.
Anecdotally, many bodybuilders and other
strengthathletesonly train each muscle group specifically between one and two times per week,
sometimes even less often. On the other hand,
weightliftersareknown to perform exercisesinvolving their quadricepsfor severalsessionsper training
Teschtl45l
day.tt+el
has remarkedthat it is not known
if bodybuildingregimensare superiorto the training
regimens performed by powerlifters and olympic
lifters.Training eachmusclegroup once a week has
been shown to result in increases in muscle
SportsMed 2@7:37 (3)
242
However,
csAt7e,r34,r46l
and lean body mass.tla6,laTl
the resultsof Wirth et al.tl34land Vikne et al.tTelare
supportedby data from Mclester et al.,tla7lwho
using experienced subjects showed superior increasesin lean body massfor threetraining sessions
per week versus only one per week per muscle
group,evenwhen the total weekly volumeremained
the samefor the two groups.
Hkkinen and Kallinent6rlalso investigatedthe
effects of different distributions of the same total
volume, in a group of trained female subjectsof
which some were competitive strength athletes.
Thesesubjectsunderwenttwo three-weekperiodsof
strengthtraining for the quadriceps,with 3 training
days per week. During one period, the subjects
trained once eachtraining day and during the second
period the same subjectstrained with the sametotal
daily volume but separatedinto two sessions.During the first trainingperiod,the subjectsdid not gain
in either strengthor quadricepsCSA, but during the
secondperiod,the subjectsincreasedsignificantlyin
both maximum static strengthand CSA (4.0Voand
O.l9Voper day, respectively).Although some interesting trends can be discernedfrom the data discussedin this section, there is clearly a need for
further researchon training frequencyin both highly-trainedand less-trainedsubjects.
4.2 Intensity
The studies reviewed in this article show that
there is a remarkablywide range of intensitiesthat
may produce hypertrophy.Still, there seemsto be
some relationshipbetweenthe load (or torque) and
the rate of increasein CSA, at least for dynamic
external resistancetraining, but this relationshipis
not a straightforward one. In figure 2 and figure 10,
it can be seenthat the rates of increaseare generally
higher for intensities>6OVoof IRM than for those
<60V0of lRM, although caution is warrantedbecauseof the few datapoints<6OVoof lRM. However, it appearsthat intensitiesof =70-85V0of IRM
are sufficient to induce high rates of increaseand
that even heavier loads do not necessarilyresult in
greaterCSA gains. In the categoriesof accommodating resistance,where the effort is usually maxi-
o 2007 Adis Dqto Informotion BV. All righls reserued.
Wernbom et al
mal from the first repetition, there was no relation

betweenthe torque developedand the rate of gain in
CSA, regardlessof whethereccentricmuscleactions
were included in the analysisor not. This was true
for both the quadriceps(figure 5) and the elbow
flexors (data not shown). This is contradictoryto
someof the studiesin the literature,[48-lsll
but consistentwith other reports.[5s'62'68'6e]
It shouldbe noted that no accommodatingtraining study was found
in which the level of torque was <6OVoof MVIA.
In contrast, the study of Farthing and
Chilibecktlsll seemsto confirm the importanceof
the force developed during training for the hypertrophic response.In their study, subjects trained
their elbow flexors in an isokinetic dynamometer,
with concentricactions for one arm and eccentric
actions for the other arm. One group trained with
fast (180'/s) and another group trained with slow
(30'/s) speed,with all groups training three times
per week and progressingfrom 2 to 6 sets of 8
repetitionsover the course of the training period,
which lasted 8 weeks. The hypertrophicresponse
was evaluatedwith measurements
of muscle thicknessby ultrasound.The greatestincreasein thickness(=13Eoat the middle level) was found in the fast
eccentric group, followed by the slow eccentric
group(=1Vo),the slow concentricgroup (=57o)and
the fast concentric grovp (=2Eo).The authors interpreted their results as a confirmation of the theory
that greater force production leads to greater hypertrophy.
This interpretationmay be prematuresince the
protocols differed greatly in terms of torque-time
integral and in the volume of work performed. Furthermore, it is possible that a local overtraining
response was developing in the slow eccentric
group. Support for this possibility comes from an
earlier study by Paddon-Jonesand colleagues,t3sl
who usedvery similar eccentrictrainingregimensto
those in the study of Farthing and Chilibeck.tr5rl
Their results generally showed increasesin elbow
flexor torquefor both fast and slow eccentrictraining after 5 weeks _oftraining, but at l0 weeks the
torquevaluesof the slow group were eitherhaltedor
evenbackto the baselinevalueswhile the fast group
SportsMed 27:37 (3)
continued to gain. The authors suggestedthat the

great cumulative stress of the slow protocol had
causedan overtraining-likeresponse.The resultsof
Shepstoneet al.lr38lconfirmed the findings of Farthing and Chilibecktrsrlregardingthe superiorityof
fast versus slow eccentric training for the hypertrophic responseof the elbow flexors. However, this
study is open to the same interpretationregarding
the possibilityof a local overtrainingresponsein the
slow eccentricgroup, sincethe differencein torquetime integral between the fast and slow protocol was
even greater(>1O-fold)than in the preceedingstudies. Moreover, the slight differencesin peak torque
betweenthe fast and the slow eccentric velocities in
the studiesmentionedhere arguesagainstthe level
of torque as the primary explanation for the differences observed in the hypertrophy of the elbow
flexors. Recruitment differencesbetween fast and
slow eccentric velocities cannot be excluded as a
contributing factor; however, at presentthere is not
enoughevidenceto supportthat this occurs.Nevertheless, the considerable difference in hypertrophy
between fast eccentric and both fast and slow concentric training in the study of Farthing and
Chilibecktl5rllends support to the hypothesisthat
the force developedby the muscleduring training is
an important factor for hypertrophy.
At present,there are no accommodatingresistancetraining studiesthat have investigatedthe impact of different eccentric velocities and/or different
levels of torquedevelopmentduring eccentricmuscle actionson the CSA of the human quadricepsas
measuredby scanningmethods.There is also a lack
of direct comparisonsusing accommodatingconcentric training at different velocities and/or different levels of torque development.The studiesreviewed in this article suggestthat hypertrophycan
be inducedwith a rangeof concentricvelocities.The
upperlimit of concentricvelocity that is still capable
of inducing hypertrophy is not known, but type I
fibre hypertrophy has been observedafter training at
2400lsttsz)
and type 2fibre hypertrophy after training
While being quite
at velocitiesas high as 300o/s.tls3l
high compared with the cadenceof conventional
resistancetraining, thesevelocitiesare still well be-
o 2007 Adis Dolo Informolion BV. All righls reserued.
low the reported maximum velocity of unloaded knee extensions,which may reach values of
=JQQo/5.tlsal
In supportof the use of moderatelyfast concentric training, an early study by Thomed s1 al.lrssi
showeda clear trend (but not significant)for hypertrophy of type 2 muscle fibres (=30-35V0)in the
vastus lateralis in both the healthy and the injured
limbs in individuals undergoing moderately fast
(180'/s) isokinetic concentrictraining after reconstruction of the anterior cruciate ligament (ACL).
No such trend was apparentin either limb in the
group that trained at a low velocity (60'/s). The
training was carried out three times per week for 8
weeks, with the slow velocity group progressing
from 3 setsof l0 repetitionsto l0 setsof l0 repetitions and the fast group from 3 setsof 15 repetitions
to l0 setsof l5 repetitionsduring the time courseof
the study. A CT scan was also performed of the
quadricepsand vastus lateralis at mid-thigh level,
but unfortunatelytoo few scanswere availabledue
to problemswith the scanner,thus making it impossible to confirm or reject the trend ofthe fibre CSA
dataat the whole musclelevel. However,a studyby
Frobseand colleaguestl56l
also supportsthe use of
moderatelyfast concentrictraining for the quadriceps, at least in the rehabilitation setting. These
authorsinvestigatedthe effectsof isokineticconcentric training in patientsthat had undergonereconstruction of the ACL and they showed hypertrophy
of the quadricepsin responseto moderate(150'/s)
and fast (240"|s)protocols,which was at leastequal
to that of the slow protocol (60'/s). Thus, it appears
that hypertrophycan be inducedin the humanquadricepsas a result of concentrictraining at velocities
of up to at least 24o"ls and torque levels as low as
=50Voof MVIA.
However, the largest increase (=l\Vo at midthigh level), as well as the by far highest rate of
increase in quadriceps CSA (0.447o per day) for
accommodatingtraining, was noted in the study by
who trained subjectswith 5 sets of 10
Rafeei,troal
concentricmuscleactions at 9OVoof the maximum
torque at 60o/s,three times per week for 6 weeks.An
was that
interestingfeatureof the studyof Rafeeitl0al
244
the subjects trained at the same absolute torque

