Physiology 01

Sport and ....

Exercise
'/ion

Larry Kenn~y
k H. Wilmore

w~
vid L. Costill
Intro~.,.. Jction
Much Of the history of exercise physiology in the United States can be traced to the effort of
a Kansas farm boy. David Bruce (D.B .) Dill. whose interest in physiology first led him to study the composition
of crococtile blood. Fortunately for what would eventually grow into the discipline of exercise physiology. this
young scientist redirected his research to humans when he became the first research director of the Harvard
Fatigue laboratory in 1927. Throughout his life he was intrigued by the physiology and adaptability of many
animals that survive extreme exer•
cise and environmental conditions:
but he is best remembered for his
research on human responses to
exercise . heat. high altitude. and
other environmental factors. Dr. Dill
always served as one of the human
·guinea pigs" in his own studies.
During the Harvard Fatigue Labora•
tory·s 20-year existence, he and his
coworkers proctuced approximately
350 scientific papers along with a
classic book titled Life. Heat. and
Altitude.8 Dr. David Bruce (D.B.) Dill (a) at the beginning of his career; (b) •• director
of the Harvard Fatigue Laborato,y at age 42; and (c) at age 92 )tlst prior to
After the Harvard Fatigue Labora- hls lourthret1'9menL
tory closed its doors in 1947. Dr. Dill
began a second career as deputy director of medical research for the Army Chemical Corps. a position he
held until his retirement from that post in 1961 . Dr. Dill was then 70 years old- an age he considered too young
for reliremenl- so he moved his research to Indiana University. where he served as a senior physiologist until
1966. In 1967 he obtained funding to establish the Desert Research Laboratory at the University of Nevada at
Las Vegas. Dr. Dill used this laboratory as a base lor his studies on human tolerance to exercise in the desert
and at high altitude . He continued his research and wri ting until his final retirement at age 93. the same year
he produced his last publication. a book titled The Hot Life of Man and Beast.10

T
h e lnu nan hody is ;m am .11.in g m achine. A~ you
im p ulses d own th e sp in al cord to the k b'll, O n n·ac hi11g
si1 read ing lh is int mduciion . co1111tl e~~ pcr ft·ctly
th e m usclc·s. th.-s.e n e u ro ns rell•;L~l" ch e rn ical 111es.,cn•
n1o rd i11;1ted ;md i11tegra tn l en• n ts are occu r•
gen th a t nnss th t· gap between th e nen·e ;md 111usd e,
ri ng sinmltaneolL~ly in rou r body. T h e,1.e n ·e n t.\ ;1Jlow
each ne u ron exc-itin g ;1 numbe r of indhid ual nmsd e
complex functions. su r h a.~ l1e,ui 11g. s.eeiug. b re;uh iug.
cells or fihe rs. O n n· th e tll" n ·e impulses crrnc., thi., ga p.
a nd in fonn a tio n proces.sin g. to n ,ntinut· \\i th o ut <Ill }
they spread alo n g the le ng1 h o f 1•;1eh nms.cle lil>er ;m d
cun:1e:iotL\ effon . lf yo u stand u p. wal k o ut the d1M,r. and
.itt ac h lo spedalizrd renvto ~. Bindi11g o f the m e~
j og around the b lock. a lmos1;ill yo u r lMidy's ~)'Stems ,1ill se11ger to its n -,;·epmr .'1-1.·L, iu to motion a seri es of ste in
be fil lled in to ac1ion. enabli ng you t,, sufft•s.,full ys hift
th a t ac th,u e th e musde fi h1·r•s co nt rnct ion processe.'I.
from rest to exe rc ise:. If you continm· this routin t· n-gu-
wh ic h imu h·e s11t·cifif pro td11 nml,·n1Ie-actin and
larly for Wf"d :s or mmu hs and grnd u;1lly innea se the
myo.sin--au d a n ,·lalx1r.lle e 111·rg)· S)'Uem to prmid e
du r.uion a nd imensity of yo ur jogging. yo ur bod >· \\i ll th e foci n ecL~\.ll)" to s11s1.1in a single n mtr;tt"tio n and
;1dap1 M> th at you fa n per fo m1 bc.'tl t.''1". T h e n ·in lit: th l· .subseque nt contr.tCl io ns. It i.\ at this lc"'\·d th;u other
two b;t-\ic com poncnl5 of th e study of exerci_,e ph~-sicil• m o le cul es. su ch as ade nosine tri phos pha11· (ATP) and
,~,._,-: tl w acm e respc.m scsof the lx1dy to cx1·rcL,1· in all it-,
pl wsplmcreali ne (PCr). becom e critic;il for prmid in g
fomu a nd th e acL1. p1at io 11 of 1hose ·'}'5te111s to repeated tlw ene r1-,•y n ece s.\itl)' to fud con trnciio n.
nr c hro nic ex,•n-ise. often called cxt·rci'\t:" tr.tini ng.
In s11p1Min of th is sus1ai ne d ;md rh ythmic musc u\;,r
Fo r •·xampl,·, as th e point g uard d irecL, her le am conu,1ctio11 a nd rel;1x.ati o n. mu hip le additio n al 11)'51e1m
d m•m th t· b..L,kr tba.11 cou n o n a fa,t break, her lxMlr .ire t·;11le d into ac tio n. for e x.a mplt·:
111 a kes 111a n y ou lj us lnu, n ts 1h a1 re q u ire a se r ies of
complex intc r.inio ns imuhi n g m a ny bod y .'1}'5 le 111.~. T he skdt·tal syste m pnJ\idn th e b,1.sil" frnm cwurk
Aclj1l\tm e 111.~ occu r e\'en a l th e cellula r ;md m o lecular aro un d wh ic h muscles act.
le..-els. To e nable th e c1M1rd in a wd leg 1nu:1e:le ;1etio ns T iu- cardi,,\~L'ie ul;u·s),1.te m d elin :r:s fue l to wo rk-
as slu: m m·t·s rn pidlr d o\\1.l roun . n er\"e cell\ from th e
ing m11scl1· ;md to ;1II of the cells o f the body and
hr.tin. refc1n·d to a., moto r m·ttru n.~. conduct dec1rical re m on-s \\<LSI(' pro c\ufts.

Ail lnttoduction to Exercise and Sport Physiology

T he ca1dim.1.'1C ular and rt$pird lOI)' ~)1ile 11L~ wo rk

toge th e r to provide oxygen 10 th e cells a nd Acute and Chronic
rt• mm·e carho n d ioxide. Responses to Exercise
T h e inl t.1,'lllllt"llt.111· S)1'- le m (ski n ) help-, maimai n
T h t, study of exe1·cis1· and spo n ph p,iolugy in'l"oh-es
lxxly te mper.u u re by ;1lhmin g lhe exch an ge of
heat betwee n the body a nd iL~ surroun d in gs. learn in g th e con cepts a.~soci.i ted wit h two dist inct
exerci.Q• p att erns. First. exercise phy:<io logisb are nften
T he nen·o us an d e ndocrin e sr1e m s n1t1rdin.ite
n m cemt:d \\ith h ow th e ixMI)· responds to an i11cli\id11;1I
thi., acti\i ty. while hel p ing to ma in1ain fluid a nd bout uf exercise. such a.~ n umi n g o n a t readm ill for a n
elcnro lyte lmla n ce and a.'i.~isti n g in th e regul.i- ho u r o r lifting weighLs . .-\n indi\i du al bout u f exercise is
tio n of blcMMI pressure.
callee! acute exercise, and th e re.~Jllm scs to th.it excrfise
For cen1u ri1·s. scie ntisL, ha\'e smdi ed how th e human IMm t aJ"e refe rred to a.~ acute res pons.es. Wh e n exam in-
l1t1d y f1111ctio ns ,11 1·est durin g h ealth and dise;1.,e . ing the ;1n 11e response m exl·rcise. we· are con cen1ecl
Durin g the p a.~ t I 00 years or so, a specialize d g roup of \,i th !11: .~ ~~~(-~.:t,n'.~~~li~',te rt$J>ol ns~ m . ;md som e limes
 Af1 Introduction to Exercise and Sport Physiology

23 of 642
The cardi1>\-a.~ular and respir.1!01")' S"}1ilt·11L\ work Acute and Chronic
together to pro\'ide oxygen 10 the cdb and Responses to Exercise
re mon: car bon dioxide.
TI1e inteb'lnnenta~· system (skin ) helps ma intain
body temper.1ture by allowing th e exc h,'.uge or
h eat betwee n the body and iL~ surrmuulmgs.
T h e nen·o u~ and e ndocrin e systems c oordinate
~~:: ;: : L'.~:,~:~-:: ;~~E::£!~;~.I: ;.~: :\~:
concen1ed y,"Jth ]i cw: 1he body responds 10 an indhid11al
bou1 or ext·rcise, suc h a.\ nmning o n a treadmill for an
thL, a cthlty. while helping to maintain nuid and
ho ur or lifting weights. An indi\idual bout of exerci~t: is
rlectrolyte balance :md assisting in th e reb'li la-
callcd acute exercise. and the n :.~ponses to 1ha t ext·rcise
tion or blood p rn., ure.
hmu are n ·krrt·d to a.s acute responses. \\'hen examin-
For centuries, sciemL,L~ have studied how the hum:m ing th e anue n·spouse to exercise. we arc concen1ed
bo dy func ti o ns at rest during h ealth a nd di., ease. y,"Jth 1he l><Kly"s im m t·dia te respOIL'ie to, and st1metimcs
During tlw past 100 yt,ar.1 or so, .1 spedalized grou p nf its rrfm·ery from, a single cxerc L~e ho ut.
p hysiologl~L~ ha\·e focused 1heir studies on how the lx1dy The otl1er m;tjor are a of inten:sl in exerci:o;e and sport
fouc1i011 s during physical activity and spun. T his i111ro- ph~-sio loi-,'}' is h ow the lxxl y ropt111d\ m·er time 10 the
d11c tion prt·.'1e11L\ a historica l o\'eniew of eu·rdse a nd s1n·.s.~ or n :peated bouL~of exercise, stime1i1m,s referred
sport physin lo1,')' and the n explains srnm· ba.\ir n mn·pL~ to as ch ronic adaplalion o r training cffec1s. \\rhen one
th at fonn the fo undation for the chap1en1 th:11 follow. pt·rfonns regular exerci'il· m·er a pcri,KI o f days and
,wc k.,;. the lx1d y adap ts. T h e ph>~iolobrir;1l ad:1pt.atio ns
1hat occur "1th ch ronic exposun· lO 1·x1· rci'>l· or trnin-
Focus of Exercise and i11g impro \·e ho1h exercise c:tpad1y;md 1.fficic n cy. With
Sport Physiology rcsist:mce 1rnining. th e nULS("les bt-rom t· strongc r. \\"ith
aenibic trJining, 1hc heart and lungs lx.·rotne more e lli-
l::xe1TiSt.· and sJKln physiolo!,,')' ha\'e e\"ol\·ed from the cie nt. and e ndur.mn· cap.1d1yof 1hc 11111., des increa.~es.
fomlament.il disciplines of anatolll}" and phr inlob~-- A~ disc1t<iM1l later in 1his intn)( hll'IOI"}" d1apter. th ese
.-\11;11omy l~ th e study uf a n o rg-.tnL~ m·., stru cture. or adaptations a re h ig hl y spcdfic tu the t)l >c of tr.tining
morphology. While ;m:11omy fon1ses on the lu~ic stn1c• th e person does.
mrc- of\"ariou.~ body parts and their i11te1Trl.t tio nships,
phy!>;ology Ls th e study of body Jm1 rtio11. Physio lo,.,risLs
smdy how the hody·s org.tn sr1t·11Ls. ti s.~ues, cell,, and
lhe mo lecules within cells work and huw thei r func•
G)t.£41b
1io ns arc- inte gr.11cd to n·g11bte the lx1dy".~ imern:11 Exercise physiology evolved from its parent disci•
elJ\ironmt·nt. a pron·s.~ called h o meosta.,is. Bec.111sc pline, physiology. The two cornerstones of exercise
physiol••b~' foc 1N·s on th c fun nio ns o f body su-u ctures, physiology are ( 1) how the body responds to the
und erstand in g ana to m y is es.~ential to kan1ing physiol- acute stress of exercise. or physical activity. and
Ob'}"- Furtht'nnore. lxith ana10 111y ;u1d physio]Ob')' rely (2) how it adapts to the chronic stress of repea ted
o n a working knowledge of biology. ch cmi.•IIT, phr ic.s. bouts of exercise . that is. exercise training . Some
:md other l>.t~ic science.~. exercise physiologists use exercise or environmen-
Exercise physio logy is the smdy or how tlw body•s tal conditions (heat. cold. altitude . etc .) to stress
fun ctimL, are alte red whe n we are exposed to exe1-cise. the body in ways that uncover basic physiological
a challenge 10 ho m eosl;t~i-~. &canst, tht· 1·1J\inmmc111 in mechanisms. Others examine exercise training
which o ne pt·rforms exercise h a.~ :1 l;1rge impafl. cmi- effects on health . disease. and well-being. Sport
ronml'ntal physiology has 1·111,-rgcd ,t~ a ., uhdiscipline physiologists apply these concepts to athletes and
of excrcL<it' phys inl,,),,')", Sport ph ysio logy f11rther applies sport performance
the cm1n·p1.,,,f e:r.:erc L~e phrsi(1\ob'Y to e nhancinJ.:" sport
11t·rforman ce and optimall)· Ir.tining th e athlelt:. Thus.
spun ph~11io logy derfres iL~ principle.• from exercise
phy-.inh~•y. lkcau<it' exercis,.• physio lob•yand spun physl- The Evolution of
olob'}' are so c\os,.•ly rel,ued and integr.tte d . it is oflen Exercise Physiology
hard to clearly distin gui.sh l1t·1wec n them . Ber.1use 1he
same unde r lying scie utilic principles apply. exercise To suukuL~. conte mpor.111· e xercise ph>11iolu1-,'}' m:1r
:md sport physiology are often consid e red 1og1·tlwr, a.s st.-em like a ,-Jst collectio n or new idea.~ llt:\'l:r before
1heyare in thistt·xt. studied with rigorou~scientific scrutiny. On thc conu-.iry,

Physiology o! Sport and Exercise

l'.owen:r. 1he infonuation in this l)()(ik n·presents the di~cm·1·r t he exisience ofindhidnal nnt,;c le fi11t· n1 until
hklong dfu rts of hundre<h of outst.andiu g scil'ntisL• Dutd1 .'w:i.,111is1 Anton ,,m
l.tTHWenhoel.. i111 n)(llln"(I
who ha\"l: collecti\·clr helped piece toi;t·th e r what we the micrnscn1x: (c:1. 1660) . How 1lws1· fibt·rsshonem:d
cmTently I.now about tht: s,.:it:n cc of human 111on·ment. 1 1
T hr tlwories and h n1ot heses of 1111)(len1 ph~"SiologisL• :::~~1;t:::~t:~1!o;~:h\\~c:\,:\ :,1;~l::.;1i~:1•1.1: 1~):1t1~:~;-::::~I~.~:.~~
h,l\"e been sh<1J1t·d by the t·fforls of scie 111i~L~ who ma,· in~s of muscle proteins could Ix· studied by clt"ctro n

;~i\~!£:,:}ig~ii'.II:~:}i:~\}qff,ifIs
microscopy.

