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Anabolic Processes in Human Skeletal Muscle:
Restoring the Identities of Growth. Hormone
and Testosterone
Daniel W, D. West; Stuart M. Phillips, PhU, FACSM
Abstract; TestosteroneEUppIementation acts via numerous mechanisms asa highly potent anabolic agent to skeletal muscle. Although
growth hormone {GH} strongly affects collagen synthesis and lip olysisr as weli asjricreasjng lean body mass, it ts not an abollc toward
the contractile fa, myofjbrlJIar) muscle tissue in healthy Individuals. However, there Is a persistent belief {both in scientific literature
and among recreational vyeightltfters) that exercise-induced release of 6H er»d tsstcsiercnE yndarpins rnuBCHlsr hypErtrophy with
resistance training. This Is a premature assumption because although pharmacological GH supplementation can Increase muscle
strength or size In individuals with clinical GH deficiency, there is no evidence that transient exercise-induced changes In GH have
the~sarne effects in Individual* with normal GH letfsferExerase paradigms are designed based on the assumption {not necessarily
evideneed-basad mechanisms) thatGH and testosterone facilitate anabolic processes that lead to skeletal muscle protein accretion and
hypertrophy* Our recetit work disputes this assumption. Instead, our data indicate that exercise-induced hormonal elevations do not
enhanceJntnacellular markers of anabolicsignajfng orthe acute postexercise elevation of myoflbriliar protein synthesis. Furthermore,
data fromour training study demonstrate that exercise-induced Increases in GH and testosterone availability are not necessary forand
do not enhance strength and hypertrophy adaptations, instead, our data lead us to conclude that local mechanisms that are Intrinsic
to the skeletal muscle tissue performing the resistive contractions {ie, welghtlffting) are predominant in stimulating anabotism. The
purpose of this article fs 1) to provide a brief overview of the mechantsmsof action of testosterone and GH; 2} to discuss the inability
of physiological exercise-induced elevations in these hormones to have s measurable impact on skeletal muscle anabolism; and 3) to
describe factors thatwe believe are more Impartantfor stimulating hypertrophy In human skeletal nrascle. Clarifying both the role of
hormones in regulating muscle mass as well as the underlying basis for adaptation of skeletal muscle to resistance exercise, will hopefully enhance and supporttbe prescription of resistance exercise as an integral component of a healthy lifestyle.
Keywords testosterone; growth hormone; hypertrophy; muscle protein synthesis
__________
DanletW. D,West!
Introduction
Stuart M. PhBllpy, PhD, FACSM1 fop^t phases of positive net protein balance, which can be generated by the elevation of protein
letabollsm Rweirch
synthesis in response to repeated bouts of resistance exercise and protein ingestion, underpin muscle
Ktnesloiow McMastfirtjntverstty,
hypertrophy. In addition to the stimulus provided by loaded muscle contraction and amino acids, the
Hamilton, Ontario, Canada
hormone testosterone is considered, highly anabolic. Training response in young men is attenuated
when testosterone is pharmacologically suppressed to approximately oner tenth of, normal levels,'
Thus, testosterone contributes in some way to the anabolic processes that are initiated by resistance
training in young men. Interestingly, vronwn hare testosterone levels that are approximately 10-fold
lower basally (not unlike the levels seen in the study by Kvornteg et all), yet a recent large study rep ores
comparable rates of hypertrophy to men and women after resistance training,2 Does this .mean, then,
Correspondence:
Stuart M.PWffips, PhD. FACSM.
Department of Ktneslolo^y,
McMajter University,
Hamilton Ontario
i8S4Ki,carach.
^?, ....
[email protected]
that testosterone is contributing to anabolic pathways in men but not in women? Or are women more
_ - . . . . . . ,
., ,
, , r t - x 4.
7^.1
^T
,
•,
efficient in using their inherently lower levels of systemic testosterone? Fnndamental questions such as
these remain unanswered as the role of hormones in regulating muscle mass continues to be elucidated.