throughoutthe study. Another notable feature was
the generousrest periods, with 5 secondsbetween
each concentric action and 120 secondsbetween
each set. The level of effort was probably high
enoughto recruit most of the motor units, while the
high but not maximal force level and the generous
rest periods may have minimised muscle damage
and fatigue of fast fibres, thus allowing the muscle
to hypertrophy already in the initial stagesof training.
Wernbom et al.
relativelyhigh tensions.It is also possiblethat some

fast fibres are preferentiallyrecruitedduring eccentric actionsand that this may occur at loadsas low as
25Voof MYlA.tr58l16ut, it may be too simplisticto
estimatethe stressimposed on each muscle fibre
merely by the magnitudeof the externalload.
In summary, although there may well exist a
level oftension below which no hypertrophyoccurs,
the relationshipbetween the training load and the
hypertrophic responseis complex. Achieving recruitmentof the greatestpossiblenumberof motor
units in the targetmuscle(s)and making thesemotor
units fire at high rates and for sufficient lengthsof
time are obvious prerequisitesfor inducing significant hypertrophy. Still, it appears that maximal
loads are not necessaryto ensure that these conditions are met providing that the training is performed with close to maximum effort in at least one
of the sets.Thus, the resultsof this review support
with intensitylevelsof
the typical recommendations
70-85V0of maximum when training for musclehypertrophy, but also show that marked hypertrophy is
possibleat both higher and lower loads. However,
placinghigh mechanicalstresson the working muscle may result in local overtrainingif the durationof
work is long. Some of the possible interactions
betweenthe level of tension,duration of exercise,
mode of exerciseand muscle damagewill be discussed in section 4.6. The impact of intensity in
more advancedathletesremains poorly defined due
to the lack of objectivescientific data.
Regardingdynamic external resistancetraining,

the rangeof intensitiesthat can producehypertrophy
is even more remarkable than in the caseof accommodating resistance. 5g6i"rt83'8al have shown
markedincreasesin CSA in responseto loadsas low
as 20Voof IRM when the exercisehas been combined with partial restriction of the blood flow by
meansof thigh torniquets.Even so, considerationof
the recruitmentof motor units during fatiguingexercise with low loads reveals that the results of the
studiesof Takaradaet al.t83land Abe et al.t8aldo not
necessarilydisprove the theory that tension is a
major determinantof the hypertrophicresponse.A
study by Greenhaff and co-workerstlsTlshowed a
greatly increasedrate of glycogenolysisin type I
fibres and a marked decline in force and near total
depletion of phosphocreatine in both fibre types
during intermittent electrical stimulation of the
quadricepswith the blood flow occluded.In contrast,the declinein force during the sameprotocolof
stimulationbut with intact circulation was ascribed
4.3 Volume
almost solely to fatigue in type 2 fibres. Although
Greenhaff et al.tl571used electrical stimulation inA notabletrend in the severaltypes and modesof
stead of voluntary activation, their findings are of
strength training reviewed in this article was the
relevancefor the developmentof fatigue under cir- occurrenceof a plateau in the hypertrophic adaptacumstanceswhere the blood supply to the working tions after a certain point of volume or durationof
muscleis limited, for exampleby a tourniquetand/ work had beenreached.In someof the results,there
or by the raised intramuscular pressureduring the is evena suggestionof a declinewhen the volume or
continuously performed coupled concentric-eccen- durationis extendedbeyondthe point of the plateau.
tric muscle actions that conventional resistance Again, it must be noted that no studieswere found
training usually consistsof. With a decline in force that investigatedthe effectsof I or 2 setson muscle
in type I fibres, more type 2 fibres would have to be CSA or muscle volume of the quadricepsor elbow
recruited and towards the end of each set. the re- flexors.That said,figure I l, for the total repetitions
maining force-producingfibres could be exposedto for DER training of the elbow flexors, suggestsa
o 2m7 Adis Doto Informolion BV. All ights reserved.
SportsMed 2m7;37(3)
245
dose-response
curve where greatergains in muscle
massare noted initially with increasingvolume (or
duration) of work, but with diminishing returns as
the volume increases further. Overall, moderate
volumes (=30-60 repetitionsper sessionfor DER
training) appearto yield the largestresponses.
musclethicknesswere demonstratedafter as little as

I set of 8-12 repetitions of specific exerciseper
musclegroup.Becauseof this and given the relative
paucityof data,especiallyregardingthe early part of
the volume continuum (i.e. l-20 total repetitions),
there is clearly a need for further researchon the
impact of training volume on whole muscle CSA.
However, two notableexceptionstl25'1261
also appear in figure 11, which demonstratethat high rates This appearsto be true for both previouslyuntrained
and well-trainedindividuals.
of growth can be achievedwith a relatively small
However, recent data from Ronneshd s1 al.tt6zl
number (=12-14) of repetitionsper sessionunder
provide further support to the notion of a dosesomecircumstances.
In the first of theses1udiss,ll25l
responserelationshipbetween the training volume
very high loads (=90-l00Vo of IRM) were usedfor
and the hypertrophic responseof the quadriceps.
both the concentricand the eccentricphasesand in .
These
authorsreported superior increasesin quadrithe other study,ttz6lextremely high loads (progresscepsCSA for a total of 6 sets(ll3qo) versus2 sets
ing from 130 to 230Voof IRM) were used for the
(7,6Vo)of quadricepsexercise(two exercisesof I or
eccentricphase.As can be seen in figure 10, the
3 setseachat 7-l0RM) at an exercisefrequencyof
majority of the other datapoints were distributed
three times per week for I I weeks. It deservesto be
between 60 and 9OVoof lRM. A further example
noted that the subjectsin this study received a protedemonstrating that significant hypertrophy can be
in supplementprior to each workout. Becauseit is
inducedwith a surprisinglysmall numberof muscle
currentlyunknown how protein supplementation
inactions at very high loads, at least in previously
teractswith training volume, theseresultsmay not
untrainedsubjects,can be found in a study by Hawnecessarilyapply to strength training that is perkins et al.,tl48lwho showedthat a total of 9 maximal
formed without supplementation.
eccentric muscle actions was sufficient to induce
significant increasesin thigh lean mass, while 12
4,4 Mode of Troining ond Type of
maximal concentricactionswas not. Thus, the relaMuscle Action
tionship between volume and the hypertrophicresponsemay differ betweendifferent levelsof torque
In the scientific literaturerelating to the area of
and/or types and modes of strength training. The
resistancetraining, one sometimesfinds categoric
discrepancybetweendifferent studiesin terms of the
statementssuch as 'eccentrictraining producesthe
volume neededto induce hypertrophy may, in part,
greatestmuscle hypertrophy'. This review demonbe related to differences in the total duration of
stratesthat given sufficient frequency,intensityand
muscleactivity per session.In many studies,neither
duration of work, all three types of muscle actions
the velocity nor the durationofeach repetitionwere
can induce significant hypertrophy at impressive
reported.
rates and that at present,there is insufficient eviTo date,relativelyfew studieshavedirectly com- dencefor the superiority of any mode and/or type of
paredthe effects of different volumes of work on the muscleaction over other modes and types of trainhypertrophic response as measured by scanning ing in this regard.Using dynamicexternalresistance
methodology.Thesefew studiestl6'l5e-l6ll
usedless training as an example, one would be tempted to
accuratemeasuresof muscle massratherthan mus- concludethat, if anything,pure eccentrictraining is
cle CSA or volume, or scannedonly parts of the actually inferior to both concentricand concentricmuscle groups, and it is therefore difficult to com- eccentrictraining, as judged by the degreeand rate
parethesewith studiesin which whole musclescans of hypertrophyobservedin the studiesincluded in
were performed with MRI, CT or UL. However, in
this review. If one insteadconsidersconcentricand
two of these studies,tl5e'1601
significant increasesin
eccentrictraining with accommodatingresistance,
O 2m7 Adis Doto lnformotion BV, All rights reserued.
Wembomet al.
If force is an importantstimuli in resistancetrarnmaximal eccentrictraining has been slightly more

