PHYSI

OLOGYOFMUSCULARSYSTEM 2014-
15

PHYSI
OLOGYOFMUSCULARSYSTEM
 Muscleisacont r
acti
l
et i
ssueanditi
soneoft hemostcomplexandor deredtissuei
nthebody.
 Musclei shighlyadaptablebychangingthenumberandsi zeofcellsaccor dingtodemand.
 Musclenotonl yundergocont r
acti
onbyshor teni
nginl
ength,theyalsocont r
actwit
houtshort
eninginlength
bydev elopi
ngf orce
 Therear ethreetypesofmuscl es–skeletal,cardi
acandsmoot hmuscl es.
 Totalmuscl econtentforms45t o50%oft hebody massofwhi ch40%i sbytheskel
etalmusclesandt he
remaining5- 10%i sbysmoot hornon-st
riatedmuscleandcar di
acmuscl es.
 Skelet
al musclesar ev olunt
aryinfunction,whi l
ethecardiacandt hesmoot hmusclesarei nvol
unt
aryi n
thei
ract i
vit
y.
Funct
ionsofSkel
etal
Muscl
es
1. Met
abol
i
sm ofskel
etal
muscl
escont
ri
but
est
othemai
ntenanceofbodyt
emper
atur
e.
2.Skeletalmusclespr ovi
det heforcef orsomat i
c( body )mov ement s,provi
decont r
oloflary
nx,
pharynxandpar tsoftheoesophagus, t
hushelpingi nv ocali
sation,degluti
ti
onetc.
3.Byadj ust
ingdiff
erentbodyparts,helptomai ntai
nbodypost ure
4.Externalanalandurethralsphi
nctershelpinurinaryandf aecal conti
nence.
5.Externaleyemuscl esfixtheeyedur ingheadmov ement s,thushel pstabl
eimagef ormati
onon
theretina
6.Mast i
cationandt onguemuscl eshel pinpr ehensi on,mast i
cationet c.Gastro-
int
est
inaltr
act
muscl eshelpinperist
alsi
sthatmov et hecontentst hrought heGIt ract
7.Heartandmuscl esi nbloodvesselsdistri
but
eandr egul atebloodt hroughoutthebody
STRUCTUREOFMUSCLE
 Eachmuscl
econsi
stsofmuscl
ecel
l
s,connect
ivet
issue,
bloodv
essel
sandner
ves
 Eachskel etalmuscleiscoveredbyat hi
nmembr anousconnect i
vet i
ssueout ercover
ingknownas
epimy sium.
 Enclosedwi thinthe epi
my si
um, each muscl eismade up ofa numberofbundl esknown as
fasciculus;eachsuchf ascicul
um i scov eredbyanot herthinmembr anouscov eri
ngknownas
perimy sium.
 Eachf asci culum i
nturni
smadeupofanumberoff i
breswhi charethemuscl ecell
s.
 Endomy sium i sthe connectiv
et issue structur
et hatcov ersthe indivi
dualmuscl e cel
l
s.The
connect ivet i
ssuesofthemuscl e–epi mysium andper i
my si
um consistofcoll
agenf i
bresandsome
elasti
cf ibres.
 Thecel lmembr anecover
ingthemuscl ecel
l,alsoknownasmuscl efibr
eiscalledassarcolemma.
MUSCLEFI
BRESorMUSCLECELLS(
Myocy
tes)
 Thesear
ethe phy
siol
ogi
caluni
ts oft
he muscl
e,whi
ch cont
ain or
ganel
l
es l
i
ke sar
cosomes
(mitochondri
a),sarcoplasmi
cr et
iculum wit
hanadditi
onalorganell
ethemyof
ibr
il
.
 Largenumberofsar cosomesispl acedinbetweenthemy ofi
bril
swhichi
ndi
catethel
argequant
it
y
ofener gy(
ATP)r equirementforthecontracti
onoffi
bri
ls.
 Skeletalmusclecellscontai
nmul ti
plenuclei(
100-200nuclei/cel
l)
.
 I
nv er
tebr
ates,
thematur
eskel
etalmuscl
ecel
lshavethenucl
eij
ustunderthecel
lmembrane
 Thesmoothandcardi
acmusclecell
saremono-
nucleat
edandthenucleusisl
ocatedi
nthei
nter
ioroft
hecel
l
.
 Thel
engt
hofskel
etal
muscl
efi
brer
angesf
rom 10t
o30cm;
diamet
err
angesf
rom 5t
o100µm
 Thefi
bresinskel
etalmuscl
er unfull
lengthofthemuscl
ewi
thoutanybr
anchi
ng.Eachmuscl
efi
brei
s
i
nner
vatedati
tsmi
ddlebyanerv
ef i
bre(axonfi
bre)
.
Sar
col
emma
 Thesarcolemmahasaspeci
alproper
tyofpropagat
ingtheact
ionpotent
ialsi
mil
artonerv
ecell
s.Attheend
ofeachfibre,t
hesar
col
emmalsheathfusestoformtendonfi
bre.Bundl
eoftendonfi
bresint
urnfor
mst he