Studies using pharmacological doses of testosterone and growth hormone (GH) help clarify these
hormones* mechanisms of action onskeletalmuscleand. other bodilytissu.es. For example, it isapparent
fhatsupraphysiological testosterone is highlypotent in stimulatingnrascle hypertrophy, whereas GH does
notincreasemyofibrillar protein synthesis andinstead directs anabolic action toward increasing Whole-
©THE PHYSICIAN AND SPOKTSMEDICINE* ISSN -0091-3847,October 2010, No.3. Volume 38
041509^7
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CLINICAL FEATURES
body protein synthesis (eg. connective tissue). In contrast,
:hoj^o^es./,^«hi<:h haye;beeincharacterized
' ' ' ^
challenge the common perception that systemic changes
in anabolic hormones are required for the stimulation of
muscular hypertrophy that occurs with resistance training.
tw^s-^ ••&.?«.-• vrr.-r.
•ovys 'vsvcy ••••;•} -r.-r .z-.:
payjcaa c>ccu?£)ithe absence of exercise-induced increases in
Testosterone
Evidence is undisputed that anabolic steroids are extremely
testo'steron^ do notenhi&Keajid'are nStiieoessarytorriHiscle poteatia inducingjmuscle hypertrophy. Indeed, the capability
anabolisro, Moreover-women can fwyehypertropby to a simi- of testosterone to increase muscle cross-sectional area, both
lar degree to men in response to resistance training, despite solely and in combination with resistance exercise, has been
haying testosterone levels that are approximately 10-fold demonstrated 5n a study of testosterone supplementation
lower bastxlly and with no exercise-induced testosterone in young men.* Thn highly anabolic nature of testosterone
xSspjpnse,,
^
^ ontajbKtft to ijicreping evidence is likely taterdfipdndent to some extent (Figure 1). Firstly;
s&pp^brr' ib.c%irirfegfilatibri?of ske.letSl muscle and the nature of testosterone as a steroid allows it to cross the
A5Figuri ), Th« multiple mKbantoii by -which twwaeerooe cm ace ta i nwofibtr.
' ?•£•'•"•
.EKerase-inducerf-eievatfons
in systerhjcitestQster'o.ne
'-' l'-V"•-•£: :%'.!'
Local and rogen
metabolism
Afldrogen
receptor
•^ Intraceliyjar
amino'actd
reutHizatJon
Non-g'enornic
-signalling?
t.
r
^Transcrlptidn
t MPS
t Satellite cell
proliferation
INDIVIDUAL USE ONLY, NOTT5 BEDISTWBUTED OR PRINTED FOR. DISTRIBUTION, © JTE MULTIMEDIA, LLC
CLINICAL FEATURES
Human Skeletal Muscle Hypertrophy
« :. \f.y
Hpid bilayer bitthe.sarcolemma whereat can form a complex binding to improve festing net protein balance.5'18 Thirdly,
with cytoso{ip.or:nude.^^d^0gen;,^ceptorso' to Toltimately testosterone may facilitate the accumulation of DNA that is
enhance gei{^tr^scripfio^Tlife!an3rbgenij:ece^ior contains normally re<juired-for muscle growth by activating satellite
> Simportaatsubumts; 1) a C-tenninaWoraam, which binds cells." Bgure 3 places exogenous testosterone at the bottom
iicands fee, fetosleiohfeV, 25 a central domain* which binds ofthehvpertroohvtwramid^indicatirifftlijaitisViiirWvTine/.nt
<~i:-?':>
•*•*:•*r;-: • :.-
•:'Z-j}Xs'
"rr
V."';i.
"A>,iV7T-
I7-'',-
dimeiize, translocate to 3ie nucleus, and bind to androgen-
'.Vi-y
. :V
V'.*"!
GrO'Wtn H
directly through its receptor
IGF-1 to regulate a variety
targets direcfiy'relat^f to protein \cctetion (eg^MRQfbr1'- of tifeues (Figure 2), It is weE established that growth and
addition, testosteronejrnayaLto ^-nlinnr^ trantrrmtmti of other maturation of the musculoskeletal system is mediated by
anabolic systems, such as the local production ofinsulin-like systemic changes in the GH/ IGF- i axis.u For example, a
growth, factor 1 (Jut-IJ^Secondly, phannacological doses of deficiency or excess of GH during growth can result in overt
testosterone increase the rate of muscle pjrotein synthesis.7'8 musculoskeletal changes (eg, slowed or accelerated changes
mcr.ease^
Jnstatare,respecdvdy).Rirthennore)GHplaysaBi important
lizatkm of a^no4dd4-td%feriMatentocl^prbfe3K'syJith%siss role in regulating body composition madultlife; rh'GH sup-
if
ir« 2, A taSc'orervtaY of tte afettt oTgrowtfi hormone (GH) onrouWpteBssues lo tNa bod/v
••:&
.ftSl!