effective than maximal concentric training in the ing, it follows that a certain level of torquemust be
few studies that have directly compared these two
reachedfor some minimum duration for significant
types of training. However, the hypertrophic re- hypertrophy to occur. In the pioneering study of
who comparedpure consponse has been modest in many of the studies Jonesand Rutherford,t55l
comparingthe effectsof pure concentricversuspure centric and eccentricregimensusing a variableresistanceknee extensiondevice, the authors noted
eccentrictraining and this appearsto be true both for
accommodatingresistanceand dynamic externalre- that the subjectsactivation of the quadricepswas
sistancetraining studies. Thus, the protocols that high only near the position of full knee extension.
Further inspection of their electromyogram data
have been compared may not have been the best of
each type. Again, it is noteworthy that both the suggeststhat the eccentric training was accompalargesttotal increasein CSA (l8.4Vo)and the high- nied by slightly less activation than the concentric
est rate of increasein CSA (O.44Voper day) for the training and also that the duration of high activity
quadricepsfor the category of accommodatingtrainwas shorter.Thus, it is possiblethat less than optiing was noted for a pure concentric training mal internal force production and short total duragroup.tlo4lThis was also the secondhighestrate of
tions of high activity contributedto the finding of no
increasefor any mode of quadricepstraining, sur- difference between the eccentric and concentric
passedonly by the shorterstudy of Abe et al.t84lThe training, and to the modest hypertrophy for both
findings of Rafeei,tr@lof greaterhypertophyat the protocolsin their study. The eccentricand concenwhole muscle level as well as at the muscle fibre tric exerciseregimens in the studies of Housh et
may have sharedthe sameproblem, as the
level for near maximal concentricversus submax- al.t68'6el
imal eccentrictraining,expandedon an almostiden- resistancein their device was greatestnear full extension.Nonetheless,the studiesof Jonesand Ruthtical study from the same researchgroup,tto3lin
which greater fibre hypertrophy was found for con- erford,ts5lSmith and Rutherfordt62land Housh et
show that at least for the mode of dynamic
centric training when compared with eccentrictrain- al.t68'6el
ing when both regimens were performed at the same externalresistancetraining,the greaterloadsthat are
possible with eccentric training (compared with
torque level.
concentrictraining) do not necessarilytranslateinto
The divergencein the resultsof concentricversus
greatergainsin musclesize.
eccentrictraining betweendifferent modes(DER vs
Among the accommodatingmodes,the isoineraccommodatingresistance)may be due to differencesin the characteristicsofthe resistancefor each tial flywheel knee-extensionmodel of Tesch and
hasso far consistentlyinduced
mode. As discussedin the introduction,when using colleaguest2T'87'105'll8l
external resistance(e.g. free weights, weight ma- hypertrophy of the quadricepsCSA at high rates
chines), the torque is not necessarily optimally (0.174.22Voper day). It is not immediatelyobvious
matchedthroughoutthe movementto the individu- why this mode seemsto be more effectivethan most
al's strengthcurve. Herzog et al.tto+lcalculatedthat of the isokineticregimensthat have also usedmaxrmal eccentricactions.In the flywheel studyof Tesch
the internalforcesofthe three vastusmusclesofthe
quadricepsare at their highest at knee angles of
et al.,Iz1lthe subjectswere instructedto resist only
=60-80o of flexion (full extensionis definedhereas gently during the first part of the eccentricaction
and then apply maximum force. The torque-angle
0o of flexion), whereafter the forces drops to lower
curvesin the samestudytzrtshow that high eccentric
levelswith decreasinganglesof flexion. At 0-20'of
=2040Eo
maxtforces were reachedonly during a rather short arc of
forces
are
low,
only
of
flexion, the
=20-25", from =65-90o offlexion. In contrast,durIchinose
s1
3l.tr6sl
reported
that
the
mum. Similarly,
ing isokineticeccentricexercise,maximum effort is
vastus
lateralis
was
maximal
at
70o
of
force of the
gslusually
applied from the start and dn16ll66't671
flexion.
@ 2m7 Adis Doto Informotion BV. All rights reserued.
Sports Med 2m7j 37 (3)
lected at similar average velocities show that high

eccentricforcescan be achievedthroughan arc ofat
least 40o. However, maximum eccentric efforts at
extendedknee angles are often perceived as uncomfortable.lr68lIn the isokinetic study of Seger et
al.,tlO2l
four of five subjectsin the eccentrictraining
group had frequentcomplaintsof knee pain during
the training, which may have affected the hypertrophic adaptations. Interestingly, Holder-Powell
and Rutherfordt168l
showedthat much of the discomfort associatedwith maximum isokinetic eccentric
exercise could be avoided if the subjects started to
resistlater in the arc of motion, from =45oof flexion
insteadof 15" of flexion. Also, the eccentricpeak
torque was significantly higher when the range of
motion was 45-95o, comparedwith l5-95o of flexion. It could be that the relatively low volumesand
short ranges of high-force eccentric exercise in the
flywheel studies worked in favour of producing
hypertrophy, whereasthe isokinetic-eccentricprotocols may have resultedin too much stressand strain
on the tissues.
ations can result in far greaterpeak loads than the

nominal load.tl2elThe significance,if any, of these
very high but momentary forces for the hypertrophic
responseremains to be explored. We speculatethat
differences in torque profile, torque-time integral
and motor unit recruitmentaccountfor someof the
differences in the hypertrophy observed between
studieswhere the training variableshave beennominally similar. Also, if strenuouseccentrictraining is
performed at a high frequency, the hypertrophic
responsemay becomecompromised. These interactions will be discussedin sreaterdetail in section
4.6.
In summary,all modesreviewedhereseemcapable of inducing marked hypertrophy, at least in the
short term. This is not to say that some method or
combinationof modeswill not emergeas superiorin
the long term. The ideal proportions between the
different types of muscle actions are still a subjectof
debateratherthan a scientific certaintv.
Other factors to consider when comparing the

isokineticand flywheel modesare the accelerations
and decelerationsthat occur in the latter mode but by
definition not in the former. It hasbeenhypothesised
that accelerativeand decelerativeforces are important componentsof the stimulus for muscle hypertrophy in resistancetraining.tl0elTo date, there is
little evidenceto supportthis hypothesis.Collectively, the successes
of both the dynamic,isokineticand
isometric modes in producing muscle hypertrophy
does not appear to support accelerationsand/or decelerations as being particularly important for the
hypertrophic response.However, although the angular velocity in isokinetic training is controlled,the
fascicle velocity in the working quadricepsvaries
markedly through the range of motion.tlT0lFurthermore, isokineticsusually involve a brief build up of
speedbefore the isovelocity phase is reachedand a
short braking period after the isovelocityphase.tlT0l
Thus, from a muscle point-of-view, accelerations
and decelerationsoccur even during 'isokinetic' exercise. It should also be noted that with weightbasedresistancetraining, accelerationsand deceler-
Becausetoo many studiesdid not report the rest

periodsbetweensets(and repetitions),we optednot
to try to evaluateany trends.However, someelaboration regarding the potential impact of rest periods
is possible.Closely associatedwith rest periods rs
the role of fatigue in strength training. Regarding
strength,some studiestlTl'1721
have shown that short
rest periodsbetweensetsand/or repetitionsare superior to longer ones, whereasother studiestlT3l
have
concludedthat longer periods are superior to shorter
ones, while yet other studiestlTal
have reportedno
difference.Upon closerexamination,it appearsthat
when maximal or near-maximalefforts are used,it is
advantageousto use long periods of rest. This is
logical in light of the well known detrimentaleffects
of fatigueon force productionand electricalactivity
in the working muscle. If high levels of force and
maximum recruitment of motor units are important
factorsin stimulatingmuscle hypertrophy,it makes
senseto use generousrest periodsbetweensetsand
repetitions of near-maximal to maximal efforts.
It is interestingthat for the accommodatingand
isometriccategories,the studiesin which the highest
@ 2m7 Adis Dsto Infomotion BV. All rights reserued.
4.5 RestPeriodsond the Role of Fotigue
Med 2m7;37(3)
Sports
248
Wernbomet al
rates of muscle growth were foundtl04'1061

did include long rest periods. Furthermore,in the DER
training studytezlthat reported the highest rate of
CSA increase(O.26Voper day), very long rest periods (10 minutes) betweenworking sets were used,
but in this study the volume was periodised,which
may also have impacted on the results. It is also
worth noting that maximum isokinetic-concentric
exercise performed with little rest between each
muscleactionis associatedwith a markeddeclinein
peak torque during each working set, whereaslittle
or no decline occurs during maximum eccentric
exercise.tlT5-1771
Hence,it can be hypothesisedthat if
the rest periods are too short during near-maximal
concentricexercise,the training effectswill be compromised.
latter group trained with considerablyless effort in