Historical Aspects
a nu~1be~ of kt·y scientisL\ :md many pin1tal sd1·111ific of Exercise Physiology
~o nt11lmt10ns th ,11 hruugh1 al)(mt sib'lliric;mt ,1d\"ann·s Ext·rci.'IC.· ph>-si,,11,b'}· is a rd:uh·e n ewcomer to th e world

!:~~;:;: : :~~l:~f:~:~:~~: ~:I:~:~f;~.~: ~: : :~~

m_ou_r kr~owled ge of th e- p_h ysiological resp onst·s to
1·xnnse. nu- folloY.ing secuon rcnects on the his1on ·
:md onj1N a f,·wnfthe peo ple who sha pcd th e field 0 J
 1
1;i;;.~io!~;.:·&:(.7'~L; exer~i.-.tc ,;hysiol:'b'}"and s1~u·1 ph}~j.. The Evolution of
u lo).,') ' an: so dosdr related and in legrJted, ll i., often
hard to clearly disthlb'llillh between th e m. EkcatL'I.C: the
Exercise Physiology
same underlyi n g scientific principles apply, ex1;n:i.'I.C· To s1udt·nt,. contempornry exercisr ph p1iology may
and spon ph}'lliology arc ofte n cm1.~idcn·d together, il' seem likr a \<L~t collc<lion of new ide:L\ n en:r before
they are in this text. llt udied \\i lh rigoro1u sdt·ntific scrutiny. O n thecontr.tl')',

Physiology of Spo,t and Exercise

howe,,e1; the infom1ation in this hook repn·M:nt~ 1hc di.~o,·e r t he existence of indhidual 111\L'\('ie fiberi; until
lifelo ng e ffort, of humln·ds of m11s1andi11g -~ ienti.~t, Dutch scicntillt Anton ,·.in Leeuwenhoek introduced
who ha,·e ro llecth-dy hel1 >ec\ piece together what we the microscope (ca. 1660). How these fibers shortened
C111Tcntl y ki10wabout the K ienre of human 1110\"emem . and c rea ted force wou ld rr-111:1i11 a lll}"lllery until lhe
The theories aud h)1>oth escs of mode rn ph~"lliologist~ middle of th e 20th CelltUl)', whe n th e intricate work-
h:we bet:n sha1)(•d by tht: dfort, of Kie nt ist~ wh o may in~ of muscle proteins could he studied by del·trun
he long forgotten. \\'hat Wt' cmuid cr orib-inal or m·w is microsco py.
mos1 often :111 assi111il:1tio11 of pre,ious findin~ or tht·
applir;uion of basic .'1cien ce to problems in cxerci~,
ph ysiologr A, \\ith t:\'e'1' diKipliuc. then· are. of course,
Historical Aspects
a number of kC)· 1\Cientillt~ and m any pivotal srit·ntific of Exercise Physiology
contributinns th:u hruuglu abo ut .,ib'llificant ach-;mces
Exercise ph}"llioh'b'}' is a rclati\'e n L•wcomer to th e world
in o u r knowlt·dge of the physiological respo n.'IC.s to ofscience, o1hho ugh as early as 1793, a celebrnted paper
exerci~.-. l11e follo\\i ng :1,ectiou rellcct, on the hilltory by5ei,.'l.1in and L.1,ui.~icr dt.'!l{Tibcd theoxygt·n comump-
a nd on just a few o f th e people who sh aped thl" field of
tion of ;1 }Uung man:\, mea.~ured in the rr-sting state and
exerrisc physiolt'b')", It is impossible in thi.sshor1 ll-t:Ctio n
while he lifted a 7.3 kg (16 lb) weig ht nu merom tillle5
to do justice to th e hundn·ds of pioneering llCil"nti~t,
for 15 1nin. '" At rest the m au 11:1,ed 24 L ufoxyge n per
who pa,l:d the way and laid 1he foundation for n1oden1 hour (L/ h). which innea.•·,ed 10631./ h during exercise.
exercise.-- phpiiology.
l..nuisier belicn·d th:u the 5ite of oxygen utili1.ation
and carbon dioxide production ,1-.1., in th e lungs. Even
Beginnings of Anatomy though thi~ con ct'pt wa., donhlt·d by oth er ph}"lliol11b'l.\t,
and Physiology of tlu, time , it remai,u·d acn,pted doc trin e u n til the
middlt, oftl1e 1800s. when lle,-e1~1I German phr iologi.\tll
O n e of the earlit·st de.~ rip1io1u of h11111;111 a natom y
demon~trnted that combustion of oxygen occurrt.-d in
and phys iology .,,-J.s ClaudhL, Gale n·., G reek u•xt D,
ti,o;,,ues throughout the e ntire body.
/mri1u. publislwd in the first centul)'- A, a physici,111 to
Although many ad\'anct·.\ in the undef!ltandin g of ci1~
the gladi,uors, Galen had :un p le oppon1111i 1y to s1udy
cul:uio n ;md respirntion tKCllnL"II during th e 1800s, few
and ex1>eii1uent on human anatomy. Hi., t11eori es of
effort, \\·ere mo1de to focu, o n the physiol~'}' of physicoll
anatom y a nd ph}"lliolOb'}" werr- .' IO \\idcly ;iccepu·d 1h;11
acti,ity. Howe\"er. in I 888. an appar.uus \\-Js de>\Cribed
thq re111:1ined unc hallenged for nearl y I .400 years.
that enabled scientist\ to 5ludy subject., du ring 11101111-
Not until 1he 1500s were any truly sib'llifica111 cont rib11-
1;1i n climbing. n·en th ough the subject, had to carry ii
1io11s lllade to the u nderstanding of both th e stn1c1t1re
7 kg ( 15.4 lb) -g.111t>m eter- on th e ir bark.,."
and func-tion of the human body. A l,111d111 ark text
Arguably the first publi5hed textbook on exe rcise
by And reas Vesali1L,. titlt·d fflbrim H11mn11i Corpori1
ph)"lliology. J>hpiolOf!J of &xlily Exnrll,. \\'all wri n en in
/Slrnr/11~ of th, lfom1111 &,dy/. prnc111ed his find ings
French hy Femam\ LtGrnnge in l889Y Considering
on htmian ana tomy in 154'.\. Ahhough Ves.1 litL,• book
the small :1111011111 of research on exercist: lh:u had been
focn,ed primaril~· o n anatomical descrip tions of \'ari-
ronductcd up to that ti111e. it i., int1ib'l1ing to read the
nus organ., , h e occa.,iona lly att1·mptt.-d to explai n their
alllh or's ar couu ts of .suc h topics a.~ -Mu.~ 11l,1r \\'ork. •
functiom a., well. Briti.,h historian Sir !-.lichacl Fos 1r-r
-1-·:uigue.- "Hahitua1io11 to Work.- and -The Offire of
sa.id. -111is book ill lhe beginning. not o nly of modem
th e Br.1in in Exerci.'!.C.~ l11i.s early :uten1pt to explai n
anatumy. but of modern ph)"llioh'b'}'- It e nded. for a ll
th e rnpo nsc to exerci.'I.C \\-.is. in many \1.1)-S. li mited to
time, the long reign offourteen ce nturi1·., ofprect-de111
speculation and 1heo11·- A.ltho ugh some b:lSic ronceplll
and beg;m in a t111e seJL'\.C the renai.'l.'l.lllce of 1ne di ri11e-
(p. 354). 1·' of exercillC biochemislT)' Wt•re e merging ;11 that t ime,
l.:1Gr.1 nge wa., 1p1ic k to admit th:u m any details wt·rc
Most early <llle111pts ;11 explaining phy.sioloi..'}· wt·re
5lill in th e fonn;uivestagt·s. For example, he stated that
either incorrect or so \'ab'lle th at they could be n)J}sid-
-,ital combtt\lion [t·nergy 111etabolism ) ha., become
ere d no more th;m s1>eculatiou. Attempt~ lo explain
\ "CT)" complicated of l:ue: we 111ay !I.I)' th.it it i.,.so,newhat
how a mu~ le gc11e1~\ll.'!I force . for example. were 11su;1ll y
perplexed. omd th:u it is d ifficult to g h-c in a few word§
limited to a d ewription of it, ch ange in lli1.c and shape
,1 dear aud concille s11m111tn)' of it. It i., a ch apler of
d u ring action bccau'IC obsen~llions were li111ite d lo
what could be llecn .,,;th the 11.1ked eye. Ye t fro111 su<·h ph}"llio\Ob'}' wh ich is being rewri1te 11, and we cannot at
th is 111o m ent fom1u l:1u, o u r co11cl1L,io11~~ (p. 39!"1)_ 1~
oh:1,erntions. liieronyinus Fahricius (ca. 1574) su g-
ge.sted thal ,1 mu.,;cJe·s contr""Jctilt· power resided in iL, lkc;nt~ the earlr 1ext hy l.:1G1,mge offered only
fibrous te n dons, not in it, - nt•sh .- A:n:urnnist, did not limited phys io logic-.il iusight, n·g;u-ding bodily func•
lions during physical ac1ili ty, it m ight he argued that

A.V.HIII
OctOber 16, 1923. was a significant milestone in thi> h;"''""' ,,f ,., ~~ - " ·" -
 A.V. HIit
October 16, 1923. was a significant milestone in the history of exercise
physiology. A.V. Hill was inaugurated that day as Joddrell Professor of
Physiology at University College, London. In his inaugural address he
stated the principles that subsequently shaped the field of exercise
physiology:

"It is strange how often a physiological truth discovered on an

animal may be developed and amplified. and its bearings more
truly fou,d. by attempting to work it out on man. Man has proved ,
for example, far the best sub}&t for experiments on respiration
and on the carriage of gases by the blood. and an excellent
subject for the study of kidney. muscular. cardiac and metabolic
function. . Experiment on man is a special craft requiring a
special understanding and skill. and 'human physiology,' as ii may
be called. deserves an equal place in the list of those main roads 1921 Nobel Prize winner
which are leading to the physiology of the future. The methods. of Archibald HIii (1927).
course, are those of biochemistry, of biophysics. of experimental
physiology; but there is a special kind of art and knowledge required of those who wish to
make experiments on themselves and their friends , the kind of skill that the athlete and the
mountaineer must possess in realizing the limits to which it is wise and expedient to go.
Quite apart from direct physiological research on man , the study of instruments and
methods applicable to man, their standardization, their description, their reduction to
routine, together with the setting up of standards of normality in man are bound to prove
of great advantage to medicine ; and not only to medicine but to all those activities and
arts where normal man is the object of study. Athletics. physical training. flying . working.
submarines, or coalmines. all requ ire a knowledge of the physiology of man. as does also
the study of conditions in factories. The observation ot sick men in hospitals is not the best
training for the study of normal man at work. It is necessary to build up a sound body of
trained scientific opinion versed in the study of normal man , for such trained opinion is
likely to prove of the greatest service. not merely to medicine, but in our ordinary social and
industrial life. Haldane's unsurpassed knowledge of the human physiology of respiration
has olten rendered immeasurable service to the nation in such activities as coal mining
or diving; and what is true of the human physiology of respiration is likely also to be true
of many other normal human functions ."

During the late 1800s, many theories were proposed to explain the source of energy for muscle
contraction . Muscles were known to generate much heat during exercise. so some theories suggested
that this heat was used directly or indirec tly to cause muscle fibers to shorten. After the turn of the
century, Walter Fletcher and Sir Frederick Gowland Hopkins observed a close relation between muscle
action and lactate formation .'' This observation led to the realization that energy for muscle action is
derived from the breakdown ol muscle glycogen to lactic acid (see chapter 2), although the details
of this reaction remained obscure.
Because of the high energy demands of exercising muscle, this tissue served as an ideal model
to help unravel the mysteries of cellular metabolism. In 1921 . Archibald V. (A.V.) Hill was awarded
the Nobel Prize for his findings on energy metabolism. At that time. biochemistry was in its infancy,
although it was rapidly gaining recog nition through the research efforts of such other Nobel laureates
as Albert Szent Gorgyi. Otto Meyerhof. August Krogh, and Hans Krebs. alt of whom were actively
studying how living cells generate and utilize energy.
Although much of Hill's research was conducted with isolated frog muscle. he also conducted
some of the first physiological studies of runners. Such studies were made possible by the technical
contributions of John S. Haldane, who developed the methods and equipment needed to measure
oxygen use during exercise. These and other investigators provided the basic framework for our under-
standing of whole-body energy production, which became the focus of considerable research during
the middle of the 20th century and is incorporated into the manual and computer-based systems that
are used to measure oxygen uptake in exercise physiology laboratories throughout the world today.