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PHYSI
OLOGYOFMUSCULARSYSTEM 2014-
15

muscl
etendonwhi
chai
dst
heat
tachmentoft
hemuscl
eint
othebones.
Sar
copl
asm
 I
tisthefl
uidpresenti
nsi
deeachmuscl
efi
bresi
mil
ari
ncomposi
ti
ont
othatoft
hei
ntr
acel
l
ularf
lui
d.The
myof
ibr
il
saresuspendedi
nthi
sfl
uid.
Sar
copl
asmi
cRet
icul
um
 Theendoplasmicret
icul
um oft
hemuscl ecel
li
scalledassar
copl
asmi
cret
icul
um (
SR)
.
 ItstoresCa2+
2+
 Ca i sreleasedthroughspecialchannelsintosarcopl
asm
2+
 Ca - ATPasepumpr est
oresCa2+f r
om sarcoplasm t
oS.R.
 Ithasanast amosingchannel-
li
kest r
ucturethatsurr
oundseachmy obr
il

 Theplasmamembr aneofmuscl
ecellinvagi
natesandextendsi
nwardandisclosel
yassoci
atedwi
thSR.
Theseext
ensi
onsofplasmamembranearecal
ledtransv
erset
ubul
esorTtubul
es.
`
T'Tubul
eSy
stem

 TheTtubul
esarecommuni cati
onli
nkbetweenthecellmembr aneandmy ofi
bri
lsl
ocatedwit
hineachmuscle
cel
l
.
 Whentheplasmamembr aneissti
mulated,
theTtubulesalsorecei
vetheacti
onpotenti
alandthi
ssignal
sthe
SRtor easeCa2+ i
el ntothecy t
opl
asm whichinturninit
iatescontr
acti
on.Theseeventsoccurwithi
naf ew
msecsothatall
myofibr
il
swithi
nacellcancont
ractatthesamet ime.