comparisonto the first group. The resultsshoweda
dramatic difference in the extent of hypertrophy in
favour of the group that trained in a continuous
manner(l2.9%oincreasein quadricepsCSA), versus
the groupthat restedin the middle of each set(4.0Vo
increasein quadricepsCSA). The authorsspeculated that both increasedrecruitment of motor units
and a greateracute hormonal responsecould have
contributed to the greater hypertrophy seenafter the
continuousprotocol. However, along with the rncreasedstresson the muscle with shorterrest periods at submaximal resistance,the potential for overtrainingmay alsoincrease.In a studyby Follandand
colleagues,tlTal
conventionalresistancetrainingwith
multiple sets to muscular failure and very short rest
Although eccentric exercisegenerally produces periods (30 seconds)led to considerabledelayed
little acute fatigue, it appearsthat it is dependenton onsetmusclesorenessduring the first week of trainthe trainingvelocity (and probablyalso the work-to- ing. With this type of training, caution with the
rest ratio), with fastereccentricvelocitiesproducing training frequencyand volume appearsto be warless fatigue than slower velocities (see Tesch et ranted.
al.lr75lfor a discussion).The difference in acute
The impact of rest periods may extend beyond
fatiguedevelopmentbetweenconcentricand eccenthe
effects on fatigue and motor unit recruitment.
tric muscle actions and also betweenfast and slow
Using
a rat muscle model, the researchgroup of
eccentricmuscle actionshave obvious implications
Faulkner
has,in a seriesof studies,llT8-1801
investigatfor comparisons between these modes in regard to
ed
the effects of electrical stimulation againstthe
training effects. On the other hand, when using
submaximal resistance,the size principle dictates deleteriouseffects of denervationon muscle mass.
that motor unit recruitmentand firing ratesareprob- They showedthat denervatedmuscle is sensitiveto
ably far from maximal until the muscle is near both the total number of muscle actions and the
distribution of loading. For example, 100 muscle
fatigue or unless the repetitions are performed with
the intentionto executethe movementvery quickly. actionsper day generatedat a constantintervalover
24 hourswas sufficientto maintainmusclemassand
The importanceof exercisingwith near-maximal
effort when using submaximalresistancein conven- force, but the same number of muscle actionsdisjust 4 hoursper day (and consequently
tional strength training has been elegantly demon- tributedover
20
hours
of
rest
in between)failed to maintain mass
stratedby Goto and co-workers.l85l
In their study,
and force.Although thesefindings may not necessatwo groups of untrained subjectsperformed 5 setsof
l0 repetitionsof dynamic knee extensionsat a load rily extrapolateto intact innervatedhuman muscle,
they show that skeletalmuscle,at leastunder some
of lORM (=75Voof IRM), two timesper week for l2
circumstances,
is sensitiveto both the total number
weeks. One group performed all l0 repetitionsin
eachset in a continuousmannerto muscularfailure, of muscle actionsand the distribution of them. Fuwhile the other group performed 5 repetitions and ture studiesshould examinethe potentialimpact of
then rested for 30 seconds before performing the both shorter(seconds)and longer (minutes,hours)
remaining5 repetitions.Thus, althoughthe volume rest periods on skeletal muscle hypertrophy and
was matchedbetweenthe groups, the subjectsof the hypertrophicsignalling in this light.
@ 2007 Adis Doto Informotion BV. All rights reserved.
SoortsMed 2m7;37(3)
4.6 Interoctions Between Frequency,

Intensity,Volume ond Mode
After acknowledging that the training volume
seems to influence the hypertrophic responseto a
certain extent, the question arises of what aspectof
volume is the most important determinantof this
response?Is it the total mechanicalwork performed
or is it the time-tension integral of the activity?
Basedon the availableevidence,we suggestthat the
time-tensionintegral is a more importantparameter
than the mechanicalwork output (force x distance).
The elbow flexor eccentric training studies of
Farthing and
Refsnes,tl261
Paddon-Joneset a1.,1381
Chilibecktr5rland Shepstonss1 4l.tl38Jprovide in_
sight into some of the complex interactionsthat
takes place between the training variables of frequency, intensity, volume and mode of resistance
training. If one considersthe last two of thesestudies, one finds that the external work output (expressedas total number of repetitions x the torque
developed) was likely to be similar between the
slow and fast eccentric groups becauseof the marginal differencein maximum torquebetweenthe slow
and the fast velocities.In contrast,the total duration
and the torque-time integral between the groups
were vastly different, =6-10-fold greater for the
slow groups compared with the fast groups. The
studiesof Farthingand Chilibecktrsrland Shepstone
et al.tl38lare difficult to comparewith eachother in
termsofthe degreeofhypertrophy achievedbecause
different measuresof musclemasswere used(muscle thicknessvs muscleCSA). Still, the hypertrophic
responsenoted for the fast eccentric group appears
to be larger in the Farthing and Chilibeck studyttstJ
(=l3Vo increasein muscle thickness)than the 8.57o
increasein CSA reported by Shepstones1 3l.tl38l
Becausethicknessmeasuresonly one dimensionof
the muscle, the increasein elbow flexor CSA was
likely to be greater than l3vo. If the muscle grew
equally in width as it did in thickness,the result
would be an increasein CSA of 27.1Vo.This scenario seemsunlikely, becausea tricepstraining study
by Kawakami et al.tlsu showed that an increaseof
3l.7Vo in elbow extensorareawas accompaniedby
an increasein thicknessof 27.OVo.Ifthe sameratio
o 2007 Adis Dolo Informotion BV, All rlghls reseNed.
249
of =1.17betweenthe increasein CSA andthickness

is applied to the elbow flexors in the study of Farthing and Chilibeck,tl5llone arrivesat an estimated
increasein CSA of =157o for the fast eccentric
training group.
The traininggroupsof Farthingand Chilibecktrsrl
progressedfrom2 setsof 8 repetitionsto 6 setsof 8
repetitions,resultingin an averagetotal durationof
muscle activity per sessionof 22 and 132 seconds
for the fast and slow eccentricgroups.respectively.
In the study of Shepstoneet al.,tl38lthe progression
was from I set of l0 repetitions to 4 sets of l0
repetitions and the corresponding average total
durationsof muscleactivity per sessionwere 14 and
146 secondsfor the fast and slow eccentric groups,
respectively.We suggestthat the slightly greater
total duration for the fast group in Farthing and
Chilibecktlsll versus Shepstoneet al.tl38lwas responsiblefor the greater hypertrophicresponsein
the former study.On the other hand, with the slowvelocity protocols,the cumulativedamagingeffects
of the long durationsof maximum eccentricexerclse
may have counteractedthe hypertrophy so that this
becameless in comparison with the fast training.
The study of Refsnes,l126l
using a dynamic constant
resistancetraining model in which the eccentric
phasewas overloaded(progressingfrom 1307oto
=230Voof IRM during the time courseof the study)
also atteststo the effectivenessof short durations of
maximum eccentricexercisefor inducing increases
in elbow flexor CSA. In this study, the volume was
carefullyprogressedfrom 2 setsof 2 repetitionsto 5
setsof 4 repetitionsduring the 8 weeksof the study,
resultingin a maximum durationof =14-16 seconds
of near-maximaleccentricwork. The velocity in the
eccentricphasewas moderate,=8G-90"/s.The concentric phasewas loaded with only 307o of lRM,
and the contribution of the concentricphaseto the
hypertrophicresponsewas thereforeprobably small.
The subjectsincreasedtheir elbow flexor CSA by
2l.5Vo(0.38Voper day), an impressiveincreaseespecially when consideringthe very briefduration of
work.
The risk for overtrainingwith long durationsof
high-forceeccentricexerciseis supportedby a study
SporlsMed 2@7;37(3)
250
by Amiridis et al.,tr82lwho compared different

modesof training in a group of young elite female
basketball players after a period of very strenuous
training for the knee extensors.During the first 12
weeks,all subjectsperformed8 setsof 8 concentric
repetitionsat10%oof lRM and 8 setsof 8 eccentric
repetitions at llo%o of IRM in the leg press, 4
sessionsper week. At 12 weeks, the subjectshad
significantly reducedperformancesin both the leg
press and the countermovementjump, indicating
that they were overtrained.During the second 12
weeks of training, the subjects were divided into
three groups that performed different modes of recovery training,4 sessionsper week.The first group
trainedwith 8 setsof 8 concentricrepetitionsatTOVo
of lRM; the second group completed4 sets of 8
concentricrepetitionsat 7}Vo of I RM and 4 setsof 8
eccentricrepetitionsat llo%o of IRM; and the third
group performed 8 sets of 8 eccentric repetitions at
llo%a of lRM. Comparedwith valuesfrom the first
12 weeks overtraining, all groups increasedtheir
performance in the leg press and the countermovementjump, but only the pure concentricgroupnoted
significant increasesin leg-press strength (39Vo),
isokinetic strength (ll43Eo) and vertical jump
(l1vo) in comparisonwith the pre-trainingvalues.
Although no morphological data was presentedin
this study,it is likely that somedegreeof overtraining at the muscle level was responsiblefor the poor
performanceat 12 weeks. It is also interestingto
note that despite reduced total volume in comparison with the first 12 weeks,neitherthe pure eccentric nor the combined concentric-eccentricgroups
experienced any supercompensationin performance,whereasthe pureconcentricgroupdid. Thus,it
would appear that moderate-forceconcentric training was better tolerated than high-force eccentric
training, at least for the moderatelyhigh volumes
and the rather high training frequencyused in this
study. Taken together,the results of these studies
support the common recommendation of using
somewhatlower frequenciesand volumesfor highforce eccentricexercisethan for conventionalresistancetraining.
o 2m7 Adis Doto Infomotion BV. AII righls reserued.
Wernbomet al
On the other hand, studiesby LaStayo and colleaguesll83'ta+l