Physiology of Sport and Exe,cise

the third edition nf:1 lt."Xt by f.A. Bainbridge titled 'fh, the phy:,;iology of eudurnnce t:xercise ;uul described
PhysiologJ ,if ,\J rurnillrf,'xm-iM~honld Ill' comidert."d th t." the ph~11kal requireme nts forsutTeM in en·111s such ;n
e;1rli~t scientific text o n thissubjen.' Interestingly, that dislann: numing. S.:m1t• of the 1110<11 0111.'it:111ding HFL
third t·dition "-,._~ written by A.V. Bock and D.B. Dill, at
tht." requt,.~t of AV. Hill. three kt,y pioneer,1 of exerci.'IC
phy11iol~1-· di.'ICussed in this intmdm·tory chapter.
~:;;c~:jfi::j~;.:~::,:•~1~:::.l:~el~~:!l!~~,:::;i \:~::\~:.1
on Whitl' Mountain in C.·difornia {v.ith ;m a ltitude of
3,962 m. or 13.000 ft) . These and other stud it~ prmided
Era of Scientific Exchange the founda tion for fmure in\"estig.uions on the effect5
and Interaction of the en,irmunt,ut on php•irnl perfonn:1nce :ind in
exerci!l.t' :ind spun physiology.
Frnm the t•arly 190(k through tht." 19'.\0s, the medical In iL, early yean;, the HFL focullt'd primarily on
and sdentifo: t·n,iromnent in the United States " "<l.~ gene-ml problem5 of e xerci'll', nutrition, and health.
d1a11ging. This "~\.~ an er.1 nf ren1lutio11 in the edu- For e~1'.nple: tl1w fi~~.t..~'.\1di~~ '.''.'. t·xercise ;md :1gi11g
-
26 of 642
Physiology of Sport and Exercise

the third Nli1io11 ofa 1ext h}· EA. Bainbridge tith:d Th~ the physiology o f l'ndur,mce cxcrd~ and described
PhyJ.iology of ,\11urn/nrf.x,rri.vs hould be considered the the php;ical requirement.~ for SII C<"C'S.~ in C'\·e n t.~ .'l uch as
earli r:st scientific text on thi.c..subjcct .' Interestingl y. that dist.ance nmniug. Some of till" mosl OUl.'11.UJdiug HFL
third edition \\-.t.S written by A.\'. Buck a nd D.B. Dill, ;u irffestig-dtions were conduc ted 1101 in dw l.1ho r;i1t111'
the reqm:st of A.V. Hill, three key piunecr5 of ext•n-iil<' bu1 in the Nei.·.tda desen, on the Mi Mis.~i ppi ddt.i, and
phyi1ioloi,,,1• discm sed in this introductory chaptt·r. o n White Mo untain in California (\\ith .111 :1hitmk of
3.962 m , nr 13,000ft). ThC'5C' and othe rstudit'5 pro\'id,·d
Era of Scientific Exchange tht· fouud:nion for future in\"estigations on lhc dft-ct.~
of the emironmcnt on phy,iical performance and iu
and Interaction exerci.~- and sport php1iol1)j,,'}"-
From the early 1900s throu gh tht• l!l:-\0.<1, the m edical Jn its early year.1, the HFL focu.'ll'd primaril>• on
and scientific cmironmcnt in tht· United S1a1cs \oo.1., genern.l problem.'I of ext•rci!\C, nutrition, and hea lth.
changing. T hi.~ wa., an cr.t o f rc.."\·olutio n in 1he t"rlu- For example. the first s111dit•s o n exercise and aging
rn tio n o f mcdir.tl students. led by chaugt·s a l Johns we re conducted in 1939 ~ · Sid Robinson (see figure
Hopkins. More medical and grnduate progr..1111.c. h;l.c..,:d 0.1 ), a student at the Hfl.. On th e l>.1.~is of his studies
thdr cduc:uioual c nck;1,ufl! on the Europea n modd of uf subject., rnnging in age from 6 to !I I ye;1r.1, Rubimron
experimentatio n and dc..-,.·dopment of Kie ntific insights. dc!ICribed the effect of aging on maxim;1\ heart rntc
There were imponant a,h-,mces in phy,iiology in areas ;md oxygen uptake Y But with tht· ousct of\\'orld War
such as bioenergetics. g;.t.~ exch ange. and b lood chem- II. Hc nde rllo n and Dill realized 1hc l·ffL"s JX)te ntial
istf}' that sened as the ba.~is for ;ich;m ces in the php;i- con1rilm1ion to the war effort , aud rt•st·;u-rh at the
olt)j,,'}' of exercise. Building on ,olh1bor;11ions forged HFL II Mlk ;1 different direction. Han,ml Fati).'lLe Lah
in the late 1800s. internctions among l;ibordlories and scicnti.~ts ;md support perso nnel were instnuncntal in
scic111ists we re promoted, and international me-dings funning ne w lahor.tlOriC"S for the Anny, Na,)·, a nd Ann}'
of org-dni;r.ations such as the lnter11a1ional Union of Air Co11>s (now the Air Force). l11ey al!ltl publi~ hcd the
Phrsiologic.11Sciences c reated an :um ~ph cre for free me1hodol()j,,rit.'5 llC'CC'5.Q.I)' for rdemnt milil.ll)' re5e,1rch,
scientific exchange. discm sion. and ckha1c. mt."lho<b th at arc s1ill in use throug h oul the world.
Toda y's exerci.'le physiology student.'I "·ould be
amazed .1t the me thod~ and devices u:1.e.·d in the early
Harvard Fatigue Laboratory days of the Hfl. and :u the time and elll'll()' commit-
No laboratory lrns h;1d nmre impacl on !ht' field of ted to conduning research projec ls iu those da>·s.
exerci.,e phys iolob'}' th;m th e Han-a.rd Fa1igu e l ,;1bor.1- What is n ow acco mplished in milli:1.e.·coud s wi1h the
tory (HFL).A \'isil by A.\I. Hill to Hamud Uni\'l•nity in a id of computers a nd automatic an:, lpcrs lite rally
1926 h;id ;i significant imp;Kt on the foundin g and 1•;1rl r demanded dap~ of effort by HFL scienti.~,.~. Me;t~un:-
actidtics of the HFL. whkh \\;t.~ established a yt•;ir ];u c r menl5 of 0X}'gl"II 11pt.1ke during exerci.'l.l', for cx;unple.
in I !127. lnterC"Stingly. till" 1·.1rly ho m e oft he Hfi. "'l~ the required colkcting ex pire d air in Dongl;1.~ l>.1gs a nd
hmwmenl of Han-a.n:l"s Busim:ss School.audits st;1tt•d analFing it for oxyge n :md carlx m dioxide hy ming a
early mi.~.~ion wa.~ to comhKt n·se:u ch on -fatigue- ;md m;muallyopcr.Ut-d c he mical analyzer. "i 1hout the hdp
othl'r ha;r,;uds in industrr, Creation o f this labor.uory ofa computer, o f courllC' (see figure 0.2). ·n1e :mal ~'llis
w:1.~ due 10 the insightful planning of world-famom of a si ngle 1 min sample of expired :1ir required 20 to
bioch e mist, Lawrence J. (LJ.) Henderson. A young 30 min o f effort hy one o r mo re labor,Uol)' worke n .
bioche miM from Stanford Unil·c,,.i1y. D.1,id Bmce D.R. Today. scie ntist,; make such me.1snre m e nts a lmos t
Dill. " 't.~ appoi nted as the lint dirttmr of research , .i instantaneously and with little phrical effort . One
title Dill held until the Hl-"Lcloscd in 1947. mlt~t man·d :11 1he dedicati o n. dilige nce. and hard
A~ nol t"<l e:1rlier, Dill had :1iclt-d Arle n -Arlie- Bock in work of the HFl.'.~ exercise php;iolob'}' pioneers. Using
writing the 1hird edition of8;.1inhridge 's text on exercise the r:quipmt·III ,mcl methodsa"1il;1blc ;u th e time, HFL
phys iology. J,;1tcr in h is career Dill credited the writ- scientist.~ published appruximatdy 3:"10 rellC'arch papers
ing of th,11 tcxlix)ok with -.'limping 1he program o f the m·era20-yt•;1rpcriocl.
Fatigue l.abor;uory. - Although lw had little expc.·rit'IKC The HFL w.is an intdkc tn a l e 1wiro nment th a t
iu applit-d 1111111.111 physioloi.')·, Dill's creative thinking attrncted yo un g physio logistl'i and physiology doc-
and ability tn sunound himst·lf "ith young. 1alentcd toral stud1•nt., from all over th e g lobe. Scbolan from
scientist.~ n-c:ued an emironmcn1 d1.1t would lay the 15 countries worked in the Hl-1. between 1927 a nd
foundation for moclen1 ex,·n-i!\C and emiro111ncntal its closure in 1947. Most we nt on to develo p their
phys iology. J,' or example. l·ffl pen.onnd examined o\\11 l:1boratories and Ix-come noteworthy ligures in

An Introduction to Exercise and Sport Physiology

27 of 642 AA Introduction to Exercise and Sport Physiology

Ii .. rvard FaHgue laboratory and

(b) as a Harvard s1udent and athlete in 1938

'
.a.. -
-... .
'-'•
;IJ~p,-
' ~--::in "'° -
....
. • '
(a) Early measurements of metabolic responses 10 exercise requ ired the collection ol expired air in a
sealed bag known as a Douglas bag. (b) A sample of that gas then was measured for oxygen and carbon dioxide using a
chemical gas analyzer, as illustrated by this pho1o of Nobel laureate August Krogh.

exercise physiolob')' in die United Stales, including Adolph. Walter B. Cannon. Peter Scholander, and
Sid Robim1on, Hen11· Long.' llreel Taylor, Lawn,un· Rudolfo Marg-.nia. along with S{_'\"er.tl other not.able
Morehom;e, Rohen E. Johll!mn , Ancd Keys, St<.'\'Cll Srnudin a,i:m sdenti~L~ discusS(_·d later. Tims . the Hl-1...
Hon;nh, C. Fr.tnk Consolazio, and William H. Forhcs. plmllt·d seeds of intdlect at home and around the
Notable inten1;uional !ICit'nli.'ll'I who spclll time at 1he world 1h:11 resulted in :m ex plosion of knowledge and
HFL indurkd Augu.~l Krogh, Lucien Brouha, E<hmrd intt:n,st iu lhis new lit·ld.

Physiology of Sport and Exercise

{0)i•l41b In addilion to 1hcir work at GIH. both Hohwf1-

Chri.'l tensen and Astrnnd interacted with ph p;iulogisl5
Founded by biochemist L.J. Henderson in 1927 and at the Karu\insk.a Institute in Stockholm. s. . <cdcn , who
d irected by 0 .8 . Dill until its closure in 1947, the studied clinical applications of cxerci~. It i.'I h,..-d 10
Harvard Fatigue Lab trained most of the scientists single 0111 tht:" most exceptional coutributi o11.'I from
who became world leaders in exercise physiology th is in.'l titute, lmtJuna.'I St-r,r-mnm'.'I (figure 0.3r) rein-
during the 1950s and 1960s. Most contemporary troduction of th e biopsy needle (ca. 1966) to .'!ample
exercise physiologists can trace their roots back m11.~lc ti~~uc .,.,~,1.~ a phutal poi111 in th e study of huma n
to the HFL. 11111.~lc biochemi~try and muscle murition. This u·rh-
nique , which in\"ulves "l<lilhdrnwing a tin y .'l:Ullplc of
11111.~le tissue "l<lith a needle in.~rtc.-d into tht:" 1111L~lc
through a .~mall ind.'lion, wa.~ originally imruduced in
Scandinavian Influence the e:uly 1900.'I 1o s111dr mlL~ lilard~strophr. The needle
In 1909.Johanue.'I limlbcrg nt.ablished a labornturr biop~·cuabled phr1 iol01,-isl5 lo cond1Kt hislologiral and
,1,.,. h,,,.,..,,.." f,.,·,il,· h ,-,.,.,1;,.,, ,.,.,.,,,.,t fnr ~.-;,.,. ,;r,,- 1,·,,.-1,, . ..,·,. _,I· ,,r,-,.,f l111mm,rn'rh,•f ,n·. , l11 r" 11,.
 28 of 642
In :1ddit io11 to their work at G UI , both Hohwf1-
@t-J41b Christe nsen and Asu-;md inter.iclt'd with physiologi.d s
Founded by biochemist L.J . Henderson in 1927 and ;II the Karolinska Institute in Stockholm, Sweden, who
directed by O.B. Dill until its closure in 1947, the studied dinkal applic;itious of exercist'. It is hard to
Harvard Fatigue Lab trained most of the scientists singk out till' most exceptional contributio ns from
who became world leaders in exercise physiology th i.~ im titlllc. but Jonas Bergstrom's (figure 0.:ir) n·i n•
during the 1950s and 1960s. Most contemporary troduc1ion of the biopsy net•dle (ca. 1966) to i\alllplc
mu!IClc 1i.~sm· v,-.1.~ a pi,·otal puint in lhe smd y ofh111m111
exercise physiologists can trace their roots back
1n11K lc bioc he mi.,try a nd muscle nutrition. This tech-
to the HFL.
nique. whic h i1l\'Oh1:"s withdrnwing a tiny s.unplc of
muscle ti.•1sue with a needle inS(:rted into the nHL"W:le
through a sm:111 incision, v,•,1s orii,.rin;illy introd uced in
Scandinavian Influence the early 1900s w stndy muscnlardptrophy. The mxdle
biopsy enabled phr~iologi~L, to conduct histological and
In 1909.Joh;umcs Lindberg t:5tabli.,hcd a labo1-dtory
biochemical ~tuclic.1 of human 11111., ck before, during,
that bcc;unc ;1 fertile breeding grou nd for scicntifir
contribu1ion~ at the Uni\'enityorCopen hagt·n in Den- and ;ifter exercist:.
O ther im.1sh·e studies of blood circula1ion "·err
mark. Lindberg and 1920 Nobel Prize ,oinn<'r August
.'11bst•quently nmd uctt·d by ph}'lliologists at GIH and lit
Krogh teamed up lO conduct many cla.,;.,ic experimen ts
the K.1roli11sb Instit ute. Ju~, as the Hfl... had been the
and publ i.,lu:d ~ minal papen on topics r,mging from
mecca of exe rc ise pht'lliology researc h lx·twet·n 1927
the metabolic fuels for muscle to g-.t.s 1:xcha ngc in the and 1947. the Scaudina,i.a n labor.i tories were eq11,1lly
lungs. Thi, work w.t, co111i nul'rl from the 1930s into the
noteworthy beginning iu the late 1941k. J..fany leading
1970s by th ree yo ung Danes. Erik Hohwi1-Christcnsc11,
in,·estig;1tions O\·er 1he pasl :15 ye ars wen· coll;ibo1-;i-
Erling A.~mllli.~e n, and ~farius Nicl~n. tio ns between American a nd Sc:111di11a,ia11 exercise
A.~ a rcsuh ofcont:Kts between D. B. Dill aud August
phpiologisL~. No rwegian Per Schol;mdt•r illlroduced ii
Krogh. these three D;mish phylliologisL, c:1mc to the
g-.t., analyze r in 1947. finn Ma rtii K;1r.oncn publisht"d
Hfl . . in the 19». where they smdicd exercise in hot
a fonnul;1 for calcul;11ing exercise heart r,ue that is
cmironme nt., and at hig h altimde. After re turning
still "iddy usc."ll today. (For a more de miled listin g of
to Europe. eac h man established a separ..lle line o f
the Scandina,ian contribution.~ lO e xercise physiology,
rese:irch. Asmussen and Nidsen becaml' professors at
the Uni\·er.iity of Copcu hagen. where A,mu.~<1e n .1tud- consult A"rnnd 's rnil'w. 1 )
ic.-d the mechanical properties of ntlL'ICk and Nicl~en
("onducted studies on control of body temper,uure. Development of
Sod1 remained acth"c at the U11h1:rsity ofC..ope1 1hage n ·s Contemporary Approaches
AugtL~I Krogh Institute until their re tirements. Much ad,~.mccment in exercise phys iologt· must
In 194 1, Ho hwf1-Chri.~tenscn (see figure 0 .3(1) mon~d
be credited to impro,·e menL~ in tec hno logy. In the
to Stockh o lm 10 lx•come the fiT5t php io loi,.1• professor
la te 1950s, He nry L T:1ylor and EL, worth R. Buskirk
at the College of Physical Education at Gpmrn.dik-och
publis hed two semin;1 I papers" •~ describing the
ldrotL~h0giikol,111 (GIH ). In the late 19~. he teamed
oiteri;i for meaM1ring 111a.xi ma l oxygen uptake a nd
v,ith O le Hansen to conduct and publish a 11eril.'S of
t'Slablis hing that mea.~ure as die ~gold st:md:ird~ for
fh1:' stud it's of (";irbohydr.tte a nd fat metaboliMn du ri ng
c,1rdiorespirntory fitnL'S.~- In die 1960s, dcn~lopment
cxerci.'IC. T helle stud ies arc still cited frequently ;md arc of electronic ;111alpien. to measure rcspir.t tor)' g:.1.'1cs
nmsidered to be a111ong the fiT5t and mO!lt important
made stml~i ng e1ll'rb')' 111et;100lism much ea.~ier a nd
sport nu trition studies. Hohwi1.Chri.,tenscn introduced
more produc1i,·e than before. T his techno llJb•y a nd
Per--Olof A~tr.md to the fidd of exerci11e php;iolog)'.
r..1diotcle111elr)' (which IL~es r.1dio•trnns:rnit1cd sib'llals).
A.,tr..md . who conducted numerous studies rdatcd to
used IO monitor heart rnte and bod y tempern111re
ph~'llical fitm•.~~ and cndur.mcc capacity du ri ng the
duriug e xercise. were developed a~ a re:-su h of the
)9:,(k a nd 19fi0s, bec:une the direc tor of G )H after
U.S. space pn1gr.u11.Altho11g h such instrumen ts took
Holu,i..CJ1risten11en retired in 1960. Duriug his tenure
nmch of die l.1bor out ofrt'!IC'arch. they did not a lter
at GI H. Hohwft-Christensen mentored a 1111mlx•r of
the directio n of!ICientilic inquiry. Un til the late I !ll;()s,
o uL,tauding scientists, including Bengt S.1hi n, who w;l~
111o:11exercise phy~ioh>t,,')' studk'S focused o n the who le
tht· 2002 O lpnpic Prize v,iuner for his man y contribu- body's n-spo nsc to cxerci'IC. The m;tjority of inn.""Sti-
tions to tht· field of excrci.'IC ;md clinical physiology
g-..uions invoh-ecl me:1.~un,1nen ts or such \"i1riabks ~I'
(sec figure0.3b).