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CLINICAL FEATURES
Daniel VY O, Ytetand Saari M, Philips
lead to an inaccurate interpretation that there is a cause-and- 7 days of GH administratioa stimulated collagen-producing
effecfcrefetkpsjii^. B.ased on.Ms logk^'.GHrcculd be viewed osteobksts and activated bone remodelling,^ Circulating GH
'
andIGE-1 are thought to also increase collagen synthesis by
mass..jn. elderly populations. Although GH replacement cart fibrobiasts andincreasedfibxoblast cell division in the extracelije'usedtotecover lean body mass in individuals with climca! lular matrix and tendons.2* ';;:a-:Xs:^^T^fe-^Safebi ?M&iS<3'Sj-i:i jVaJJSjj. J=;^ a s&* •S^>« • :'.*%-.-., -•-•4-.,;.-.v-;-1
'£>'
• . f ' o T o y y n g a o .
ISu^lemen^tiqn;^
•
''effect bft:Tnasd«lhy^e;r|roplay;'%theiv accjo^yjating evidence
-»•:
fib» hypertrophy fbllovnng resistance training to elderly men. suggests "thatGHnTOcdanspjhiiarily to mcreaseihe synthesis of
These findings are fax agreement with studies by Iferasheski whole-body prqteins and connective tissue, in particular
increasing water .retention and altering body composie^^i^jvho^fouad.&af^^
was'increased, tfcfere'was'jrwj augmentation of •muscle protein. tioru Accordingly, GH supplementation does not increase
synthesis with thGH sxtpplementation in. both young and old layofibrillar protein synthesis or muscle strength, in healthy
men. Moreover, in sedentary elderly men, 16 weeks of GH individuals.Thus, Figure 3phces GHat the top ofthe pyxamid
and resistance exercise conferred no added strength gains effectors Kgtilatinghypertrophy.indicatingthatitscontr/ibuacross 9 exercisesversus resistance exercise plus placebo.KNot tion. is negligible,
Exercise-Induced Anabolic Hormones
raass were greatefin the GH group, trteauthors demonstrated
tSat?a''disjfopordonate ammuit ofthe mass was because of
increased total body water,1* whach may be partly due to a
,-ti!.^.-M>f&C«S,^'f$'<:&^f^r"^f-p-.
%•'• «*•;& /jiVV:'-; ..
., .
j:e.a CQnnective tissue that
becaus'fe SH acutely arid chronically alters the regulation of
¥Jfcs&{S.'%S!=
..'.f.
c^Siposition by stimulating: Itpolys3sw and increasing lean
significantlyincreased lean body mass byapproximat«ly2!
However, the increased mass did not translate into strength
changes, implying that GH can promote fluid retention, but
not affect functional (ie, strength.-promoting) muscle protein
accretion. Indeed, GH-induced increases in lean body mass
accretion of l«an tissue that arises from elevated whole- body
.,> ..-.-, .- , _
____,——,_
~
in cases :61 clinical deficiency can Help recover muscle mass
f; •.-.:<•:.-•;' •*• :•-#.:./:. -H'^Tf .-""• ••'• •''.-':• "•••• "•>
"
'
'
i
anabolic
effects:are
directed toward
the synthesis
of collagen,
_ _^^
, ,v!
^
'
~
notjcpntracble myofibrUlar proteins,-" tor example, Wallace
J
?v
Resistance exercise progranfvariables can be .manipulated to
alter the acute postexereise blood concentration profile of testosterone and GH,Specifically, boutsof resistance exercise that
arehighin volume,actiyatealargemusclemass,and requires
high degree of effort transiently (-40 min) to elevate the circtiIatinglevels.of;te||osteronearldG.p.Werecendy measured the
'
.