using eccentric cycling at submaximal intensitiesfor long durations(=20-30 minutes)
have shown very rapid and large gains in MFA
(=5U60Vo). Recently, a case reporltl8slwas published that showedthat marked hypertrophyis evident also at the whole musclelevel after this type of
training. In theseregimens,up to =100G-2000eccentric muscleactionswere performedduring each
training session,threetimes per week. The absolute
intensitylevel was reportedin watts,so it is difficult
to quantify the forces in terms of percentage of
MVIA. Nevertheless,thesestudiesl183-1851
show that
given careful and gradual progressionof exercise
intensity and duration, human skeletalmuscle can
tolerateand adaptto prolongedsubmaximal-eccentric exercise.
Overall, we feel that the trendsobservedin this
review are consistentwith the model for trainingovertrainingcontinuumproposedby Fry,tt8el*6"r"
the optimal training volume and also the volume
thresholdfor overtrainingdecreaseswith increasing
intensity. The study of Abe s1 3l.t8alis especially
intriguingin this contextbecauseof the combinatron
of very low intensityand extremelyhigh frequency.
To the best of our knowledge, no studies to date
have investigated the effect of performing two
strengthtraining sessionsduring the same day on
skeletalmuscleprotein synthesis,so it is not known
whetherthere is any additional benefit in doing so
comparedwith performingjust a single session.If
the sensitivityof skeletalmuscleproteinsynthesisto
mechanicalstimuli is regainedduring the sameday
and if thereis room for further elevationsof the net
protein synthesis,then it would make senseto perform more than one sessionper day. This could
explain the results of Hkkinen and Kallinen,l6ll
although the effects of tapering down the volume
(and hence,the total stressper sessionimposedon
the muscle)alsoremain a possibility.As pointedout
in the discussionconcerning rest periods (section
4.5), mechanisticinvestigationsconcerningthe effects of different _distributionsof loads and rest
periodson skeletalmusclemassand/orintracellular
hypertrophicsignallingare largely lacking.It is also
SpodsMed 27:37 (3)
uncertainwhether the mechanosensitivityof skeletal

muscledecreases
with long periodsof strengthtraining. Until these and other dose-response
relationships become more fully characterised,it will remain difficult to prescribe a proper 'dosage' of
training for eachmode of training and type of muscle action for the specific purposeof inducing hypertrophy.
Regarding training for hypertrophy in already
highly-trainedindividuals,thereis at presentinsufficient datato suggestany trendsin the dose-response
curvesfor the training variables,It hasbeensuggested by some authorstl8Tl
that the volume neededto
induce optimal gains in strength, increaseswith
training status,so that advancedtrainers and elite
athleteswill have to perform far more sets(>10 sets
per muscle group) than untrained and recreational
lifters (={-5 sets per muscle group). Other authors
emphasisethe importance of load and the type of
muscleaction.Refsnestls8l
has reportedpreliminary
findings from unpublishedstudies,which indicate
that very well-trained athletes respond to eccentric
overload training with greater hypertrophy than after conventional training. Recently, a study by
Vikne and colleaguestl3Tl
demonstratedsignificantly
larger increasesin elbow flexor areain well-trained
individuals after pure eccentric training (ll%o increase)than after concentrictraining (3% increase).
It shouldbe notedthat the volume was not equalised
betweenthe groupsand it also seemslikely that the
total duration of work was markedly longer for the
eccentricgroup,thus resultingin largedifferencesin
time-tensionintegral betweenthe protocols.Hence,
although the results suggest a clear superiority of
pure eccentricexerciseversuspure concentricexercise for inducing hypertrophy in well-trained subjects, other variablescannotbe ruled out as contributing factors.Seemingly at odds with the observations of Refsneslls8land Vikne et al.,tl37]
Brandenburgand Dochertytl33lshowed no difference in muscle CSA after eccentric-overloadtraining for the elbow flexors and the elbow extensors
compared with conventional training. In their
study,tl33lcoupled concentric-eccentricrepetitions
were performed in both groups and the total volume
@ 2007 Adis Doio Informotion BV. All righls reseNed.
251
was similar between the groups. Needlessto say,

more researchis neededregardinginteractionsbetween variablesin both trained and untrainedsubjects.
Finally, a comment on the interactionsbetween
strengthand endurancetraining is warranted.It has
been recommendedthat both strength and endurancetraining should be included in a well-rounded
training programme.tlselThe additive effects of
strengthand endurancetraining on variousparameters of health status have been shown in several
studles.lleo-le2l
Recently, it has been demonstrated
that while strength training by itself can lead to
increasedarterial stiffness, this negative effect can
be offset if endurancetraining is performed concurrently.tte3lFrom a strengthtraining point of view,
there is an interest in how to train concurrently
without affecting strength and hypertrophy negatively. It has been suggestedthat strengthtraining
should be performed first, in order not to compromise the quality of the strength-trainingsession.treal
However,this order may not necessarilybe the best
choice for inducing increases in muscle mass.
Deakintre5linvestigatedthe impact of the order of
exercisein combinedstrengthand endurancetraining and reported that gene expressionassociated
with muscle hypertrophyrespondedmore strongly
when cycling was performedbefore strengthtraining, insteadof vice versa.Interestingly,in the study
of Saleet al.,trI rl performingcycling first seemedto
induce the greatestincreasein muscle area. Still,
becausethe lack of studiesinvestigatingthe effects
of the order of exercise in concurrenttraining on
hypertrophy,no firm conclusionscan be drawn on
this issue.
4,7 Time Course of Muscle Hypertrophy
Strengthgainsas a resultof a period of resistance
training are usually attributedto two major factors:
(i) neuraladaptations;and (ii) hypertrophy.tre6l
Until
recently,the prevailingopinion has beenthat neural
adaptationsplay the dominant role during the first
6-7 weeksof training, during which hypertrophyis
usuallyminor. However, as notedby Staronand coworkerstle6land by Sale,tleTl
it appearsthat the hySports
Med 2@7:37(3)
Wernbomet al.
pertrophy processbegins earlier than this, as trends

for increasedfibre CSAs can be observedat 2 weeks
into the training period. In the studies of Mayhew et
al.tl631
and Rafeei,tl0alsignificant increasesin fibre
CSA (type | = l2-l4%o; type 2 = 26-28Vo) were
observedfor the concentrictraining groupsas early
as 4 weeks into the training. In the latter study,llO4l
further hypertrophy had occurred at 6 weeks, which
was also manifested at the whole muscle level
(=l8Vo increase in quadriceps CSA at mid-thigh
level). In the study of Abe et a1.,t841
significant
increasesin muscle volume were noted afterjust 2
weeks
and
other
investigaseveral
have also demonstratedsig_
1isnst13,27,54,87.105.118,1281
nificant hypertrophy at the whole muscle level after
short periodsof training (3-5 weeks).Thus, thereis
now plenty of evidencethat significanthypertrophy
can take place early on given proper frequency,
intensity and volume of training.
that the majority of the increasetook place during

the first l0 weeksof training.
Little is also known about how different training
variablesand modes interactwith the length of the
training period. In some strength-trainingstudies,
the increasein muscle volume is delayed,while in
others,the rate of growth is rapid. We speculatethat
less-damaging
training modesmay allow the hypertrophy responseto start earlier. Regimens that include eccentricmuscle actions,especiallythoseinvolving maximal effort, appear to require a careful
initiation and progressionof training to avoid muscle damageand muscle protein breakdown.In line
with the resultsof Foley et a1.,t2001
who noteda longlastingdecreasein elbow flexor musclevolumeafter
an acute sessionof high-force eccentric exercise,
Willoughby e1 al.t2orlfound decreasedmyofibrillar
protein content in muscle biopsiestaken from the
vastus lateralis after an acute session of 70 nearmaximal eccentricactions for the knee extensors.
Based on the observation of positive muscleThis decreasewas accompaniedby increasesin casprotein balance after an isolated sessionof resispase 3 activity and in the expressionof ubiquitin,
tanceexercise,Phillipstle8lhas proposedthat a gain
which the authors interpreted as indicating that
in active force-producing myofibrillar proteins
apoptosisand increasedproteolysishad occurredin
could occur after a single strength-trainingsession
the exercisedmuscle.They also reporteda repeated
and that this increase may take place without a bout effect for most of the parametersafter a second
change in fibre CSA. In line with this idea, Wilsessionofan identicaleccentricprotocol.Nevertheloughby and Taylorlleelreportedan increasein myless,a trend towardsdecreasedmyofibrillar content
ofibrillar content in muscle biopsiesobtainedfrom was evidenteven after the secondsession,although
previously untrained young men after just three this was not significant and certainly of a smaller
strength-trainingsessions,with sessionsseparated magnitudethan after the first session.In contrast,in
by 48 hours of rest. As argued by Phillips,tle8lthe the study of Willoughby and Taylor,treelwhere a
idea that early gains in strengthare due exclusively conventionalresistance-training
model for the quadto neuraladaptationsseemsdoubtful. Judgingfrom riceps was employed, the myofibrillar content apthe studiesincluded in this review, the hypertrophy peared to increase from the very first workout,
processactually seemsto be most rapid during the reaching significance after the second session.
first 6 weeks, after which the rate declinesslowly. Clearly, more researchinto the time course of the
Becausethe majority of studieshave only investi- hypertrophicprocessis needed,especiallywith refgateda time period of up to 12 weeks,it is difficult
erence to the effects of different resimens and
to assesshow the rate of protein accretionis affected modes.
by longer periods of training. A study by Sale et
al.tll2l suggeststhat relatively high ratesof increase
4.8 Hypertrophic Response of the
in muscleCSA (0.22-0.24Voper day) may be possiQuodriceos versusthe Elbow Flexors
ble to maintain for periods of up to 20 weeks,but
unfortunatelyno mid-point datawere availablefrom
It haslong beenrecognisedthat somemusclesare
this investigation.Therefore,it cannot be excluded very responsiveto the stimulusof strengthtraining,
o 2m7 Adis Dolo Informollon BV. All righls reserved.
SporlsMed 2m7;37 (3)
while others seem more stubborn.One explanation