Es

An Introduction to Exercise and Spor1 Physiology

29 of 642 AA lr\lloduction to Exercise and Sport Physiology

l:l(C'11fd¥•1ki (a) Erik HohWU-Christensen was the first physiology professor at the College ot Physical Education
at Gymnastik-och 1drottsh0gskolan in Stockholm. Sweden. (b) Bengt Saltin. winner of the 20:'.)2 O lympic Prize. (c) Jonas
Bergstrom (left) and Eric Hultman (right) were the fi rst to use muscle biopsy to study muscle glycogen use and restora1ion
be!Ofe, during . and after exercise.

o xrge n uprnke, heart r.u e, bod y u:mper.uure, and 0.4 r) a l Wa~ hin1,,rto n S1a1e Uni\'e ri;it)' fi ri;t used mt~ and
swe al r,ue. Cdlular ~ Pon,,;es 10 exerci,;e received lit de mice to smdr musd e meta bolism and 10 examine fac•
anention. tors rdate d 10 fatigue . Their public-, uions a nd their
In the th ree biochemists eme rged "'ho tr.tining or gr.tduate a nd pnstdoctornl student~ ha\'t'
we re 10 ha,·e a major in1pac1 on the field of exerdse resuhe d in a mrn·e binche111ic-,tl approac h 10 e xercise
phys iology. John Holl~· (fig\lre 0.4a) at Was hington physiology research. HollOSZ)· wa.~ ultimately a"~,uded
Unin:r.;ity (St. Louis), Charks '"TipMT ipwn (figure the 2000 Olympic Prize for his contributions to exerci'll'
0.4b) at the Univer.;ity o f!CM-J , and Phil Gollnick (figme physiology and health.

Exercise Ph slolo MIiestones

Physiology has always been the basis for clinical
medicine. In the same mamer, exercise physiology has
provided essential knowledge for many other areas,
such as physical education, physical fitness, physical
therapy. and health promotion. In the late 1800s and
early 1900s. physicians such as Amherst College's
Edward Hitchcock Jr. and Harvard's Dudley Sargent
studied body proportions (anthropometry) and the
effects of physical training on strength and endurance.
Although a m.mber of physical educators introduced
30 of 642
Exercise Phvsiolog Milestones
Physiology has always been the basis for clinical
medicine . In the same manner. exercise physiology has
provided essential knowledge for many other areas.
such as physical education . physical fitness, physical
!herapy, and health promotion. In the late 1800s and
early 1900s. physicians such as Amherst College's
Edward Hitchcock Jr. and Harvard's Dudley Sargent
studied body proportions (anthropometry) and the
effects of physical training on strength and endurance.
Although a number of physical educators introduced
science to the undergraduate physical education
curriculum . reter Karpovich. a nussian immigrant
who had been briefly associated with the HFL. p layed
a ma jor role in introducing physiology to physical
education . Karpovich established his own research
fac ility and taught physiology at Springfield College
(Massachusetts) from 1927 until his death in 1968.
Although he made numerous contributions to physical
education and exercise physiology research, he is best
remembered for the outstanding students he advised.
including Charles Tipton and Loring Rowell. both recipi -
ents of the American College of Sports Medicine Honor
and Citation Awards.
Another Springfield faculty member. swim coach T.K
Cureton . created an exercise physiology laboratory
al the University of Illinois in 1941 . He continued his
research and taught many of today's leaders in physical
litness and exercise physiology until his retirement in
1971 . Physical fitness programs developed by Cureton
and his students , as well as Kenneth Cooper·s 1968
book. Aerobics, established a physiological rationale
for using exercise to promote a healthy litestyle.7
An other contributor to the establishment ol exercise
physiology as an academic endeavor was Elsworth R. "Buz· Buskirk. After holding positions as chief
of the Environmental Physiology Section at the Quartermaster Research and Development Center
in Natick. Massachusetts (1954- 1957), and research physiologist al the National Institutes of Health
(1957- 1963). Buskirk moved to Pennsylvania State University, where he stayed for the remainder of
his career. At Penn State . Buz founded the lntercollege Graduate Program in Physiology (1966) and
constructed The Laboratory for Human Performance Research (1974). the nation's first freestanding
research institu1e devoted to the study of human adaptation to exercise and environmental stress.
He remained an active scholar until his death in April of 2010.
Although there was some awareness as early as the mid-1800s of a need for regular physical
activity to maintain optimal health , this idea d id not gain popular acceptanc e until the late 1960s.
Subsequent research has continued to support the importance of exercise in slowing the physical
decline associated with aging , preventing or miligating the problems assoc iated with chronic d is-
eases. and rehabi!itating injuries.

M Introduction to Exerc ise and Sport Pnysiology

 An Introduction to Exercise and Sport Physiology
31 of 642

(a) John Holloszy was the winner ot the 2000 Olympic Prize for scientific contributions in the field of
e)(ercise science. (b) Charles Tipton was a prolessor at the University of Iowa and the Uni versity of Arizona. and a mentor
to many s1udents who have become the leaders in molecular b ,ology and genomics. (c) Phil Gollnick conducted muscle
and biochemical research at Wash ington State University.

Al about tine" time that Bt·rb"!;tmm rd!llnxltictd the int.1ke \'!I , enell,'"fexpenditure) during Weston',,; attempt
to w;lik 400 mi (644 km) in !i,·e dap•. Although the ,,;tucly
needle hiop~1• pn)(:edun·. exerciM· physioh,i,,rist~ who
were m·l1 1r.1incd a.~ bioc hc1ni~L~ emerged. In Stock• reso1'·ed fl"\\· questions ;1bout mu-.ck metabo li,;m dming
holm. lkngt Salt in realizt·d the ,-.1.luc of this procedun• exrrcise. it did de1111111-~tr,1te Ihat some body protein is
for stud)i ng hum au muscle stn1c1urc and biochemistry. lost during prolong,:d hea\'y exe1-cist·. 1t
He first collalx,1-,ued \\ith Ber~lrom in lhe late l!M;()s Throughou1 tht' 20th r e nlury. :tthlete5 were used
to study die l°Hc.'Cu of die! on m11.<1ele e nd111<11Kt· ;md repealedly IO a.~sess the physiologkal capabilitie.~ of
musde nutritiou. About the ...ame time. Regboie Edger- human st~n1,,'1h and end111-,1.nce ,md to a.,n·nain char,1.c-
ton (Unh·e~ily of(~tlifomia at Los Angdes) and Phil terist ics needed for record-setting perfonnances. Some
Gollnick WL·re tL,;ing rat,; m stud~· tin· charnch·ristics of allt·mpt~ ha,·e been made to u.,;e tlw 1ecl111olo1,,'}' and
indi,idual 11111-~de fibe~ and their rt"Sl)t)IIS(_"S In uainiug. knowledgt· d eri\'ed from ext·rcise physiology to predict
5.tltin subs.cquently combined his knowledge of the pc-rfonnance, m prrscril>t· tr.tining. or m idrutify :uh-
lt•tes\\;lh exceptional potential. In most ca.-.es, hoWt.'\'er,
biopsy pnKethm· with Gollnick ·s biochemical talent~
These n·se;uclwrs wen· responsible for many early theM." applicalilllL~ of physiological te5ti ng are of little
studies 011 human muscle fiber r har,1.cleristics and tL'le more than academic intere,,;t bcc-,111q" ft,w labornlol)' or
field 1esL,; can accurntdy a.'l.~<."SS all tlu- qualitit·s requirt-<I
during c·xercisc.·. Although man~· bioclwmi~L~ ha\'e tL<;ed
exen:ise to sh1dy metal>tili~m. fl"\\' han· lmd mnrc impact to becomt· a champion.
The inlent oft hi~ M.""ction ha.~ been to prm-ide rt·ad<:"r.;
on thrcurrent direclion of human ext•rc::i"le physioln),,'}'
with an O\'ef\;ew oft he personalities and teclmologies
than Bergstrom, Saltin. Ti pion, Holl ~ ·.and Gollukk.
For mon· than 100 years, athlt·tes ha\'t• sern·d a.~ that ha,-e hdped to sh.1pe 1he field of rsercist· physinl-
subject, for study of the upper limiL~ of human t·ndur- Og)'- Naturall y, a comprehensh·e n•,if'\\' of all the sden-
ann:. Perh.1ps the li~t phyi; iologir;1I s111dit·son athletes tisL~ and n· ...ea.-ch ;as.~odatc·d ";th 1his field i, not Pos-
occurred iu 187 1. A1Lo;tin Fli111 studied one oflhe most sible in a tt·xt intc·nded ,L~ an int rod u ction HJ exerciu·
ph)'!liulu1,,')'; bm for thnst· studc·nt~ who wish to take an
Cl'kbratl'd athletesofth;u e1-... Edw;ud P;1yso11 Weston,
in-depth look ;It the hi.~wrical background in exerci«-
an endurancl' runnrr/ walkt·r. Flint 's inn·stigation
imuln:d mea.,uring \\'1·s11m ·s 1·ne1l,'Y b;1lanet· (i.e .. food ph~':\iolng)', there an· sl."\·ernl good so urn·s.

Evolution of Exercise Ph
The history of exe rcise p hysiology
has. in some ways, been d riven by
advancements in technologies adapted
from basic sciences. The early studies
of energy metabolism during exercise
were made possible by the invention of
gas-collecting equipment and chemical
analysis of oxygen and carbon dioxide.
Chemical determination of b lood
lactic acid seemed to provide some
insights regarding the aerobic and
~-~n,,.,1-, ;,. .,.,.n,,rt" nf m11c:r11li:ir ,;ir,tivitv
 3 2 of 6 42
..xercise Ph
The ~istory of exercise physiology
has. 1n some ways, been driven by
advancer_nents in technologies adapted
from basic sciences. The early studies
of energy metabolism during exercise
were made possible by the invention of
gas-collecting equipment and chemical
analysis of oxygen and carbon dioxide .
Chemical determination of blood
lactic acid seemed to provide some
insights regarding the aerobic and
anaerobic aspects of muscular activity,
but these data told us little regarding
the production and removal of this
by-product of exercise. Likewise. blood
glucose measurements taken before.
during , and after exhaustive exercise
proved to be interesting data but were of limited value for understanding the energy exchange at
the cellular level
Prior to the .1960s. t~ere were few biochemical studies on the adaptations of muscle to training.
Although the held of biochemistry can be traced to the early part of the 20th century, this special
area of chemistry was not applied to human muscle until Bergstrom and Hultman reintroduced and
popularized the needle biopsy procedure in 1966. Initially. this procedure was used to examine gly-
cogen depletion during exhaustive exercise and its resynthesis during recovery. In the early 1970s.
as noted earlier. a number of exercise physiologists used the muscle biopsy method. histological
staining . and the light microscope to determine human muscle fiber types.
Over the last 30 years, muscle physiologists have used various chemical procedures to under-
stand how muscles generate energy and adapt to training. Test tube experiments (in vitro) with
muscle biopsy samples have been used to measure muscle proteins (enzymes) and to determine
the muscle fiber 's capacity to use oxygen . Although these studies provided a snapshot of the fiber's
potential to generate energy. they often left more questions than answers . It was natural. therefore.
for the sciences of cell biology to move to an even deeper level. It was apparent that the answers
to those questions must lie within the fiber's molecular makeup.
Although not a new science. molecular biology has become a useful tool for exercise physiolo-
gists who wish to delve deeper into the cellular regulation of metabolism and adaptations to the
stress of exercise . Physiologists like Frank Booth and Ken Baldwin have dedicated their careers to
understanding the molecular regulation of muscle fiber characteristics and function and have laid
the groundwork for our current understanding of the genetic controls of muscle growth and atrophy.
The use of molecular biological techniques to study the contractile characteristics of single muscle
fibers is discussed in chapter 1.
Well before James Watson and Francis Crick unraveled the structure of DNA (1953). scientists
appreciated the importance ol genetic_s in pre?etermining _the structure and function of all living
organisms . The newest frontier in exerc ise physiology combines the study of molecular biology and
genetics. Since the early 1990s. scientists have attempted to explain how exercise causes signals
that affect the expression of genes within skeletal muscle.
In retrospect. it is apparent that since the beginning of the 20th century. the field_of exer~ise
physiology has evolved from measuring whole-body function (i:e .. oxyge~ consump_ t10~. respira-
tion. and heart rate) to molecular studies of muscle fiber genetic expression. There 1s little doubt
that exercise physiologists of the future will need to be well grounded in biochemistry, molecular
biology. and genetics.