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PHYSI
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Tr
iad
 Inskelet almusclecel l
s,twoTt ubulesarepr esentineachsar comereandt heyrunpar al
leltothej uncti
on
betweent heAbandandIband.
 TheSRi sdi l
atedattheAband- Ibandj uncti
onandt hispar tofSRi scalledast erminalci
sternawhi charein
closecont actwithTt ubules.
 Thear r
angementofTt ubulesy stem atthecent rewit
hci sternaeofSRonei t
hersideisdescribedast ri
ad.
 TheTt ubul escontainextracel
lularfluid.
 TheSRandTt ubuletogetherarecal ledsarcot
ubularsystem andi tsurroundseachmy ofi
bri
l
.
 Incardiacmuscl ecellseachsar comer econtai
nsoneTt ubul elocat
edadj acenttozdisk.
 TheSRcont nsCa2+pumppr
ai ot einswhichact i
velytr
anspor tsCa2+ionsfrom thecytoplasm i
ntotheci st
ernae.
2+
Thecist ernacontains98%oft het otalCa oft hecell
.Wi thintheSR, heCa2+i
t sstoredboundt ocalsequestri
n
-apr oteinwi thhighaf f
init
yforCa2+.Wi thinthecytoplasm t heCa2+ bindst oapr otein-calmodulin,which
tr
anspor tscy t
oplasmicCa2+t oitssiteofact i
on.
ELECTRONMI
CROSCOPI
CSTRUCTURESOFMUSCLEFI
BRES
My
ofi
bri
l
 Eachmuscl ef i
brecont ainssev er alhundr edt osev eralt housandmy of i
bril
s( cyli
ndricalst r
ucturesthatextend
throughoutt heent iremuscl ef ibr e)andt heyconst i
tut emor et han80%v olumeofskel etal muscl ecel
l.
 Eachmy ofibri
lis1t o2µm i ndi amet erwi thlengt happr oximat ingt hemuscl ef i
brelengt h.
 My ofibrili
sanor ganel leoft hemuscl ecel landshowsspeci alf unct onofcont
i raction.Themy of
ibri
lsare
arrangedi npar allel t
ot hel ongaxi soft hecel l
.
 Eachmy ofibri
lisconst i
tutedbyt wot ypesofmy ofilament s
 t hickormy osinmy ofilament s
 t hinoract inmy ofilament s
 Thinf il
ament sprimar i
l
ycont ainact in,tropmy osinandt roponi npr oteins
 Thickf i
lament spr i
mar ilycont ai nmy osinpr otein.
 The my of i
lament s ar e
arranged i n an or derly manner
whi chr esul t
si nr egular
repetiti
onofdense( dark) cross-
bands and l ess dense (li
ght)
bands. This
arrangement pr oduces cross-
stri
at i
onst hatar eseenmi croscopi call
yi nt heskel etal andcar diacmuscl es.
 Eachskel etalmuscl efibrecont ai nsabout1500mol ecul esofmy osinand3000mol eculesofact i
n.
 Themy osin( t
hick)andact in( thi n)f i
lament spar tiallyint erdigit
at e
 Thel i
ghtbandsseenundermi croscopecont ainonl yt heact inf i
lament sandt heyar ecal edas‘
l I’bands
becauset heyar ei sotropi ci .
e.t heydonotpol arisev isi
bl elight.
 Thedar kbandsar eknownas` A'band,whi char eani sot ropici.e.t heypol ari
sev isi
bl elight.Theycont ai
n
my osinf il
ament sandt heendsofact infilament swher et heyov erl
apt hemy osinfil
ament s.
 Smal lpr ojecti
onsofmy osi nf ilament scal ledcr oss- bridgespr otrudeoutoft hemy osinf i
lament sal
ongt he
entirelengt hexceptt hev er ycent reoft hemy osinf ilament .
 ThecentreofIbandscont ai
nadar kli
ne-thezl i
neorzdi sk.
 Theendsoft heact i
nf i
lament sareattachedtoZdi sks.From thez-disk,t
heacti
nf il
amentextendsi
nboth
dir
ecti
onsandi nter
digit
atewi thmy osi
nf i
laments.Thez- di
sksar efi
l
ament ousprotei
ns,passi
ngcrosswi
se
acrossthemyofibri
l
.
 Thatporti
onofthemy of
ibri
lthatli
esbet weentwosuccessi v
ez-disksi
scal edassar
l comere.
 Atrest,
whenthemuscl eisrelaxed,t
helengthofsarcomer eisabout2m.

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PHYSI
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 Thesar
comer
eist
hephy
siol
ogi
calcont
ract
il
euni
tofmuscl
e.