to this phenomenoncould be that musclesthat are
frequently used in everyday activities are already in
a trained state,thus leaving less room for improvements in strengthand size. For example,the soleus
muscle appearsto be relatively unresponsiveto resistance exercise in comparison with the vastus
lateralis and the biceps brachii.t2@lRegardingthe
latter two muscles, it is commonly held that the
elbow flexors are in a less-trainedstate than the
quadriceps.tl28'2031
Sdies where the hypertrophic
responseof the quadricepsand the elbow flexors to
similar training regimens have been directly compared tend to support this theory.ta8'6'2slThe trends
reportedin this review for conventionalresistance
training for the quadricepsversusthe elbow flexors
lend supportto this observation,as the CSA of the
elbow flexors tended to increaseat a greater rate
(0.2OVoper day) than the quadriceps(O.ll7o per
day). Further support comes from a study by Turner
and co-workers,t2o3l
who found marked hypertrophy
in the elbow flexors (24Vo increase in CSA) in
responseto endurancetraining for the upper limb
(arm cycling to exhaustion for 30 minutes, five
timesper week for 6 weeks),while leg cycling at the
same relative intensity and duration had negligible
effectson the mass of the lower limb. Notably, the
rate of CSA hypertrophy for the elbow flexors observed in this study (0.57Voper day) surpassesall
strength-trainingstudies included in this review,
exceptthat of Narici and Kayser.tl2sl
The differences
betweenvariousmusclegroupsin the physiological
responseto similar training regimenswarrantssome
caution in generalisingfindings from one muscle
group to another.Future investigationsshouldstudy
whether the dose-responserelationships differ betweenthe elbow flexors and the quadricepsin regard
to the major training variables.
4.9 The Stimulusfor Muscle Hypertrophy in
Strength Troining
It is beyondthe scopeof this article to discussin
any detail the pathways or networks of intracellular
signalsleadingto hypertrophyas a resultofa period
ofincreasedloading ofthe muscle(s)involved.Sev-
@ 2007 Adis Dolo lnformotion BV. All rights reserued.
253
eral excellentlevievv5[205-2081
and original investigadonstel'20e-2131
on various aspectsof this topic are
available.However, a brief discussionof some of
the physiologicalstimuli occurringduring resistance
training that may trigger the hypertrophicpathways
is relevant.Over 30 years ago, it was suggestedby
Goldbergand co-workersl2l4l
that increasedtension
development(eitherpassiveor active) is the critical
event in initiating compensatorygrowth. MacDougalltzr5lnotedthat the loading of the musclemust be
very high in order to result in hypertrophy,but that
the total durationduring which the muscledevelops
tension also affects the magnitude of the hypertrophy response.He supportedthis observationwith
the resultsof the study of Sale and colleagues,tl2sl
who showeda tendencyfor training with 6 setsof
lO-l2RM to result in larger increasesin elbow
flexor muscle CSA than trainins with 6 sets of
l-3RM (33Vovs247o).
Two studies by Martineau and Gardinert2l6'2171
have provided insight into how different levels of
force and different durations of tension may affect
hypertrophicsignalingin skeletalmuscle.Using rat
muscle preparations, these authors noted that
mechanicallysensitivepathwaysreactedin a dosedependentmannerto the level of force, so that larger
increasesin intracellular signaling were seen after
eccentricactionswhen comparedwith isometricand
concentricactions.t2l6l
In a follow-up study,l2rTl
they
showedthat the samepathwayswere also sensitive
to the time-tension integral in a dose-dependent
manner.Interestingly,this was the caseregardlessof
whetherthe total durationwas distributedinto a few
long durationsof stretchor many short ones.Also,
the rate of stretch had no effect on these pathways.
In the latter study,t2l7l
they remarkedthat both peak
tensionand time-tensionintegral must be included
in the modelling of the mechanical stimulus responseof skeletal muscle. Some of the pathways
that are now recognised as crucial for the hypertrophic responsewere not assessed
in the studiesof
t6'2t'1
1
Martineau and Gardiner,l2
and little is currently
known about the responseof thesepathwaysto the
variablesof peak tensionand time-tensionintegral.
One of these is the phosphatidylinositol-3kinase/
SporfsMed 2@7:37 (3)
Wernbom
et al
protein kinase B/mammalian target of rapamycin

which has several downstream
Pathway,t20s-2081
targets,among them the signalingmoleculep70 56
kinase (p70S6K). A recent study by Eliasson et
al.t2r8lshowedthat the phosphorylationof p70S6K
in the human quadriceps 2 hours after resistance
exercise was greater after a total of 24 maximaleccentricactions compared with 24 maximal-concentric actions and to 24 submaximal eccentric actions. However,this was in the absenceof nutritional supply. In contrast, Cuthbertson and colleaguestzlel demonstrated similar increases in
p70S6K and muscleprotein synthesisafter eccentric
and concentric exercise. Importantly, in this study
the subjectsreceived protein and carbohydratesupplementation immediately post-exercise.Another
difference is that a much greater volume of work
was performed compared with the study of Eliasson
e t a l . l 2 l8 l
Based on the data reviewed in this paper, we

speculate that hypertrophic signalling in human
skeletalmuscleis very sensitiveto the magnitudeof
tension developedin the muscle. Hence, for very
short durationsof work, the increasein musclesize
will be greaterfor maximal-eccentricexercisethan
for maximal-concentric exercise of similar durations, as in the studiesof Farthingand Chilibecktr5rl
and Hawkins et al.tl48lThe responseis presumably
also dependenton the total duration of work and
increasesinitially with greaterdurations.Thus, both
short durations of maximal eccentricexerciseand
somewhatlonger durationsof concentric,isometric
and conventionaldynamic resistanceexercisecan
result in impressive increasesin muscle volume.
However, especially with maximal eccentricexercise, damagealso seemsto come into play as the
duration of work increases even further and the
acute and/or cumulative damage may eventually
overpowerthe hypertrophicprocess.This could be
an explanation for the modest hypertrophy reported
in severalisokinetic training studieswhere the eccentriccomponenthas beenmaximal and of moderate-to-longtotal durations.As discussedby Rennie
and colleaguest2osl
and Jonesand Folland,t22ol
other
physiologicaleventsassociatedwith muscleactiva-
o 2007 Adis Doio Informoiion 8V. All rights reserued,
tion (e.g. temporarily increasedCa2+ levels in the

cytosol, metabolite accumulation, ischaemia and
acutehormonalchanges)may also act as signalsfor
adaptation and interactions between these and
mechanically-induced
signalingseemplausible.The
potentialrole of acutehormonalresponseshas been
reviewedby Kraemer and Ratamess.t22ll
Apart from mechanical forces, growth factors
and hormones,anotherexercise-related
stimuli that
has beenshown to affect the hypertrophicsignaling
in skeletalmuscle (as assessedby phosphorylation
of p70S6K) is heat stress.t222l
Interestingly,both in
cultured muscle cells and in skeletal muscle, heat
stressand mechanicalstretch has been shown to
interactso that protein expressionand concentration
is higherafter a combinationof the two stimuli than
either alone.tz23'2241
These authors suggestedthat a
stress-inducedheat-shockresponsemay modulate
the exercise-inducedadaptationsof skeletal muscles, for example when combining vascularocclusion with resistanceexercise.If an interactionbetween heat stress and mechanical stimuli occurs
during strengthtrainingwith restrictedblood flow, it
could, at leastin part, explain the successof low-tomoderateintensity training during theseconditions
in inducing hypertrophy,[83'84'r3rl
even in highlytrainedathletes.tT6'2251
4.10Suggestionsfor FutureReseorch
The trendsobservedin this review could serveas
a startingpoint for experimentsaiming to establish
efficient modelsof training for the purposeof gaining and/or preserving muscle mass. Major challengesfor future researchare to isolate the impact of
each of the resistancetraining variablesand to investigatethe interactionsbetweenthem, as well as
the effects of various training strategies(e.g. perrodization, tapering,changesin type and mode of
exercisein order to 'shock' the muscles).We also
recommend that future investigationsshould describe the exerciseprotocols in greaterdetail than
hasgenerallybeenthe caseup until recently.Consequently,variablesEuchas speed,rangeand duration
of each repetition and rest periods between repetitions and setsshould also be reportedin additionto
SportsMed 2m7; 37 (3)
255
the commonly recognisedvariables of frequency,