t
o a n acute
-'j,
it hormonal
l .... ?*v" s ' ' f •ib r -:..v.« --; ' ' / ' v ' , ' ' " "
••^epvirorunents:25 In o ne .cortpttjcm, arrn ejce rcise'aloh
formed; -ftis resulted in GH and testosterone concentrations
that remainednear basal lerds. Intheother condition, the performance of identical arm exercise ^-as immediately followed
byahighvolome of leg exercise ata modente-to-heavy load,
which activatedalargerauscleniass. Arm exercise was selected
to precedeleg exercise so that there wouldbeno possible confounding influence of central fotigue (ie, participants would
be able to exert maximal effort during arm. exercise in both
conditions without any residual fatigue from the leg exercises),
Ihe leg exercise stimulated a large increase in endogenous
GH and testosterone, which peaked approximately 15 minutes after «xerci$e, These hormones returned to basal levels
by 60 miautes after exercise. As expected, the rate of muscle
protein synthesis was elevated after exercise (ie, a main effect
was observed); however, despite drastic differences in the
postexerdseconcentrationsoftestosterone and GH, there were
-no.cUfferenws between-condltions. ^erefbre, acute increases
".:" ' ^.'* ••"";' ^ •' •"'• '* .^'*'' ''*^- - --"- ':'" ' ;• v ••"'•''' I'- • '•* ' '•'• •'
•'itiitliie'a^ulaMi^-oEGH and testosterone d3d.npt'ejihance the
V rate 6frnuscle.pr<>teinsyntJiesisJp tiie biceps muscle exercised.
1
:i:
100
exercise»irnieie.nn"dings'are in agreementavithtKe report that
1
'©THE.PHYSICIAN^ND SPORTSMEDICtNE . ISSN"00?l-3S47, October2010,No. 3, Volume 38
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CLINICAL FEATURES
Human aretetal Muscte Hypertrophy
Figure 3,ftom ej««itt>«x!rift5oucf«torath« w&^oce
which is not associated with hypertrophy. This Is in contrast to
the specific nature of various muscular adaptations that occur
in response to exercise, as demonstrated by Wilkinson et aU50
Perhaps it makes more sense that G0 would exertmore general
metaboliceffects such as'iskb'stratefnobiiization and connective
Mfntanat
"FactorsThat Drive'Hypertr'oph^'
In general, there are numerous inputs contributing to the
£ vv.wgiia!ing milieu that takes place in each muscle cell that is
'"' •'•'• stimulated frorn loaded contractions. We know that these
loaded contractions activate signaling cascades that increase
mRNA translation, which is thought to be the rate-limiting
step to increasing muscle protein synthesis.31'33 However, the
k is iwtewood that tha« ftcwrs oton Yfflrtt In tonwrt and not in IsoMon:
&rtJNmK«ifl,tfwreUth-apoiltiom of cerah c«npofl«nu are. somfiwiat specuNxty?H<m«/ef,cofKttJuSi"?ttcartft wtS kJ^j^Twwcomgopaits^i v.tiflwiicSjo.ps'jRvraM
JQCUS of ^^.^ for t}]ereKulation of muscle taass remains to
°
be fully elucidated because a siirular early activation of pxo-
t^involvedintransfetionalcoHtrolwasrecentlyobserved
totyp<«ro!%ww ctewrmine 4w potaitbt of i)«feal wijiio (orespondto fectoc after both, aerobic and resistance exercise,54 which are known
noced ort tft^ pvTaniy^&wccl**.— rillitwKA exeixhe,
,
T.
T
, ,.
•iL-5 V?."/
tf'-R
!-=•
We have rec^tgfepnMmStthe je.5ctt^eJSnjQngs'in S^rainiaig
study in wM^'*thwe2'diSeren?')ioroional cb'nclilSc>ri§'w^re
elicited repe^tdljgalfrlf bouts of elbow flexor exercise. In
toproducedwergentphenotypes,leading the authors to suggestgenetranscriptionmaybemoreiEiportantthaaoriginalfy
thought.