12

An Introduction to Exerc ise and Sport Physiology

:::;:;:i,:i;~;: :!J~;;,:::~::.7' :~:7:~;:,

1 1 1
, )~:::~~::·:::,

Artilli/1.' Althoug h the ;1uthors .~ul~c•fll\'dy :1,t•lcctc.-~

·~~~~'.~'t:,
tl1<1se ~it-ntific stud ks tlwy considered w011hy of p11blt-

Women In Exercise Ph slolo

As in many areas of science. the
contr i butions of female exercise
physiologists have been stow to g_ain
recognition . In 1954, Irma Ahymmg
collaborated with her fu~ure husban~ .
P.-O . Astrand. to pubhsh a classic
study that provided a means to predict
,. · - _____ ,,, . 1.~,.., ,.,,1-,.....,,,-,im::>I hP~r!
 An Introduction to Exercise and Sport Physiology

In 1968, D.B. Dill \\TOie a c hapter, ~Hi.\tn~· or the

Roscoe.· Brown Jr., lhe finu Afiiran Americ-,m exercise
Physiology or Exerc ise,- that detailrd many or the
physiologist. coaulhore d C/11.uirn/ St11di,J ,.,, Phy.,irnl
events and M:ienti.\ts who contributc·d tn this fll"ld
Artillity.' Although the authors subjectin•ly selerted
befon· the foundin g or lhe HFL.~ In that ,1.;un e ye;ir.
those !oeien1ilic studies lhl"}' considered wonhy or publi-

Women In Exercise Ph slolo

As in many areas of science , the
contributions of female exercise
physiologists have been slow to gain
recogni tion . In 1954, Irma Rhyming
collaborated with her future husband,
P.-O. As trand , to publish a classic
study that provided a means to predict
aerobic capacity from submaximal heart
rate. 2 Although this indirect method of
assessing physical fitness has been
challenged over the years, its basic
concept is still in use today.
In the 1970s, two Swedish women.
Birgitta Essen and Karen Piehl. gained
international attention for their research
on human muscle fiber composition and
function. Essen, who collaborated with
Bengt Saltin, was instrumental in adapt-
ing microbiochemical methods to study
the small amounts of tissue obtained
with the needle biopsy procedure. Her
efforts enabled others to conduct studies
on the muscle's use of carbohydrates
and fats and to identify different muscle
fiber types. Piehl published a number
of studies that illustrated which muscle
fiber types were activated during both
aerobic and anaerobic exercise.
(a) Birgitta Essen collaborated with Bengt Saltin and Phil
In the 1970s and 1980s. a third Gollnick In publishing the earliest studies on muscle fiber
Scandinavian female physiologist. Bodil types In human muscle. (b) Karen Plehl was among the
Nielsen, daughter of Marius Nielsen , first physiologists to demonstrate that the nervous system
actively conducted studies on human selectively recl"l.llls type I (slow-twitch) and type II (fasl-
responses to environmental heat stress twilch) fibers during exercise of differing Intensities. (c)
and dehydration. Her studies even Barbara Drinkwater was among the first to conduct stud-
encompassed measurements of body ies on female athletes and to address Issues specifically
temperature during immersion in water. related to the female athlete.
Interestingly, at about the same lime an
American exercise physiologist, Barbara Drinkwater, was doing similar work at the University of
California at Santa Barbara. Her studies were often cooducted in collaboration with Steven Horvath,
D.B. Dill's son-in-law and director of UCSB's Environmental Physiology Laboratory. Drinkwater's
contributions to environmental physiology and the physiological problems confronting the female
athlete gained international recognition. In addition to their scientific contributions. the legacy of
these and other women in physiology includes the credibility they earned and the roles they played
in attracting other young women to the field of exercise physiology and medicine.

Physiology of Sport and Exercise

ca1ion. th e l·dited hook prmides an t-xce llc111 :<.ampling

or importalll exerci~· phylliological n-a rch from tht·
Research Tools:
early l 9CXk. Ergometers
In the early 1970s, Dr. SU'\'en aud Betty Hon-.1th \ Vhen phylliological respon5es to exercise are a.~sessed
(D.B. Dill"s so n-in-law and daughter) published a in a laboratol)· settin g, the panicipant'~ physical dfon
detailt·d history or th e HFL. including the laboratory must be controllt:d In pru\"ide ii mea.\urnble ext·rcise
and fidd studit,s conducted by the key !oeil·ntisL\ or that intensity. This i.~ ge ne r.ally accomplished through use
era.''Although o lhen. ha,·e written different \·en.ions or ergometen,;. An ergnntctcr (,rgo"' work; mrln-: me.t-
or exercise physiology hl\ lOl)',u" most tend to prmide ~ure) is .111 exercillt" de'!ict· that allows the intensit}' or
the authors· \iews or important !l("ientisL, and t"\·enL~. exercise to be c011troll~d (standardiz~d) and mea.,ured.
perhaps a.~ we han· done here. Finally. McArrlle, Katch,
and Katch 1h published o n e ofthe nu.st comprehcnsi\'e Treadmills
Physiology of Sport and Exercise

cation. th e edited book prmides a n exccllem 11.1.mpling

or i111port;111t exercise physiol~rical researc h from the Research Tools:
car-Ir 1900s. Ergometers
In the t·a rly 1970s, Dr. Ste\'en a nd Betty Ho n ~dth
When phyi1iological responses lo e xercise ilre a.~5t·.'l..~ed
( D.8. Dill's son-in-law and dau g hter) publis hed ;1
i11 a lahorator,· :i.cning. the participant's phy:..ical effort
detailed history or the HFL, including the lalx1rntor,·
1nust be controlled to prmidc a mea.-.urable exc1·cise
and fidd smdies conduncd hy the kt-)·scic ntisL~or that i111ensity. This i., ge ncrn.lly ,1cco111plis hed throug h lL~o:
11
c rd. Although otheni. ha\·c written differen t ven.ions or ergon1eteni.. ,\n ergo meter (ngo ,. work; m, t,r• mea-
or exercise ph~-siol~•y histor,·, ~."' nm,.;t tend to prmide
sure) is an exercise dei.icc th;n allows the intensity or
the authoni.· \iews or i111portant scientisL~ a nd n·cnl5. cxcrci.'IC to be cont rolled (standardized) .u1d measured.
perl1;1ps a.~ we ha\'c done he re. finally, McArdle. Katd1.
and Kat ch l b publi.~hed o ne or tlw most compre hcruh-c Treadmills
rc\iews or die evolution or exercise php;iolob')". ll1cir
Treadmills are the crgom e ters or c hoice for most
description or th e early anatomi.~L,. physiolob,j,.;L,. and
1·escarc h cn and clinicians, particularly in the United
excrci,c ph}-siologi,.;ts clc-drly illmtrntcs th e complexity
States. \\'ith these dci.ices. a motor d1i\'cs a large belt
a nd diveni.ity or thi,.; field or :~cicncc.
on which a .~ubject c.111 either w.1lk or nm: thus the~
Now that we undentand the hi.~torical basis for the
ergometcn are often called motor-d ri\'ell treadmills
discipline of' exerci.'IC ph}-siology. from whic h .~port phy5-
(sec figure 0 .5) . Belt length and width muM accom•
io l~')' emerged. we can explore some basic principles
or. and tool~ utilized in. exercise and ,.;port ph}'liiology.

Research: The Founda-

tion for Understanding
Exercise and sport scientisL~ ac tiYclyengagc in research
m better understand the m t."Chanisn1s that 1·cgulatc thc
body's physiological rt.-:i;ponst.'!li to acll!c bouL~or exercise
a.~ well a.~ iL~ ada pta ti ons to tr.t ining and det r.tining.
MoM o r th is research is conducte d at major rese;1rch
tml\·crsities. 111cdical cente rs, and specialii:cd instinnes
using .~t;uulardi1.cd rcscarrh appn>.1chcs and .'IC:lect tool~
or th e exercise physiologist.

Research Settings
Research ca n be conducted either in the labora tor,· or
in the field . Labor.uory tcsl5 are u.~ually more accura1c
because m ore specialii:ed and sophi~ticatcd equipment
can be used a nd conditio n ll can be carefully co111rolled.
A~ an example, th e direc t l,iborato l'}' measurement or
maxima l ox,·gcn uptake (\1O:i...,, ) is considered 1hc
most ;1cr11r..11e estimate or cardio n·spiratory endurance
rnpacit}', H owc\·cr, !I0llle field test-.. such _as the 1.5 mi
(2.4 km ) nm. arc al~, used to estimate VO....,.. llu.'SC
field tesL~. whkh mca.~urc the time it ta kes to nm ii .'ICI
di.,tance or the di.,tance th a t can be cm-ered in a fixed
time, arc not totally accurate; but they prmidc a reason-
able ei;timate or'~Ot..,· arc inex pe11.,l\·e to conduct, and
allow many pcopk to be tested in a short time. Field
tcsL~ can be conducted in the workplace, o n a nmning
trdck or iu a S\\imming J)(Xll, o r during a1hle ti c compe-
titions. To llleil.\Ure \'O...,.,. direct ly an d accur.udy. one
wo uld need to go to a uni\'cni.ity or clinical lalxir.ttol'}',

AA Introduction to Exercise and Sport Physiology

9,.1;1g a~ pedal r,uc dt·rrea.~s. and dccrca.~cs a.~ pedal rntc

inc reoL'ICS, IO prmide a cnnsmnt power oulput.
Treadmills generally produce higher peak values Similar lo lrcadmill~. cycle c rgomcters nfTcr ,'M>llle
than other ergometers for almost all assessed ad,·antages and disad\'a111ai,;:L·.~ compared to other
physiological variables , such as heart rate. ventila• ergom c tcni.. Exe n::i.'IC intensi ty on a q-de ergometer
lion, and oxygen uptake. docs not d c pc- nd on the subject 's body weigh!. Titi~
is important when one l~ in\'estig"dtini,;: phylliological
responses 1oa s1andard rdlcorwork (power output~ . A~
mudate th e indi\'i dua\'s bod)· size and snide length. a n example, irso mcun e ln~I 5 kg (I I lb). data d cm·ed
for example , it i, uc-arly impossible lo lc.'SI clile alhleles fro m trc-ddmill testing could not be compared \\ith data
 An lntroductio, to Exercise and Sport Physiology

91.141g as pedal rnte decrea.o;es, and decrea.'IC'!l as pedal r.tte

increases, to prmide a constant puwer output.
Treadmills generally produce higher peak values Similar to treadmill~. g.dc: ergomc:ten offer so111e
than other ergometers for almost an assessed adrdntagc:s and di.'lad\·antagc:s compared to other
physiological variables, such as heart rate, ventila- e rgometc:n.. Exc:rci.~ intc:rl.'lil}· on a g.-cle ergon1eter
tion, and oxygen uptake. doc.'I not depend on the .'l ubject'.'I bod)· weight. Thi~
i.'I impon:mt when one i~ irl\"n tig-.tting physiologica l
rc.'lponsc.'I to a .'ltandard rdte of work (power output) . A~
modate I.he indfridual'.'1 body size a nd stride length . an example, if someone: lo.'11 5 kg (I I lb), data dc:ri\·ed
For cxtunple. ii i.'I nearly impossible to tcsl elite athletes from treadmill tt'.'lting could not be cump:ired Y.ith data
on treadmill~ 1hat are too short. or obcsc .'l ubject~ on obtained before the weight los., l>c:cau.'ll' ph)'lliological
treadmills that arc too narrow or not .'lturdr enough. rcspull.'ICS to a lit't speed and grade o n the treadmill
Treadmills offer a number of :1d\'a ntagcs. Walking
i\ a naturdl acthity for a lmost C\'er,·one, so indi\id11:1l1
normally adju~t tu the skill required for y.-.tlking o n a
treadmill Y.ithin a fn,· minutes. Also, 1no:st people can
ac hie\·e their peak phy.'liologica\ \-.tlues o n the tread-
mill, ahhough i'iOlllt' a thle te.\ (e.g.• compctith·e cyclists)
ac hie\·c highe r \-.tlu" on ergomctcn that more d ffl.C I}'
111:ttch their mode of tr.tining or competilion.
Treadmills do ha\"e !!O ll1t' di~ad\-.tntagcs. TIit'}' are
genernlly more expcn.'l in~ than simpler c rgmnele D.
like the cycle ergomcten disnL~sed next. Tht'}' arc
al= bulky, require electrical power, and arc not \ 't''1'
portable. Accurate mca~ureme nt of blood prcs.~ure
during treadmill cxercio;e can be dillicult because both
the noilit' a.'l.'W>ciated with no m1al treadmill oper.ttion
a nd subject mm·emenl can make hearing thro ugh .i
stethOM:ope difficult.

Cycle E,gometera
For ma ny yc:m1. the cycle eTg0mc1cr \\-.t.'I the primary
testing dC\icc: in 11.'IC. and it is s1ill used extell.'li\·cly in
bo th research .ind clinical senings. Cycle c:rgomc:leD
can be d n igned to a llow subjc.."t'. l~ to pedal c:ithl'r in the
nonnal upright pusition (.'lee figure 0.6) or in reclining
or .'ll'mireclini ng positions.
Cyde c:rgometeD in a rcs,carch setting generally 1uc:
eithe r mechanical friclion or electric.ii re.'li.'ltancc. \\'ith
mechanical friction dt'\icn. a he h e ncompa.~sing a fly-
wheel i, lightened or loo:o;c:ned to adj1L~l 1he rc:.'l istance
ag-.tinst which the g.dist pcda l.'1. The puwc r mupm
depends on the combin.itio n of the resistance a nd !he
pedaling rate-lhe f:L~lc:r o ne pedal~. the greater the
puwer o utput. To maintain the !la lllt' power output
throughout the test. one mlL\I maintain the ~ me pc:daJ.-
ing r.tte.so pedaling r.tte m1L,t be constantly monilored.
With electrically br.tked g.de ergometeD. the re.'l i~
ta nce to pedaling i., prmidc:d by a n elec trical conduc-
tor !hat mm-n through a magnetic or electrmnagnc:tic
fidd . 1l1c: .'ltrength of the magnetic field dete rmines
the resistance to pedaling. llle.'ll' ergometen can be
contmlled so that the resimnce increa.o;es automatically FIGURE 0 .6

Physiology of Sport and Exercise

\-ar,•with lx~yweight. Afte r the weight loss, the rnte or water pa.~t the .n,imme r. who attempts to maintain
wo rk at the ~ me speed and grade wo uld be Jes.~ than body position in the flume . The pump circula1ion can
before. With 1hc: C)'Cle ergome ter. weight Jos.,; docs not be increased or dc:crea.o;ed to ,~.t11' the..· speed a t which
ha\'t' a.~ great an effect on phyl'i iologica l response to a the !lwimmer n11L~I swim . The sYoinuning flwne. which
standardized power output. llnL~. walking or nmning nnfo rtunatc:ly i~ n·r,· expen.~i\e, ha.~ at least paniall}'
i~ often referred to as wc:ight-d.ependc:nt exerci~c:. while resoh·c:d lhe problems with tethered .'lwimming and
cycling i.~ weight independe nt. has created new opporttmitin to im·estigale the spurt
of .'IY.imming.
When one i~ choosing an ergome ter. the concept of
G;)z.141c specificity is panicularly impunant Y.ith highl y tr.tined
Cycle ergometers are the most appropriate devices athlc:tC.'I. The more specific the ergomete r is 10 the
for evaluating chcnges in submaximal physiological ac111al pattt·m or mm~ment u'lt'd by the athlete: in his or
function before and after training in people whose her spun , the more meaningful Y.ill be the test r"ult,.
weights have changed. Unlike the situation with
treadm~II exercise, , c~~e ergor:neter intensity is Research Designs
 Physiology ot Sport and Exercise

,;:u-y \\ith body weight. After the weight lo~s. the r.1te or water p•L~t the s\\immcr. who attemplli to m,1intain
wurk ;u th e 11;1mt· spt:t·d ;md 1,;r;1ck would ht.· It•,;_~ 1h;111 body po11itio11 in the flume . The pump circulation can
before. \\'ith the C')Tlc ergomeler. weight loM doc.~ 1101 bc im-re;t'lf.•d or clt."'Crca.-.ed lO \'al"}' the speed at which
ha,·e a.~ b'Teat an effect on physiological response to a the swinnuer must swim. The S\\imming llume, which
s1.1 m\;u-di1.cd power nntpul. Thus. " -diking or nmning unfortunately i, H'.ry expensive. has at lea.~t 1,:1r1ialh
is often referred to a~ weight-dcpe ncknt t·xerci,c, while rcml\"ed the problems "'ith tethered swimming and
cycling is weight independent. h;L, c re;ued new opportunities to im·estig-Jte the spon
of swimming.