intensity,volume and mode of exercise.
To date, the vast majority of the researchconcerning the effects of manipulating training variableshave beencarriedout using DER training.For
example,we were unable to locate any accommodating training study that directly compared the effects of different volumes on the hypertrophic response.Becausethe accommodatingtrainingmodes
can induce hypertrophy at rates comparableto those
of conventional weight training, and becausethe
training parameters are easily standardised,these
modes are well suited for experimentsdesignedto
provide insight into the natureof the dose-response
relationships.
In humans, electromyostimulation(EMS) has
beenproven to result in increasesin musclevolume
comparable to those seen after voluntary strength
training. Since EMS bypassesthe CNS, the level of
activation of a particular muscle group can be
standardised.
Combining EMS with isokineticdynamometry could provide an opportunity to gain further insight into many of the issuesdiscussedin this
review, such as the effects of the level of torque, the
type of muscle action (concentric vs isometric vs
eccentric)and the total duration of activity. Howev-
er, becauseEMS involvesmotor-unitfiring patterns,

which are usually very different from those occurring in voluntaryexercise,findings from such studies may not necessarilyapply to voluntary strength
training.
As noted in section 2.1, investigationsconcerning very well-trainedindividualsarelargelylacking,
as are studiesextendingfor longer than the typical
8-12 weeks.Becauseof this, the knowledgeregarding the dynamics of the hypertrophy processpast
this point is limited. In short-term studies, large
increasesin the training loads for lower-bodyexercisessuchas squats,leg pressesand kneeextensions
are often reported, sometimes on the order of
100-200%o.18'55,re61
Since the gain in quadriceps
muscle volume during the same time period rarely
exceeds=l1%o, the stressper unit of muscle area
shouldincreaseby almostas much as the increasein
training weight. The significanceof the large increasein the stresson the musclesand its interactions with the volume and frequencyof training in
terms of the hypertrophic responseand the risk for
overtraining remains to be explored. Although not
discussedin this article, the issueof dose-response
effects needsto be adressedin the trainine of other
Table l. Recommendationsfor dynamic extemal resistancetraining (e.9. weight-basedresistance)for hypertrophy

Muscleaction
Con and ecc
Con and cc
Eccentric(ecc)overloadtraining
Ecc (con = optional)
Exercise
Single and/ormultiplejoint
Singleand/ormultiplejoint
8-1oRM (range:-12)
=75-80% of 1RM
Ecc = >105%of l R M
Con = 60-75%of lRM
Moderateload slow-speedtrainingConventionalhypertrophytraining
Load
Repetitions
Sets
-50%
of 1RM
failureor near 4-
8-10 to muscular
1-3 Derexercise
1-5 per exercise
from1-2 to $-6 sets Progression
kom 1-2 to 3-5 setsin
Progression
from1 to 3-4 setsin Progression
8-14 to muscularfailure
1-3 per exercise
total per musclegroup
in total per muscle group
Velocityand duration
per repetition
Slow
Ecc = 2-3 seconds
Con = 2-3 seconds
Moderate
Ecc = 1-2 seconds
Con = 1-2 seconds
Slow/moderate
Ecc = 24 seconds
Con = 1-2 seconds
Resl between sets
3H0
60-180 seconds
120-180 seconds
Frequency
2-3 sessionsper musclegroup/ 2-3 sessionsper musclegroup/

week
week
Theserecommendations
Suitabletrainingmethodlor
are for
beginnersand individualswho
noviceto moderately
trained
individuals.
Welltrainedathletes
cannot tolerate high forces
mayneedincreased
variationin
intensity
andvolume
Comments
seconds
1-3 sessionsper musclegroup/week

Mainly for advanced to elite athletes.
Progressivebut carefulincreaseof
the load and volumefor the eccentric
ohase
Con = concentric: RM = repetition maximum.
o 2007 Adis Doto Informction BV. All righls reserved.
SportsMed 2Q7;37 (3)
Wernbomet al
Table ll. Recommendationsfor accommodatingtraining lor hypertrophy

Moderatelyfast concentric (con)
training
Slow concentrictraining
Accommodatingeccentric (ecc)
overload
Mode
lsokineticor hydraulic
lsokineticor hydraulic
lsokineticor isoinertialtlywheel
Muscleactions
Con
Con
Exercise
Ecc and con (con is optionalin

isokinetics)
Ettorta
90-100%
90%
Ecc = up to 100%
Con = up to 100%
Repetitions
10-15
10
6-8
Sets
H per exercise
Progressionfrom 3 to rre sets in
tolal per musclegroup
3-5 per exercise

Progressionfrom 3 to 5 sets in
1-5 per exercise

Progressionfrom 1-2 to 4-5 sets
in total per musclegroup
Velocity
12o-24o"ls
4ffio"/s
4H0'/s
5 seconds,
'120seconds
0-5 seconds,
120 seconds
Rest between repetitions 1-2 seconds,

and sets, respectively
60-120 seconds
3-5 sessions per muscle group/

3 sessionsper musclegroup/week 2 sessionsper musclegroup/week
weeK
indicates level ol torque in relation to the maximum possible torque at the specifiedvelocity.
Frequency
a
populations,such as the elderly and individualsrecovering from sportsinjuries.
to hypertrophy further in relative terms than a previously untrainedmuscle. Conversely,a muscle that
has atrophiedbecauseof disuseor detraininghas a
4..|i Limitotions
large growth potential and merely getting it back to
its previouslevel will representan increasein musWe recognisethat it is obviously very difficult to
volume if the atrophied state is taken as a basecle
separatethe impact of each training variable from
line.
Thus, even slight variationsin training status
the effects of the other training variables.For exammay
affect the hypertrophic response to a given
ple, if one increasesthe training load (percentageof
resistance
training regimen. Also, the method of
IRM) in conventionalweight training, it will also
measuring
muscle volume or CSA may influence
affect the volume of training, unlessthis is compenresults.
With earlier scanning techniques,the
the
sated for by increasing the number of sets peranatomical
CSA
of the muscle was measuredwithformed. Furthermore,we acknowledgethat the main
out
for
intramuscularfat. Recentmethods
correcting
objective of many of the studies included in this
of
MRI
and
allow
for measurements
CT
of interstireview was not necessarilyto maximise the hyperwell
tial
fat,
as
as
muscle,
and
for
consequently
trophic responseand that the motivational level of
calculation
of
adipose
tissue-free
muscle.l227l
In
the subjects may well have differed considerably
young
healthy
subjects,
anatomical
muscle
the
area
betweendifferent studies.Closely associatedwith
motivation is whether the training is performed is only slightly larger than adiposetissue-freemusHence, any increasein musclevolume
under supervisionor not. Direct supervisionof the cle area.tz21l
as
a
result
of strengthtraining will mainly reflectan
workout has been shown to result in superior inincrease
in
adiposetissue-freemusclemass.Therecreases in strength when compared with unsupervisedtraining.tzz6lThe level of supervision fore, the data from the studiesincluded in this review were pooled irrespectiveof the muscle-scanduring training varied amongthe studiesincludedin
ning method used. However, becauseof the factors
this review.
Apart from the training regimen, the training discussedhere and the many other confounding
statusis also likely to have an impact on the hyper- factors that inevitably are present when summaristrophic response.Theoretically, a muscle that is ing and comparing the results of many different
already somewhat hypertrophied has less potential studies,the dose-responsetrends and recommenda-
@ 2007 Adis Dsto Infomotion BV. All rights reserued.
Med 2m7;37(3)
Sports
257
tions outlined in the presentreview should be regardedas tentative.
tion standt3land by Kraemer and Ratamess.trlWe