Wehaverecentlyobservedthatexerciseloadisnotaprimary
variable 5n determining .gsjg.onse to prqt;ejn synthesis. -Spe-
not'e
men witfi training^
As outUne d~prevjc
after exercise t
and/or henvoconcentrationT The biologic^ implicatioas of whlcharahighIyhypertrophic>:>?'areacti¥atedduriDgtherepetipostexercise honaoaal changes remain to be elucidated, but tions preceding muscle failure despite the relatively light load.
elevated GH and testosterone appear to respond to various Iheacclusiojithatdeveiops with, this t>ye of training UMypkys
signals associated with the exercise bout (eg, increased lactate an important role it the type H fiber recruitment and overall
concentration, neural feedback, Kypoxia24); as such, it is possible muscle hypertrophy at light loads.33139
thattheyareresjp^ridpg.asgart.of.ac^^
Although load does not appear to be a crucial factor in
fuel. Interestingly, the acute SH response after 'eXerdseappears creating an acute anabolic response, we have recently observed
to be associatedjrithan'increase in the rate of glycerol appear- that 3 sets of resistance exercise enhance both theintraceDul&r
anc^ameasweoflipclyris."Thefindingthatcarb<>hyAcatepro- signals and the rate of myofibrillar protein synthesis versus
vision blunts the elevation inLCOrtispli8f9 (an exercise-responsive 1 set.10 In other words, certain regulatory anabolic signaling
hormont involyecljn giucosgrej^seyiapjieap'-.to support fois proteins (k, p70S6Kl) appear to remain active for a longer
notion, JRurtherrhore, whik'ScerciseisaVe'U^taMishedstima- duration following 3 sets versus i set of resistance exercise at
lator of GH t|le[ase;.j[t.appears that the exercise-induced GH 70% of IEM. Specificattyi;p^OS6Kl, whose activation is corrase;a^-^elitiv«ly jK>asD£ci§c and-'is -releSsed in -response .to .Tv/related wifii'-increases jn;'myofibr^lar.:protej«-fynthe3is41.arjd-..
©THE PHYSIg!Afj :ANp\SPoJtTSMEplC!Ne\r 009JJ384?, OctoW2010, No.},Volume 38
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CLINICAL FEATURES
DanteiW tt West and StuaAM.PMIHps
Ih'us, itjippears that repeat exercise sets may allow for the
agreaterdegree)thvm so/protein oranbo^oric carbohydrate
reqtup..rnuJ(|ipIe sets to be recruited andanabolicelly active.
•'"'Tfie'^TiKerent biolotrical nut-noses of the increase in GFT
of postexercise nutrition with training-induced hypertrophy
nhenotvnes'-:ii!reH'tew'hvTan<r''iTtd Phit1in<:5<)
elevations appear tofeea metabolic'byprodttct mat ' vnth postexercise nutrition, it lias rtev«r been 'demonstratedj
tisStiS anaboTisinTinaeed^-tSe complete'iacicb'f:i relationship eniance skeletal muscle protein accretion,
between the exercise-induced hormone response and either
the acute or chronic phenotj'pical outcome has led to the
question, of v?h«re or how a. hormone such as testosterone, Adaptations to exercise are both exercise- and muscle-specific,
which is anabolic ia nature, could be occurring. Recent work suggestinglocal factors are ofprimary importance. TJie many
enzjrm.es. AizawaTet at4*115 sitggesfexercise-induced actiYation improve oar understanding of the regulation of human, skeletal
o^bca\aS3rogenrrietabolism«iidreportincMasesinandrogen moscle. What is becoming clear is that exercise-induced GH
receptor^content and intranaisculactestosterone. Ihus, as we and testosterone responses are not suitable surrogate measures
~
'^^ asvare that
ifiteimusctfer hormone concentrations (while manipulated
.-WiSil^ftrfi'
^f,;^
, ,
,
,
f
<&S;upregu!ated in concert-with the intramuscular hormone
for ligand-ieceptor interaction. Burthermore,localandrogen
production appears reasonable because intramuscular
testosterone concentrations increase with exercise in female
rats, whichhave lowsystemic levels. Although these data are
intriguing, however, human data didnot show an increase in
intrahius.cular testosterone Jevels-TOth resistance training.44
Changes i^inb^uicujar'androgen metabolism need to be
corroborated with aphenotypical outcome (eg, muscle protein
syhfliesis) and inhuman skeletal muscle to determine if there
isT^,.;i"-.
a local.hormone
component that is contributing to other
..:.;••-;.:-•(•, &•.' '> '\~. • ' • •_?:•'•-:. V- ;V* ' : ., *'*•
v
1*"^ • ^ • . _±_"i;v _ -' .^t „_..-• ,' -j!iY,j.::o t
1 _- J_ 3
aort&aod/clironic anabolic response to resistance exercise is
102
<>fsMetairauscleanabolisnxanddoaotrefiectthernechanJsTOs
that underpin hypertrophy: Adverse events associated with
theprescrip.ticppf Jestosterong,cpntinueto be reported ia the
,_
„§':«!: flKlr .