@m.m When one i.~ choo~iug ;m eq;ometcr, the concept or

specificit~· is particularly impor1;1111 "ith highly trained
;ithletcs. T h e more specific the ergonwlf.'r is to the
Cycle erg001eters are the most appropriate devices
;1rtu;1I p;nte m or mm-cment used by the ;uhlett· in his or
for evaluating changes in submaximal physiological
her Sjk)rt, the more meaningful "ill be the test re.~uh~.
function before and alter training in people whose
weights have changed. Unlike the situation with
treadmill exercise, cycle ergometer intensity is Research Designs
largely independent of body weight. In exercise physiolob'Y rcM•;u-rh, there arc two basic
1nx·.~ or research design: cros.v-S<'rtion:1\ ;md lonbritu-
Cycle eq;omcten; also ha,·e di11;1th-;u11;1ges. lhhc sub- din.il . \\'ith a cross-sectional research dt.~ ign , a noss
ject dOl•s not regularly eng-Jgc in that form or cxcrci'IC, sec lion of the population or i111crest (that is, :1 rcpre-
the leg musclt•s "·ill likely fatib'lle early in the exercise sc ntati\·e s;unp\e) i~ tested atone specific lime, ;md. the
boll!. This may pre,·cnt a subject from attaining a 1111e differences bc.-1w1:t:n subgroups from that ll.1111plc arc
maximal intensity. When cxerci~c is limited in this compared. With a lo n!,ritudin al r esearch design , the
way, re.~pmL'ICS arc often referre d lO <L, peak exercise !\;\Ille researc h subjecl~ ,1re retested periodica lly ;1fter
intensity r,uher than maximal t·xcrrise intensity. This initial testing to measure ch;mgt'll o\"er time in ,-ari.1h\1~
limitation IIM}" Ix: attributable to ilK·;1l l1•g fa 1ib'lH~. bloocl of interest.
pooling in the kg.~ (kss hlood returns to 1lu· h,·;u1).or The differences between the~ two approaches ;in•
th1· use or a smalkr tmL,de m,L.,_, during t~·cling than best unde!"ltocxl. through :111 t•x;unplt". The objcrtin· of
during treadmill cxercis1·. Tr;1incd cyclisL~. howe,·cr, a rcM·arch study is lodctcnuint· w)l!"thera regular pro-
tend to ;irhin·e th eir highest 1:H;·;1k \.1lucs on the cyde gram of dis1.111re running increas1·s the concentrntion
ergometer. or cardiopn ,tt·nin, high-density lipop1Utein cholesterol
( HDL-C) in the blcMxl.. High--density lipoprutcin cho-
Other Ergometers kstcrnl is the dt·sirJb le fonn or cholesterol: increased
concentrJtions an· ;L,socia tcd. \\ith reduced risk for
Other ergomcters allow ;uhletc:-1 who nunpc tc in spe-
he:111 di'ICase. Using the cross-sectional approach. one
cific -~JklllS III" c\"c nts to !:H;• tt·~ted in a 111:mm·r that more
could . for exam pk, tt:st a large number or people who
closcl~· approximates their 11;:1ining and competition.
fall into th e follo\\ing categories:
F'ur exampk, ;111 am1 crgomt•tcr may be 11.st·d to test
:uhlctes or non;nhletcs who \lilt: primarily thdr ;inns A J,rfoup or subjects who do no 11-aining (tlw
and shoulde rs in physical acti,ity. A nn crgomell"}" ha., control group)
aiilo been used extcnsi,·dy lo tesl and trnin athlc.-tcs
Agrouporsubj,·cL~whonm 24 km (15 mi) per
par.1l rlCd below arm k\"el. The rowing ergumell"r "~,1.,
week
dc,isc.•d 10 tot cumpc.-tith-e rowen..
Valuable research d;1ta ha\"e bt·t·11 obtained hy Agrnupor subjt•ct, whonm 48 km (30 mi) pc·1
instrumenting fflimmt·u ;md moni101ing them during week
s\\imming in ;1 poul. Ho\\·t·,·t·r. the prohknL~ <L,sociatt·d A group or subjec t~ whu nm 72 km (45 mi) per
1,ith 111n1s ;me\ constant mon·ment lcd lo the use or two week
dt·,·ices-tt·llwred swimming and sv.imming flumt·s. ,\ group or subjects who nm 96 km (6{1 mi) pe r
In tethered S\\imming. the swimmer is :m;1chcd to a wt:ck
lrnrncss connected to a ropt·. ;1 series of pullc~-s. and
comHerbalancing weights aucl 11111st S\\im ;1gainsl the One would then rump.ire the results from ;1II the
pull or the app.ir.1111s tu mainmin a const;mt pasition group!i. l>:L~ing une's conclui1io1L~ on how much nm-
in the pool. A swimming flnnu: allows S\\i111111en to 11ing1,;:•s donc. Usini,: this approach, cxcrci.st· .'ICientisL~
more closelr simul;uc their natural 1'"imm i11g st.-n kes. fou nd that weekly running results in dcmtt•d HUL-C
The S\\imming flume operntcs hy pumps 1h,11 ci rculate le\"ds. suggesting ;i positiYc h c:1hh benefit related to

16

AA Introduc tion to Exercise and Sport PhysiOIOgy

80 - 60 -

II mI
ss - -
70 -
so -
i
t oo -
as so - I·
I
«> - ~

I (I I
i 35 -
g .., - ., - I 11
 An Introduction to Exercise and Spor1 Physiology

60

55 -

24 48 72 96
Dial•""'- The relatioo between ~nths ol di~•
1ance-n.11ning training and averag_e high-density li~r01e1n
The relat~ ll:9tween .distance run cholesterol (HOL-C ) concentrattons in an expenmental
per week and average high-d84:'s1ty l,poprotem c~sterol group (20 subjects, distance t_ra!ning) and a sedent_ a ry
(HDL.C) concentratlOl'ls across frve groups: nontramrog con• (20 subjects) con trol group. This 1llustra1es a longitudinal
trol(Okm/Week). 24km/Week. 4 8 ~. 72krn/'Neek. ~d study design.
96 km,lweek. This illustrates a cross•sectlOl181 study design.