also agree with these authors on the importance of
progressionand individualisation of the exercise
prescription.Regardingprogression,we recommend
4.12Troininglmplicotions
ond Recommendotions
low volumes (e.9. l-2 sets) in the initial stagesof
training,when performingeccentric-muscle
actions,
Preliminary recommendationsfor each mode are becauselow volumes have been shown to be suffigiven in table I, table II and table III. These are cient to induce hypertrophy in the early stagesof
basedon the evidenceoutlinedin this article,as well
training and becauseexercise adherencemay be
as on training protocols that have been shown to
improved if the workout is relatively brief. Also,
increasemuscle mass. However, the tables should avoiding unnecessarydamagemay allow hypertronot be interpreted as stating that all modes and phy to take placeearlier.As the individual adaptsto
methodsare equally effective in increasingmuscle
the stimulusof strengthtraining,the overall volume
mass. Rather, the aim is to summarise different
and/orintensitymay have to be graduallyincreased
methodsthat may be suitablein different situations
to result in continuedphysiologicaladaptationsand
and for specificpopulations.For example,fatiguing,
otherstrategies(e.g,periodisation)can alsobe introlow-to-moderateload, slow-speedresistancetrainducedif even further progressis desired.
ingtTs'88'e2'llo;
haspotentialapplicationsfor the rehaIn this context, it is essentialthat the trainer or
bilitation of patients for whom the high forces of
exercisearecontrain- therapistis aware of possible interactionsbetween
conventionalheavy-resistance
dicated.For patientswho can toleraterelativelyhigh the trainingvariablesand how these,in turn, interact
forces, but for whom the metabolic and cardiovascu- with the exercisetolerance of the individual. For
lar demands of traditional strength training are too example,a training volume that is appropriateat a
severe,pure eccentricexercisemay be an alternative frequency of two sessionsper muscle group per
week may becomeexcessiveat three sessionsper
becauseof the low energy demandsof this type of
for conventional week. Conversely, a volume that is sufficient at
exercise.tls3l
Our recommendations
hypertrophy training are similar to those presented threesessionsper week may be lessthan optimal at
in the American College of Sports Medicine posi- two sessionsper week. The workout structure(e.g.
Table lll. Becommendationsfor isometric training lor hypertrophy
Low-intensityisometrictraining
High-intensity
isometrictraining
Exerciseselection
Singleand/ormultipleioint
Single and/or multiplejoint
Maximum-intensity
isometric
training
Torquelevel
30-50% of MVIA
7H0o/o ol MVIA
100o/o
of MVIA
10
1-3 per exercise
Progressionkom 1 to 3 sets in
Repetitions
Sets
2- per exercise
Progressionfrom 2 to 4- sets in
2-6 per exercise

Progressionfrom 2 to 4-6 sets
in total per musclegroup
Durationper repetition
4H0 seconds,and to muscular

failureduringthe linal 1-2 sets
15-20 seconds,and lo muscular 3-5 seconds

failureduringthe tinal 1-2 sets
Rest between repetitions

and sets, repectively
3m0
3H0
Frequency
3-4 sessions per muscle group/

weeK
Comments
Suitablefor individualswho cannot Suitablelor individualswho

tolerate high forces and with
cannoltoleratenear-maximal
restricted range of movementdue forces
to pain and/or injury
seconds
seconds
H sessionsper musclegroup/
week
25-30 seconds,
60 seconds
3 sessionsper musclegroup/week
Care shouldbe taken to avoid
excessivebreath-holding
and very
high blood pressures
MVIA= maximal
voluntary
isometric
action.
o 2007 Adis Doto Informollon 8V. All rights reserved.
Sports
Med 2m7;37(3)
258
whole-body workout versus single muscle-group

training) also has a direct bearing on the appropriate
dosageof training. Table I, table II and table II are
intended as guidelines for single muscle-group
training. If whole-body workouts are performed, the
volume of specificwork per musclegroupmay have
to be reduced so that the overall volume does not
become excessive.For further discussionon the
topic of workout design, we refer to the paper of
Kraemer and Ratamess.tll
5. Conclusions
This review demonstratesthat several modes of
training and all three types of muscle actions can
induce hypertrophyat impressiveratesand that, at
present,there is insufficientevidencefor the superiority of any mode and/or type of muscle action over
other modes and types of training. That said, it
appearsthat exercisewith a maximal-eccentriccomponent can induce increasesin muscle mass with
shorter durations of work than other modes. Some
evidence suggeststhat the training frequency has a
largeimpacton the rate of gain in musclevolumefor
shorter periods of training. Becauselonger studies
using relatively high frequenciesare lacking,it cannot be excludedthat stagnationor even overtraining
would occur in the long term. Regardingintensity,
moderatelyheavy loads seem to elicit the greatest
gains for most categories of training, although examples of very high rates were noted at both very
low and very high intensitieswhen the sets were
performed with maximum effort or taken to muscular failure. Thus, achieving recruitment of the greatest number of muscle fibres possibleand exposing
them to the exercisestimulusmay be as importantas
the training load per se. For the total volume or
duration of activity, the results suggesta dose-responsecurve characterisedby an increasein the rate
of growth in the initial part of the curve, which is
followed by the region of peak rate of increase,
which in turn is followed by a plateau or even a
decline.
It is recognisedthat the conclusionsdrawn in this
paper mainly concern relatively short-term training
in previously untrainedsubjectsand that in highly-
@ 2007 Adis Doto Infotmotion BV. All righls reserved.
Wernbom et al.
trained subjectsor for training studiesextending for

several months, the dose-responsetrends and the
hypertrophic effects of different modesand types of
strength training may be very different. The same
may well be true for other populations,such as
elderly and injured individuals.
Acknowledgements
No sourcesof funding were usedto assistin the preparation of this review. The authorshave no conflicts of interest
that are directly relevantto the contentof this review.
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Correspondenceand offprints: MathinsWernbom,Lundberg

Laboratory for Human Muscle Function and Movement
Analysis, Department of Orthopaedics, SahlgrenskaUniversity Hospital, Grna Strket 12, Gteborg, SE4f3 45,
Sweden.
E-mail: mathias.wembem@orthop.
gu.se
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The Influence of Frequency, Intensity, Volume and Mode of Strength Trainirg On Whole Muscle Cross-SectionaAl Rea in Humans

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Sports Med 2m7; 37 (3)i 22$264

BV. Afl dghts reseNed

The Influence of Frequency,Intensity,

gV. All rights reserysd.

The Influence of Frequency,Intensity,

Strength Training and Muscle Cross-SectionalArea

Strengthtraining has becomeincreasinglypopular in recent decades.Whereaspreviously strength

training periodsor training sessions.aimed

@ 2m7 Adis Dqto Intormotion BV. All rights reserued.

Sports Med 2@7; 37 (3)

the quadriceps was by far the most studied muscle

1.2Clossificotionof Muscle Actions,Troining

For the purposesof this review, the terminology

O 2007 Adis Dolo Informotion BV. All rlghts reseNed.

SportsMed 2@7;37 (3)

Strength Training and Muscle Cross-SectionalArea

idea with variable resistanceis to closely match the

@ 2007 Adis Doto Infomotion BV. All rights reserued.

type of training, but Murphy and Wilsont2elhave

increasesonly marginally with increasingvelocity

sessionsper muscle group per week, as opposedto

In the majority of the studies,the subjectswere

@ 2007 Adis Doto Informotion BV, All rights reserved.

Sports Med 2007j 37 (3)

Strength Training and Muscle Cross-SectionalArea

the pre-training values. To allow for comparisons

o 2007 Adis Dolo Informotion BV. All righls reseNed.

contributingmechanismfor whole musclehypertro-

2.1 Quodriceps Studies:Dynomic

otherwiseindicated,the analysisfor DER dealswith

sions per week, the increasewas 0.ll%o per day

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Strength Training and Muscle Cross-SectionalArea

2.2.1Lenglhof frolnlng Peilod,Avercge lncrcose

per day). No studieswere found which involved <3

o 2m7 AdisDoto Informcrtion

When the percentage values for eccentric torque

@ 2m7 Adis Dolo Informotion BV. All righls reseNed.

Flg. 5. Torque vs percentage incraase in cross-sectional area

Strength Training and Muscle Cross-SectionalArea

tween 25 and 50 units (0.1470increasein CSA per

100 150 200 250 300 350 400 450 500

2.3 Quodriceps Studies:lsometric Resistonce

o 2007 Adls Dolo Informolion BV. All rights reserved.

40 60 80 100 120 140 160 180 200

SDortsMed 2m7;37 (3)

Strength Training and Muscle Cross-SectionalArea

timulation to evoke muscle actions.In the first of

(0.16%oper day). In the secondstudy,t1221

o 2m7 AdisDoto InformotionBV.All rightsreserued.

SportsMed 2m7j 37 (3)

jects in sections 3.1.1-3.1.4. Four groups (four

@ 2007 Adis Doto Informolion BV. All righls reserved.

Strength Training and Muscle Cross-SectionalArea

3,3 Elbow FlexorStudies:lsometric Resistonce

The literature search resulted in three original

3.2 Elbow FlexorStudies:

@ 2m7 Adis Doto Infomolion BV. All rights reserued.

Flg. 11, Total number of rpetitions vs percentage increase in

SportsMed 2m7;37 (3)

frequenciesin more conventionaltraining. Furthermore, it is interesting to note that there was no