.
&}3
.
.
.
,
_
wefi as reducing risR for chronicdiseases, Future studies need
-head
comparison of the health benefits of resistance exercise versus
aerobic exercise versus pharmacology-based interventions;
perhaps then the prescription of resistance exercise would be
more prevalent, which wmld undoubtedly benefit the public
health profile.
Acknowledgments
Support for this review wasprovidedittpartfromthe Natural
Sciences and Engineering Research Council of Canada and
from The Canadian Institutes of Health Research. Daniel
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CLINICAL FEATURES
Hitman Skeletal Muscla Hypertrophy
•
••'
21 „„
References V.1' . . . . . . .
." • • '....
1. Kmm\^Tiw^s^UfZi^iKpiJ^a^i(K:$^t^o^fT^^enixis
a randomized, placebo-controlled, and blinded fotervwitlofl study. AfiiJ
,„,
u
t_.
„_,..
,,.„,
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*he collagen synthesis- to human tendon and skeletal muscle i
affecting myoflbflilar protein synthesis, JPIystol 2010;5S3(pt 2);
22, ^^acoJD,CMfleoRQUiridbergJ?A,et;aI.RespoTi$3$ofn\3rfcersof bone
ai>d collagen fcunownrto e»!rdse,.jtoirtMionnone (GH) administration,
, -vfcj^y»)» z>.utuuux,' ituig j^j.-v iiJ..xicvduuaa Ait-tmeuwuj/ <u4<iyyiu- •,;;.: jta.iwiwijam.uuiuaH grown no.apon.e-uvatTuerii.sprflu.jaies osce
hormones j?$tt. resistance exercise ecftance neluier tratoing-Induced
aiiiiactwJteVbouereriiodellrjgJJinormaihurflaQ.volunteersJ)jp/7i
muscle hypertrophy nor strength of jtteelbcw flexors. ] Affl fhyiloi.
~
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:.'""<• {f""-b. "^'.-'^*-*I2o¥^?^^tterM'Gtmrt^nOD1WMm^con!lfi<:livef:issu<:l!leSi:relse'
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6, Ch«a Y, Zajac. JD, A&cLcan HE, Androgen regulattoa of siUelUte celt 26. Godfrey RJ, Madgwlck Z, Whyte GR The exeKlte-lnduced grovrth
27, Wee }, CharltOJi C, Simpson H, *t al. GH tecretton in acute exercise
on nruscleiiprotetn synthesis )n royolonlc dystrophy. Arm Nettro1>
miyiesurt in post-exercise l^ralysjj. Gnwtfr HermIGPRef.2KSil5(6);
19«6;20(5)4.?0i5jj;i'!1 -^g 'Jj "\-( -jl>. ^ ^'"^'s ]'"' •„; \
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TestostetOTie Irtjection stlajulatesTiet protein synthesis but not tissue
aralnoacldtngesttonon acutehorrdonalresiponsedurtofrasInglebQitt of
amtno xtfiTmsgortzgnt ffiiysiai, 19S&27S(5 pt l)SE8«4-E871.
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exercise towWtaed men, NKfr/ffoji. 200S£2(-l)367-375.
9, Fenan<toA^'5rieffieS-MooreM,Paddon-7on«sD,WoIfeRR,UjbaflKf. 2.9, B!rfSP,TkpeBBliigKM,MadnoFEI^uidcajbohydrate/issentIai8rnlno
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12. Ka^nf^,^^<>^D^<^^^^G^^Aor^^^^'&\iSlSa-i&i
grofrth factorSystem tn myogencsls.jEMrfoirEei'. 1996;17(s):*Bi-Si7.
resistanceandendsranceexercbe to tiiefojslateon slgnalUngmolecttle
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14 "Rage DR, Jin IB, Va TH, Hoffinan AR, .Marcus R. Lacfc of effect of
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GH-msntIfl-5lke grovrth Bctor expression in resistance-trained elderly
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