nmning di~ta n ce. Funhennore, a ~ illu.ur.tted in fi gure

O. 7. there l\·.t.~ a d ose-response relation between d1eM""
,-ariable.'1--{he higher the wdc1."c.'w of c.'Xerci!IC tr.tining,
£n-t4W
Longitudinal research studies are the most accu-
rate for studying changes in physiological variables
the high e r the rei1 ultin g cou cenm1tion of HDL..C. h
ii1 important to reme111ber. howe\'c.'r, that \\it h a nos.~ over time. However, it is not always feasible to use
sectio n al dl"!lign. tlu:se arc diffrre nt grm 1ps uf n mneB.
a longitudinal design. and valuable information can
not th e !;;ll!le nmnL·l"ll at diffrre nt tr.tinin g vo lumeli.
be derived from cross-sectional studies.
Uliing the lo ngitudinal a pproach to te.~t die ~une
qm:5tiun, one <ould d c·sign a i1111dy in wh ich untrnined A lonb,itudi nal reM::irch design i., usually best suited
people ...~mid be recruited 10 panicip;u e in a 12-month m 5tlld)ing rha n ge i1 in ,,1riables o,·cr time . Too m a ny
di~t.11Ke-n11111ing prob.-r.1111. O ne could, for ex;unple. f:tcto r5 th:ll m;1y taint rcsuh5 can inOuen r e cross-
recmit 40 people .,.;IJing m begin numing ;md then M:ctio n al design ~. For example. genetic factol"ll might
r,rn domly a.~sign 20 IO a tr.tinin g b'Toup a nd th e rema in- inter.tr! :w th.1t those who n m long di~tances :ire a lso
ing 20 1n a ro ntrol group. Both gro ups would be fol• th ose who h owe high l-lD L-C le,·els. Also, differe nt
lowed for 12 mo nths. Blood ~m1plei1 would be IL'!ited 1x1pulatiom• migln follow different dicL,; but in a Ion•
at the beginning of th e s1udy and then a t threc-.month bri lltdinal study. diet and mhe r ,-ariables GIil be more
inter,.il~. conclmling a t 12 mou th.~ whe n the pn>i,,--r.1111 ea.~i ly ro ntro lll°d. Howc ,·er. longi111dinal studi es are
l°nded . With this design. both th e nmning g roup :md 1imc-cunsuming, are ex pc11.~l\·e to conduct, a nd are
the cont ro l group wou ld be followed m-rr th e entire not a lw:.:1pi JX>S.~iblc; ;md crns.....seclion:11 .~tudi Cli prmicle
1x:ri,xl of the i1tmly, and changes in the ir 1-IDL--C le,-rls some in.~ight into th e."l' que.5tionli.
could be d e tennim:d a<ros., each pe riod. Al:tnal studi~
h:we bee n conducted IL~ n g this long itudina l d esib'll
IO examine c hanges in HOL--C \\ith trnin iug. but th eir Research Controls
resu ll'i h a,-r not l,c:eu a.~ clear ;L~ the res ults of lhe \\'hen we cnnduc1 rellCarr h . it is impo rl ant to be as
c n )!l~r tional studies. Sec figure 0.8 a., an rxam plc. careful as pt>S.~ible in d esigning th l° 5tudy ;md collc cl-
Note 1h;11 in this figure . in conu.L, t lo ligure 0.7. 1here ing th t" d.1ta. Wt· ~1w fro m fi gure 0.8 th:11 chan ges in a
1
~~~•;~~i:1;::;~~:::.c;:,~~:c:; g~,~~i:t;i1:~
1
~:i~~,\•:.1~~~:;~:: ,-ariable O\'t•r timt" result in g from an inten·cntion such
a.~ t·xc·r< isc can be ,·c •"}' small. Yt"t, e\"en small c h:mge.5
wilh only minor f1uc tualions in thei r HDL,C from o ne in a ,-,1.riable 511ch :L\ I-ID L-C c,111 m ean a sul)!lt.Ullial
three-monlh pt·riod to the next. reduction in risk for hean discasc. Rccogni:ting thi\.

Physiology of Sport and Exercise

scit:ntisl.'i design sm d iell ai med at p rmiding re.~ultli th:11 It is equally important to contro l d ;ua co\lc c1ion.
are both accurn te and reprod ucible . This rt:quircs that TI1e equipmcnl mu51 be calibrated so th e reM:arc h cr
sturlil"5 be carefully cont ro lled. knows that th t: \.1 lues gem:rntl°d by a given piece of
Research control~are .1pplied at m rio us levds. Stan- equi pment ;ire accurate. a nd th e procedt1J"c.'S IL~ed in
ing \\ith the dt"5ign of the rCliearch pn!.iccl. the scientist collecting d a ta must be sta ndardized. For example.
mnsl detennine h ow to coutrol fon-ariation in the sub. when 11.~ing a sc.1lc to mca.'!ire the weight of subject,,
jccL~ used in the study. The scientist 1111L~t d c te nnine ifit re:1-earcht•r.5 UcL·d to calibrJte that .5(":llc by using a set
is im1x1n:1111 to control for th e sul!.it,cl.'i' sex. age, or lx1dy of calibrnted we ight\ (e .g.. JO kg. 20 kg, :-\0 kg. and
si1x. To use age a.~ an example. fo r cc nain ,,iri;1bles. tht: 40 kg) th at have been me;L~urc·d o n ;i precision sc;1le.
respon.'l,l' ltl an ext:rcise trJi ning progrnm might be dif- Thn.e weighL\ art" p laced o n th e wdghing scale to b e
ferent fo r a child or a n aged person compared \\ith a m e d in the study. indl\idnally and in combination . .1l
young o r middle-aged adult. L~ it irnpona nt to cont ro l least o nce a WL°e k to provide cen a inty that th e .-.(':lie is
for the 511l!.ic·rf5 limoking or dietary statm? Con5ider- m ea.\t1ring the weights ;iccurntcly. A\ a n othe r example,
able thnught and dii;c11,1sion are needed to make sure electronic :malrzcr.i used to measure respirntory ga1;.es
th at subject~ u~ed in a siudr an· appropria lc for the n c,:d to be ealihra1ed frcq uL·111lr wi th ga....cli of known
scicntii;L, d esign studies aimed al p ro\iding r" ulls that It i~ equ ally important to cout ro l d o1ta collection.
are both ac n u-,u e and reproducible. l11is requires that The e quipme nt 11111s t be calib rnted so th e re:o;earc her
studies be ra rd ully controlled. knows tha t th e ,.1h1e5 gene r.t ted by a gi\·en picce o f
Research control, .1re a pplie d at mrio us lrrd s. St.m- equipme nt a re accu ra te , a nd the procedures used in
ing with the design o f th e rt:5Ca n:h project. the scientis t collecting data mus t be s1ando1rdi zed. For e xa mple,
mu.~t d e tennine how lo conlro l for,-aria tion in the sul>- whe n us ing a scale to me asure th e we ig ht o f su bjects.
j ects used in the study. ll1e scienti,t 11111.,1 d e te nnine ifit rescarc heu need to calibrnte that scale by 1t.,ing a set
is impo rta lll to co ntro l for the subjt,--cL,' sex, .1ge , o r body o f calibrn ted weights (e.g., IO kg. 20 kg , 30 kg, a nd
size. To n~e age a., an example. for ce rta in \'ari able5, the 40 kg ) th at h a,·c been m ea~ured o n a p recisio n Kille.
re sponse to an c xcrci!\e training prog rnm mig lu be dif- These wcighL, .1re p lace d o n the weighin g scale to be
fe rent for a child o r .1n aged person co mpa red v.i th a used in the study, indl\ich1.1lly a nd in combination, ;11
young o r middk--..aged aduh. h it important to control lea~t o nce a week to provid e cc rt.1inty that the JIC'.ale is
for the subject"s smokin g or dieta11· status? Comide r- me ;1.~uring the weightsaccu rn tdy. A~ .111o the r example,
able thought and disc us.,io n are needed to make sure elec tro nic analyzeni used to measure res pir.1.to l)' g-.i.~es
1hat th e ~ubjccb , ucd in a sm d>· a rc appro priate for the need to be calib r,. tc d freque ntly with g-.i.,c~ of knowu
specific research question being :1.~kt•d . concent rntio n to ensu re th e accuracy of th ese a n alp cll.
For ,1Jmost all i;tudie-s. it is critic.1I to h;wc a control Finally. it is import.Int to kn ow tha t all test results
gro up. In the lo ngimdinal rMearch des ig n fo r 1he cl10- arc reproducible. In th e example illustrated in fi gure
lesterol study described earlier, the control group aCL~ a~ 0 .8, the HDL-C of ,111 indhidua l ii; m e asure d e\<ery
a compari.~on group m make certain that any d1.1nge s three m o nths. If that pc.· rson i~ tested fi\"e da >'3 in a
observed in the n mning group are attributable M>lcl y ruw before he or she starts the tr.1 ining p r°brr.1.m , o ne
to th e tr.tinin g p rogr,un a nd nut to any other fac tors, wo uld expect the HDl.rC resul ts to be ., imilar acros., all
suc h a.~ the titnt."" of 1he year or aging o f the subjects fin• d a)'3, pr O\iding die t. e xercise, slee p, and time o f
during the cou rse of the study. Experimental d esigns d .1y for testing remained the same. In fig ure 0.8, th e
o ften employ a p lacebo group. l11u~. in a i;tudy in which ,-.1.Jucs for th e contro l group across 12 m o ntl1.~ \-;trie d
a subject mig ht e xpec t to h o1,·e a benefit from the pro- from about 44 10 45 mg / dL, whereas the exerci,;c group
poi;ed interventio n. such a, th e U JIC o f a specific food inc rc-.tsc.·d from 45 to 47 mg/ dL O\"er fn·e consccutil·e
o r dmg, a scie ntisl mig ht de cide to lt.<;C three groups d a}'3. the 111e;1.~u rc m c 11L~ should not \'a l)" by m ore lhan
of i;ubjec ts: an interventio n gro up 1ho1t recei,·es the I mg/ dL for a ny o ne person if the research e r is going
actual food or dn1g. a place bo gro up 1hat rece in:s an to pick up this sm,1\1 c h.1n ge m·er time. To control for
inert i;ubsta.:nce th at looks exactly like 1he acma l food or re p roducibility of resu!L,, !l(:ienti\u gen e rall r take se\~
drug, and a contro l gro up th al recci\"es nothing. (The ernl m easurcme n L,, ll0111Ctim e5 o n different da>'3· and
la,t group o ften sen <es a, a Mtime co111rol,~ accounting then a,·ernge the resulL~. before, durin g, a nd at the end
for nonexperimcn ta lly induced ch o1 ngc,,; th;u may occur ufan inten·e ntion.
o\"erthe coune of the study period .) If th e ime n -cntio n
a nd placebo grou ps impro,·e the ir perfonn ,1 nce Ill the Confounding Factors
same le\"cl and the cont ro l group d oci; not impro\·e
performance. then the imprn\·cmc nt i., likely the result in Exercise Rasaareh
o f the ~place bo effect.- o r 1he cxpec1a1io n th a t the Many fac1ors can alte r the bo d y'., .1cut c response to a
substance will impro\"e per form a nce . If the imen·en- bout of exercise. For exa mple, c ll\i roume n ta l condi-
tion group impru,·es pc.·rfonnancc and th e placebo and tions su ch as the te mpe ra ture a n d humidity o f th ~
control gro ups d o n o t, th en we ca n conclude that the laboratol)' a nd the am o unt o f light and noi..e in the
inten·enlion d ocs impro\'C perfor ma nce. test area GUI m arkedly .1ffe<" t physiological rn ponses,
One other v.-.1.y of n mt rolling for 1he placebo effect both a t rest an d during exercii;c. [\·en th e tim ing,
is to conduct a i;111dy 1ha1 u.~ a crosso,·e r design. In volume. and contt:111 of th e la.,1 meal a nd the quantity
th i., ca.~. c:1eh group unde rgoes both treatment and and qua lity of ~lct:p the nigh1 before mus t be care fully
cont rol lria ls il l differe nt times. 1-"o r example, o n e group con trolled in reJ1C.1rch ~tu d ie1.
is ad ministe red the in1e n·ention for the first half of the To ill ustrn tt: th i~. 1.1hlc 0 .1 i;hm•'3 h uw ,.1.ryi ng em1-
i;tudy (e.g., 6 momhs o f a 12-month study) and sen -cs as ro nme n ml and beh o1,i o r,d facm ni can alte r heart rate
a co111rol durin g the la~t half of th e study. The second al rn t o1 nd durin g n uming on a treadmill at 14 km / h
group sen-cs as a control during the fi rst half of the (9 m ph ). T he subj ec t "i; h eart rate response durin g
study a nd recein:s the inten·entio n during th e :1econd cxerci~ di ffe red by 25 beats / min when th e air te m-
half. In some cases. a placebo can be IL~ in the co ntrol per.1.nire wa, increa.o;ed fro m 21 °C (70 °F) to 35 °C
phase o f the studr. 01apte r 16, MErgogen ic Aid, .1nd (95 °F) . Most physio logical mriablcs that are n o rmally
Sport. - p rovides furthe r disc11J1.~ io 11 o f plo1cebo grou ps. meas ure d during exercise are simila rly inUue n ced by

An Introd uction to Exercise and Spor1 Physiology

rm Heart Rate Responw to Running Differ With Variations In Environmental

and Behavioral COndltions
HEART RATE {BEATS/ MIN)

Environmental and behavioral factors Rest Exercise

1:
TEMPERATURE (ICM HUMIDITY)

21 °c(10 °F) 165

35 °C (95 °F) 100
HUMIDrTY (21 '"C)

'°" 60 165
39 of 642
An Introduction to Exercise and Sport Physiology

H•S:14:l•il Heart Rate Responses to Running Differ With VartaUons In Environmental

and Behavk>nl Conditions

HEART RA.Tl! ( BEATS/II IN)

Environmental end beha11loral factors Rest Exercise

1: -
RMPERATURE (~ HUMIDITY)
21 °C(70 °F)
165
35 °C (95 °F) 190

1:
HUMIDITY (21-C}
50% 165
90% 175

~I:
NOISI! LEVEL (21 -C, ICM HUIIIDnY)
Low

=
165
H~h 165
FOOD INTAKE (21 -C, ICM HUMIDITY)
SmaMmeal 3 h bef()(e eK&rcising 60 165
large meal 30 min before exercising 70 175
SLEEP (21 '"C, lfM HUMIDITY)
8hormore
6horless I: 165
175

en,iron mental nuctuation.'I, \'l'hcthcrone i,; comparin g ,'U'lal.ion rckr:s to fluctuations that occur during a 24 h
a pc:rso 11'.o1 exerci.'lc rc:.o1u \L,; from one rla )· to anot h er or day. Table 0.2 illuMr,Ue!I the diunial \~.i,riation in hr:an
com paring the rt:llpons<.'!I of two different .o111bjccu, all of r.ur: at rest. during \~.i,riom IC\'d!I of e xerci'\C, and during
th c::w: factor:s must be controlled il.'I carcfull)· a.,; passible. recm·t,ry. Bod y tcmper.i.ture sJIO,...'ll !limilar flucl\lations
Ph y.o1iologk,d reliponscli, both at reli t a n d during througho111 lhe day. A, seen in table 0.2, t~ ting th e
excrci~. al'IO \'ill)' d1roughuut the day. llu: tcnn diurnaJ !Ill.Ille per5em in th e morniug on one day a nd in th e

0,1:Jij.J.J An Example of Dlumal Variations In Heart Rate at Rest and During Exerdse
TIIIEOFDAY

Cond1t1on 2am 6am 10am 2pm 6pm 10pm

HEART RATE (BEATS/MIN)

Resting
Light eKercise
65

100
69

103
73

109
"
109
72

105
69

104
Modera1e eKerdse 130 131 138 139 135 135
Maximal exercise 179 179 183 184 181 181

Recovery. 3min 118 122 129 128 128 125

Data komT. Re~ly andG .A . 8rooks( 1990).·Se1ective pe<srstenc:eolcltc.ad'81'1rt,yll'ml;w, J)hvsiological •esponse,110e•e«:1Sft.' ~ y~ ,
7. W-67

Physiology ol Sport and Exercise

aften10011 o n the next can and ,...; JI produce different ,-ariables. Take, for instance, one of the comenuone
resu lts. Test times mlL,t be sta ndard ized to control for meas ure ments in exe rcise p hp iology, an indi,id ual's
this diurnal effect. maximal oxygen uptake or maximal aerobic capac-
Al le-.i.st one other physiologica1 cycle m ust also be ity, abbre\-ia ted \o _,. Thi~ important physio logical
con., ide red. ll1 r: nonnal 28-day menstntal cycle ofte n measure ment is th e maximal ,ulume of oxygen that
ill\'oh·es consider.i.ble ,-aria1ions in an individual can uti lize d u ring e xh a usti\·e aerobic
exerci'W:', and can be mea.,ured in liters per minute or
body weight, L/ min . Howe1,er, be(:ause a large person can utili:r.e
total body water and b lood ,·o lume, more oxygen yet not be more aeTobically fit, we ofte n
body tempernture, standardize this \alue to body weigh t in kilog rnms. that
m etabolic rnte, aud i~. milliliters peT k.ilogrnm pr:T minute. Now the nota-
tio n becomes a bit more comp lex and potentially more
h eart r.itr: and stroke n,lume (lhe amount o f conf1L,ing. We could write th e units as m l/kg/min. but
b lood lea\-ing the heart ,...;th each con trnclion). what is being dh-ided by wha t in thi, notation? Reca ll
that L/ min can al'l-0 be written a.~ L • min-', j1L, t a., the
Exe rci5e scien tislS mus t control for m enstrnal C}'cle
40 of 642
cise

,~uiablcs. Take, for instance, one of th e corncn;tonc

aft1·n10011 on the next can and will produce differenl
mcasun•mcnl~ in exercise.· physiology. an indi,idual 's
Tt'SUIL~. TrM time11 musl be standardi zed to contn>I for
maximal oxygen uptake o r maximal aerobic capac-
this diurnal effect.
i1y, abbrc,iated \'O""'-". Th~~ important physiological
,\t leasl one other physiological cr cle must also be
measurement is th c maximal \'olume of oxygen th;ll
considt."n:d. l11e nonual 28-day m c nstrn;1\ cycle often
an indhidu al can utilize during cxhaustil-e aerobic
inn1ln:s nmsidc rablc ,,uiations in
exercise, and can be: measured in liters per minute or
bodywdght. L/ min. Hffll1.•,·cr. bc.-ca1L'lt" a large pcn;on GIil utilize
total body wah·r and blood vol ume, more oxygen yet not be more aerobically fit. we ofte n
s1andardize this ,.i.lue tu body weight in kilogr.uus, that
body te111per.1tun~. is, millili1cn; per kil~....-.am per minute. Now the nota-
metabolic rnt1•, and tion bc.·comes a bit more complex and potentially more
heart r.ne and stroke volume (l he amount of confusing. We could 1'1-rite the unil~ as ml / kg/ min, bill
h\ood lea,in g die heart ,-,i1h each conlrnction) . what is being di,idcd by what in this notation? Rernll
1ha1 L/ min can also bc 1'1-rittcn a.~ L · miu-',just a.'i the
Exercise sciemists must coutrol for nu.•nstnt.11 cycle fr.action 1/ 4 = I• 4- 1• Toa,uiderron;and ambi1,•11ity. in
phase or the 11.<1e of oral contr,Kep1in:s (whic h similarly excrci.'lt" physiology we lL'le the expunenl notation any
;1111:r h o rmonal stalus), or both. when testing womt·n. time more than two ,-.ariablcs arc in\"olved. ll1c rcfore,
\\'hen older women are be ing 1ested, testing stralc..._ millili1ers per kil~'fam per minute is written a.'i ml ·
gies must take into account menopause and hrnmone kg-' • min- 1 r.ather than ml/kg/ min.