 Maximum contr
actionst
rengthdev
elopsatasarcomerel
engthofbet
ween2and2.25m.
 Atsarcomerel
engthsmor ethan3.65m (
thi
ckandthinf
il
amentsdonotoverl
ap)orl
essthan1.
27m (
thi
ck
fi
l
ament sar
epulleduptoz-l
ine)
,musclecont
racti
oncannotoccur
.
 Atthecent reoft heAband,al ight
erareai sseenwhi chist heH bandoft hemy of
ibri
landit
cont
ainsonlyt
hemyosi
nf i
l
aments.Thecentr
eoft
heHzonecont
ainsadar
kli edM l
necal
l i
net
hat
cont
ainsenzymesl
i
kecreati
nephosphoki
nase.
My
osi
nFi
l
ament
 Iti
scomposedofmul ti
plemy osinmol ecules,eachmol eculehavingamol ecularweightof480,
000.
 Eachmy osinmol ecul eismadeupofsi xpolypept i
dechai ns:
o Twoheav ymer omy sin( heav ychai n)
o Fourl ightmer omy sin.(li
ghtchai n)
 Thet woheav ychai nswr apspi rall
yar oundeachot herf ormingadoubl ehel ix.Oneendofeachoft hese
chainsisf ol
dedi ntoagl obul arst ructurecal edmy
l osinhead.Thuseachmy osinmoleculehastwofreeheads
l
y i
ngsidebysi deatoneendoft hemy osi nmol ecule.Theel ongatedcoil
edpor ti
onoft hemyosinmoleculei
s
caledt
l ai
l.
 The4l i
ghtchai nsf orm par toft hemy osinheads,t wot oeachheadandt heyhel ptocontr
olthefuncti
onof
theheaddur i
ngmuscl econt raction.
 Manymy osinmol eculesf ormt hemy osi nfil
ament .
 Thet ai
lsoft hemy osinmol eculesar ebundl edtof ormt hebodyoft hemyosinf i
lament
 Theheadsoft hemy osinmol eculeshangout war dtot hesidesoft hebody.
 Alongwi ththehead,par toft het ailoft hemy osinmol eculecalled“arm”ext endsoutwardfrom t
hebodyof
themy osinfilament .
 Thepr otrudi
ngar msandheadst oget herar ecalled“cross-bri
dges”.
 Thear msar ef lexibleandt hef l
exiblepoi nt
sarehi nges.

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 Thehi ngesall
ow theheadt obeextendedf aroutwardfrom thebodyoft hemy osinfi
lamentorbebrought
closertothebody.Therearenocross-bri
dgesatthecentreofthemy osi
nfil
ament .
 Themy osinheadfuncti
onsasanATPaseenzy mewhi chhelpstheheadt ocleavetheATP;theenergythus
releasedisusedtoenergi
sethecontracti
onprocess.
 Themy osinheadhasspeci alaf
fini
tytobi ndwithactinfi
laments.Thebi ndi
ngabi l
i
tyofthemy osi
nhead
withactinfi
lamentiscontr
oll
edbyATPaseact iv
ity.
 Int hepr
esenceofactinandATP,Mg2+i onsenhancethebindingofthemy osinheadwithact
infi
lament.
Act
inFi
l
ament
 I
tiscomposedoft
hreepr
otei
ns;
globul
aract
inmol
ecul
es,
tropomy
osi
nandt
roponi
n.
Act
in
 Theact i
nf i
l
amenti sadoubl e hel
ix
structure formed by
fi
lament ous`F'act i
nprotein.
 Eachst randofF- acti
nhel i
xhas
hundr eds of `
G'actin
mol ecules;
 AnADPmol eculei sattachedt oeachG-act
inmolecule.Itisthi
sADPmol eculethatprov
idestheact i
ve
bi
ndi ngsitetomy osinhead.
 Thebaseoft heact infi
lament si
sinser
tedi
ntothez-
discs,andtheot
herendpr
otrudesint
othesarcomere.
Tr
opomy
osi
n
 Thethinfi
l
ament sal
socont
aint
ropomyosi
nfi
lamentouspr
otei
nmolecules.
 Theyarelocatedint
hegroovef
ormedbetweenthetwochainsoft
heF- act
in.Duri
ngr
esti
ngstage,i
tcover
s
theacti
vebindingsi
tesoft
heacti
nfil
amentsothattheact
inandmy osinfi
lament
scannotbi
ndtogethert
o
producecontr
acti
on
Tr
oponi
n
 Itisacompl exof3gl obularproteins–t roponin-T,t
roponin-Iandtroponin-C
 TropninThashi ghaf fi
nitytotr
opomy osnandbi ndsthetroponincompl ext otropomy osinuni t
.
 Troponi n‘I’readi l
ybindswi thact inandt ropomy osin.Inrestinganimal s,theyinhibitATPaseact ivi
tyint he
my osinhead
 Troponi nChasav eryhighaffi
nityforCa2+i ons–eachmol eculecanbi ndupt o4Ca2=ions.Thecombi nati
on
2+
ofCa i onswi thtroponinisthetriggerthatiniti
atesthemuscl econtraction
 Thepr esenceoft roponiniandtropomy osin(tt-
compl ex)intheact i
nf i
lamenti nhi
bitsthebi ndingreacti
onof
actinwi t
hmy osin.
 Inther elaxedmuscl efibre,t
hett-compl exphy si
call
ycov erstheactivesitesofact i
nf orbindingwithmy osin
 Whenamuscl ei sst i
mul at
ed,ahi ghconcent ratonofCa2+ i
i ons( about10-6 to10-5)isr el
easedf r
om t he
cister
ni ntot hesar coplasm
2+
 TheCa i onsr eadilycombi newi tht het r
oponi n` C'oft het t–compl ex.Thi sbi ndingpr oducesa
conformati
onalchangeinthet r
oponincompl ex,whichmov est
het r
opomyosindeeperint
othe
groovebetweentwoactinst
rands.Thisexposestheacti
vesi
tesoftheact
inf
il
amenttomy osi
nand
i
niti
atescontr
acti
onsofthemy of
ibri
ls.
 Onlytheskeletalmuscl
eandcar di
acmuscl ebutnott hesmoothmusclerequi
restroponi
nand
tr
opomy osi
nforexecuti
onofcontracti
oncoupli
ng.
CARDI
ACMUSCLE
 Cardi
acmuscl ei sstr
iat
edandani nvoluntar
ymuscl ewhichhasst r
ucturesmuchl iketheskel
etalmuscle
exceptthatth
eindivi
dualfi
bresar
eshort(100-200µm)andbr anched;t
hebr anchesmergeorint
erdi
git
atesto
f mi
or ntercal
ateddiskswhi chcont
ainmanygapj uncti
ons.Thisproducesamuscul arnet
workcall
edt he
syncyt
ium‚ t
opr ovi
deforamor er
api
dspr eadofimpulsesfr
om musclefibr
et ofi
bre.