replacc111ent 1hernpies.
Reading and Interpreting
(Blz-1.;up Tables and Graphs
Conditions under which research participants are This book cout.ains references to .~ pccilk research stud-
monitored, at rest and during exercise. must be ies that han· had a major impact o n our unden;tanding
carefully controlled. Environmental factors, such of exercise ;u1d spurt phy~iulogy. Once scientists com-
as temperature. humidity, altitude. and noise, can plctt· a resc;1rch prc~cct, they submit the resull~ oftlwir
affect the magnitude of response of all basic physi- rcscarch to one of the man y research journals in sport
ological systems. as can behavioral factors such as and excrci.-.e physiology. Some of the more widely used
eating patterns and sleep. Likewise, physiological research journals appt:'.IT on the IL'it of selected readings
measurements must be well controlled for diurnal and rcfcrcnccs at the back ofth~~ book, a.~ wdl as in th c
and menstrual cycle variations. study guide on th e website www.Hu111anKinetics.cm11
/ Ph)~iol~')OfSportAndExercisc.
A~ in other areas of scicnl·e, most of the quantita-
ti,·c research is p rescnted in the form of tables a nd
Units and Scientific grnphs. Tables ;md graplL~ pro,ide an efficient \\~I}' for
Notation research en; to commu nicate the resull'i of their studi es
A sci of intcn1atio1rnl stand;m:h for uni ts and abbrevia- tu other 11eicntisL~. For the student in exerci.~c and sport
tions (SI. U.· Sr5ti:me International d'Unite5) M:rn:s a.~ physiology, a working knowledge of how to read and
the prcfcrn·d units of mea.~uremcnt in exercise and intcrpn·I tabk-s a nd gr.aphs is critica l.
Tables are lL~ually 1L'ICd tu convey a large number of
spon physiult~'}·- In th is text,ahcmatc uniu in common
data puinlll or complex data that are affected by St."\-er..tl
use (such a.~ weight in pounds) a.-c often pl"O\ided a.~
facto~. Take table 0 .1 as an example. It is important to
well. Man y uf the.Ole uni L~ an: prmidt·d on the imide
first look al the title ufthe t.1ble. which identilics what
co,·cr of this text, and conn:rsions between and among
the SI unil~ and other uuits in commo n use arc found infonnation i.~ being presented. In this ca.<1e. the tabk
is desi1,,'11ed to illustr.uc how mrious conditions affeCI
on die i1t~idc bac k cm·e .-.
In common writing and e1,c11 in mathcmatio, the hean r.a tc. al rest and during cxerci.-.e. The It-ft-hand
r.nio bc.·twccn two numbers is typic.all y writte n u.~ing column . along "'ith the horizontal subheadings (like
a Msla.shM (/ ). Fur example, in di"}" air at 20 °C, the ~Humidity (21 °C) ,- specify the conditions under wh ich
speed of sound is 343 m /5. That notation work.~ well the heart r,llc wa.~ mca.'iurcd. Columns 2 and 3 prO\ide
the mean heart rnte ,-aluc.~ that correspond to each con-
for simple frnctions or raliOl'i, and we h an• mainmim:d
it in this text. Howe,·er, that nota1ion geL~ conflt~ing dition, \\ilh the middle rolumn ghing th e resting,-aluc
and the right-mo,;t column the exercise ,-alue. In e\"t"')'
for rclatiom• among SC\'er.al, that is, more than two,

Exercise Ph slol
An important segnent of exercise physiok>gy
concerns the response and adaptation of
people to extremes of heat, cold, depth,
and altitude. Understanding and controlling
the physiological stresses and adaptations
that occur at these environmental limits
have contributed directly to notable eocietal
achievements such as construction of the
Brooklyn Bridge, tho Hoover o.n, p,assurized
aircratt, and underwater habitats for the
commercial diving industry. ··---~-·~• ...... ,
41 of 642
Exercise Ph
AA important segment of exercise physiology
concerns the response and adaptation of
people to extremes of heat, cold . depth,
and altitude. Understanding and controlling
the physiological stresses and adaplations
that occur at these environmental limits
have contributed directly to notable societal
achievements such as construction of the
Brooklyn Bridge, the Hoover Dan, pressurized
aircraft , and underwater habitats tor the
commercial diving industry.
The next generation of environmental Chal-
lenges will also require such physiological
expertise. In January 2004, President George
Bush announced the Vision for Space Expk>-
ration, a strategy to first return humans to the
moon, then send explorers IO the planet Mars,
over the next 30 years. This ambitious plan
to construct permanent hi.man outposts on
the moon beginning in 2017, followed by 2.5-
year missions to the planet Mars, will require
effective countermeasures to minimize the
phySiological changeS that put space explor•
ersatrisk.
The continuous pull of gravity contributes to Or. James A. Pawelczyk.
the growth and adaptation of postural skeletal
muscles: loads bones, which increases their
size and density; and requires the cardiovascular system to maintain blood pr&SSl.1"8 and brain blood
flow. In a microgravity environment (free fall around the earth 01 constant-velocity conditions in deep
space), the reduction in loading leads to dramatic losses in muscle mass and strength. osteoporosis,
and ex8f'Cise in~rance at rates that mimic those seen in spinal OOfd-injufed patients
A series of dedicated space shuttle flights have studied these problems in detail. In 1983, the
National Aeronautics and Space Administration (NASA) began flying the European Space Agency-
developed Spacelab module, ushering a new era of internationally sponsored scientific research into
low-earth orbit. The Spacelab Life Sciences (SLS-1 , SLS-2) missions (STS-40 and STS-58) empha-
sized the study of cardiorespiratory, vestibular, and musculoskeletal adaptations to microgravity.
Subsequently the Federal German Aerospace Research Establishment (DLR) sponsored two mis•
sions (STS-61A and STS-68), perfecting a model of multidisciplinary, international investigation that
was emulated by the Life and Microgravity Sciences Spacelab mission (STS-78). which concentrated
on neuromuscular adaptation. The 1998 Neurolab Spacelab mission (STS-90), with an exclusive
neuroscience theme, concluded flights of the Spacelab module. Or. James A. Pawelczyk, a Penn
State exercise physiobgist and mission specialist on that flight. cotaught the first exercise physiol-
ogy class from space! Even roN, as of this writing. 250 mi (402 km) overhead, an active biomedical
research program continues on the International Space Station.
For the eX8fCise physiologist the question is what combination of resistance and "aerobic" exercise
training can prevent or diminish the changes that occur in space. At this time . the answer remains
unclear. Furthermore, if physical conditioning is required before and during space exploration and as
part of postflight rehabilitation. how should exercise prescriptions be individualized. evaluated. and
updated? Without doubt, further research in exercise and environmental physiology will be essential
to complete what Is destined to be the largest exploration feat of the 21st century.

2-.=

Physiology of Sporl and Exercise

good table and grJph, the unit\ for ,•ach ,-ariahle a re o n the y--axi.\, The uniu of measure for each ,-J riablc
dearly presented; in thi5 table, h ea11 r.tte ii; expressed are de:1rly di.\played on the grJph. Figure 0 .9 is in 1he
in Mbeal5/ min, Mor bt.·aL\ JX' r minute. Pay care fol atten- fonn of a line g1-aph. Linc grJphs arc usefu l in illu.,;trJ t·
tion to the unit\ of mea.~ure IL\Cd whe n interpreting a iug pa ncnls or trend\ in data but shou ld be n \Cd only
tab le or grnph. Frum this table-a rdati\"elp imple one tu co111pare two ,.iriables that change iu a continuous
hy ~ ientific .,;tandar&-·e see that both re.,;ting aud 111,mner (forexample,across time) and only ifboth the
e xercise heart rnte are increased by incrc-a.\Cd ambi- dependent and independent ,~1riabl,·s art· munhc r:<1.
ent temperature and humidil\', whil e- noi.,;e le,·d onh- In a line graph . if the dependent \"a ri ;1hlt- gn1-s up o r
affrc led resting !wan r.lte . Similarl}; consuming a larg~ down at a constant rnle with lhe iudc1x, ndc nt ,-.triable,
m,·al o r getting lcM than 6 h of sleep also mises heart the result \\il l bea .~trnight linl•, liowl"\"CI", in physiol<ll,')'
r,ue. These daia could not easily hau: been shown in tilt." r~ po1u,· p:llll'nl h1·1wccn ,.iriables is often 1101 a
graphical fonn . .~Ir.tight lim• but a cunl' of one .,;hape or another. In
Grnph.,; can pru\'idc a better ,ic,,,· of tn.•ncl\ in d ,u a, su ch c<~'les, p<ty du~ attention to the slope of,.1rious
rcspon!;l" p<me rns, a nd comparisons of data collected paru of the cmTe as it changes across the gr.tph. For
fmm t"'u or more groups ofsubjects. For some students, instance . figure O. IOshows the co ncentration of lactate
42 of 642 uni ts for each rnriable a re o n tlu: y-,axis. l11e uni t~ o f mca.~ ure for e;1ch ,~ufablc
dearly presente d : in th is tabl e , hea rt r J le is e xpressed. are cie'J rly d isplayed o n th e grnph. Figure 0 .9 is in the
in -beaL~/ min ,~ or beaL~ pe1· min m e . Pay care ful atte n- form o f a line g raph. Line grJphs arc IL~eful in illustrnf..
ti o n to th e uni ts of mea.~u rc IL~ed whe n interpret ing a ing pau cms o r tre nd~ in d ata but should be lL~cd uni)·
table or 1,rr.1p h. Fro m th i~ table- a rd;uiwly sim p le o n e to co mpare two ,11ri;1bles tha t c ha n ge in ;1 con1inuo u!I
by scie ntific standards- we see that both n:s ting and manner (fo r example, acrus.s time ) and o n l)· ifbo th th e
e xe rcise heart rnte arc increa.-,.ed by increased ambi- d ependent and inde pende nt ,.._riables are numbe n .
ent te mpernture and hum idity, while noi se ln ·d o nly In a line graph , if the d e pende nt ,-ari able g0e5 up or
affec ted r~ting lu:a rt r.tte . Similarly, con.~umi ng a large d m-.11 at a constant ra te with the independe nt ,-ari;1hlc,
meal or gening than 6 h uf .~Jee p also r.,i~ heart th e result will be as u-aight line. Ho wen·r. in php;iology
rn tc. Thei1e data could not c:t~il}· h:n-e been shown in th e res ponse p att e m between variables is often nol a
grnphical fonn. strnig ht line but a c une o f o n e sh ape o r ano ther. In
Gr.tphs can p rO\id e a better view o f tren ds in data, suc h case s, pay ci05oC attentio n to the slope o f ,·J rimn
response patterns, a nd comparisoru; o f data collec ted parts o f the curve a.~ it ch a ngClli acros.~ th e grJ ph . Fur
fro m m u o r mon : groups of subjrct~. Fo r so me smde n t~. insta n ce, fig ure 0 .10 sh owli the conn · nt r.llio n of lacta te
grJphs can be mo re diffic ult lO read and inte rpre t: in the blood a.~ sn hjt·c L~ \.\~1lk/ nm o n a treadmill at
hut gr,1.ph.'i arc, a nd will re m ain, a critical tool in the \w
dri o us increasin g speeds. At low treadmill speeds o f
undcn1a nding of cxe rci~ phy:1io logy. Fin t. (."\·cf}' grnph 4 to 8 km / h. lac tate incre ases \"Cf}' little. However, a t
ha.~ a horl1.on1al or X-ilxis for the independent vari able about 8 .5 km / h, a thresh o ld is re ache d beyo nd whic h
a nd o n t· (o r somctim t:5 two) ,·crtical or y..uc:L'i for the lactate increasc!I more d rJmatical ly. In m an y p hysiulog i•
d epende nt ,~.triable or,~triablcs. Independent ,.uiablcs caJ rc!ip<>nse!i, bo th th e th res hold (o nst•t of respou.'K')
a re th ose: fac ton th at a re manipulated o r contro lled by and th t" slope of th e respo nse b eyond tha t th resho ld
the rese-.trche r. while dependent ,.._riables an~those tha t are impo rta nt.
change \.\i th- th at is, d e pend o n---the indcpcndc n1 Data ca n a l.m be plo tt ed in th e fo nnat of a bar
,-ariables. gr.1ph. Bar gr.i phs arc commonly used whe n o nl y the
In figure 0 .9, time ofday i.~ the indepe nde nt \"Jriable d ependent mriabh: is a number and th e indepe nde nt
and is the refon~ placed a lo ng th e X-il.Xi.~ of the grn ph. va ria ble is a catcgo')'- Bar grap hs often sh ow treatm e nt
while h eart rJle i.~ the d epende nt ,-ariable (since heart effects, as in figure 0 . 7, whic h wa~ p rc\'i uusly diK IL~~d.
rnte d,pnuU on th e time of day) and is th erefore plo tted Figure 0. 7 shows the e ffect of d~~ t;m cc n m per week
(a categu')·) ou HD L-C (a numerical respo nse) in th e
bar grJ ph forma t.
145 - - - - - - - - - - -

'"ij_
c' 140 -

I + ~ --+ I •-

l::~ I , ,-~
2a.m.
_.,...,
6 a. m. 10a.m. 2p.m. 6p.m. 10p.m.
I
a •-
§

.,!I ·-
O
0
II
4
I
6
I
8 10
I I
12
I
14
t
---:-r
16 18
'
20
Tnillmnlll af>Nd {knVh)
i:::hei.lj-j:#•l#J Understanding how to read and inter•
pret a graph. This line graph depicls the relation between l#i(ijlh:#•IHI A line graph showing the nonlinear
!he time of day (on the x•axis. independent variable) and nature ol many physiological responses. This graph snows
heart rate during low.intensity exercise (on the y-axis, that. above a threshold (onset of response) ot about 8.5
dependent variable) that was measured at that time of day knv'h. the slope of the blood lactate response increases
with no change in !he exercise intensity. sharply

@zr•t41ill
In this introduction we highlighted the historical roots and scientific underpinnings of exercise and
sport physiology. We learned that the current state of knowl ed ge in these fiel ds builds on the past
and is merely a bridge 10 the luture-many questions remain unanswered . We briefly delined the
a~ute respon~es to exerc_i se_bouts and chronic adaptations to loflg•term training . We concluded
with an overview ol the princip les used in sport and exercise physiology research .
In part I. we begin examining physical activity the way exercise physiologists do as we explore
the essentials of movement. In the next chapter, we examine the structure and function of skeletal
muscle , how it produces movement. and how it responds during exercise.

Key Terms
acute exerc ise d iurnal variation longitudinal research design
chrOflic adaptation dose-response relation physiology
control g roup environmental physiology placebo group
43 of 642

@:nm
In this introduction we highlighted the historical roots and scientific underpinnings of exercise and
sport physiology. We learned that the current state of knowledge in these fields builds on the past
and is merely a bridge to the tu1ure-many questions remain unanswered . We briefly defined the
acute responses to exercise bouts and chronic adaptations 10 long-term training . We concluded
with an overview of the principles used in sport and exercise physiology research.
In part I. we begin examining physical activi ty the way exercise physiolog ists do as we explore
the essentials of movement. In the next chapter. we examine the structure and function of skeletal
muscle, how it produces movement. and how it responds during exercise.

Key TefTTIS
acute exercise d iurnal variation longitudinal research design
chronic adaptation dose-response relation physiology
control group environmental physiology placebo group
crossover design ergometer sport physiology
cross-sectional research design exercise physiology training effect
cycle ergometer homeostasis treadmill
dependent variable independent variable

Study Questions
1. What is exercise physiology? How does sport physiology d iffer?
2. Provide an example of what is meant by studying acute responses to a single bout of exercise.
3. Describe wha1 is meant by studying chronic adaptations to exercise training.
4. Describe the evolution of exercise physiology from the early studies of anatomy. Who were some
of !he key figures in the developmen1 of this field?
5. Describe the founding and the key areas of research emphasized by the Harvard Fatigue Labo-
ratory. Who was the first research director of this laboratory?
6. Name the three Scandinavian physiologists who conducted research in the Harvard Fatigue
Laboratory
7. What is an ergometer? Name the two mos! commonly used ergometers and explain their advan-
tages and disadvantages.
8. What factors must researchers consider when designing a research study to ensure that they
get accurate and reproducible results?
9. List several environmental conditions that could affect one·s response to an acute bout of exer-
cise.
10. What are the advantages and disadvantages of a cross-sectioflal versus a loogitudinal study
design?
11 . When should data be depicted as a bar graph as opposed to a line graph? What purpose does
a line graph serve?

Study Guide Activities

In addition to the activities listed in the chapter opening outline on page 1, two other activities are
available in the web study guide. located at
www.HumanKlnet1c1.com/Physiology0tSportAndExerclse

L, ..1 The KEY TERMS activity reviews important terms. and the end-of-chapter QUIZ tests your
understanding of the material covered in the chapter.

23