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 Contr
acti
leapparat
us( acti
n-myosin)isthesameasskel et
almuscl
efi
bres
 T-
tubulesyst
em issimilarexcept
:
 T- tubulesatthel eveloftheZ- l
iner athert
hanatt heAandIbands
 not ri
ads,assar coplasmi creti
culum notwel ldevel
oped,noterminal
cist
ernsf
ormed
 i oniccoupli
ngt hroughgapj unctionr egul
atescontract
ion
 Pur ki
njefi
bres-car diacmuscl ef ibresspecial
izedfori
mpulseconducti
on,par
toft
hepacemaker
-theyregulat epat t
ernofcont ractionwhi chneedst obenon- simul
taneousandco- ordinatedin
thr
eedi mensi ons
 Cardiacmuscl ehasabundantmi tochondr i
a.
 Cardiacmuscl ef i
bresdi vide,r
ecombi neandspr eadagai nandt heycontainmy ofibri
l
ssi mi l
arto
skeletal
muscl efibres.
 Dur i
ngexci t
ation-contractioncoupl ing,Ca2+ i
sr el
easednotonlyfrom thecisternaofsar coplasmic
reti
culum butal sof r
om t he‘T’t
ubul es.
 ThisCa2+ in‘T’t ubulescomesf rom ECF( becauseofi tsconnecti
onwi t
hECF) .Withoutthisextra
Ca2+,strengthofmuscl econt r
act ionwoul dbedr asti
call
yreduced.Consequent l
y,thequant i
tyof
Ca2+inthe‘ T’tubulesandt hestrengt hofcar di
acf i
brecontract
iondependst oagr eatextentont he
ECFCa2+concent rati
on( inskel
etalmuscl ef i
bres,ECFCa2+doesnotaf fectstrengthofcont racti
on
sinceCa2+i sreleasedf rom cist
ernaonl y)
SMOOTHMUSCLE

 I
tisanon-
str
iat
edmuscl
e,i
nvol
unt
aryi
nfunct
ionandi
sfoundi
nthev
iscer
alor
gansandt
hebl
ood
vessel
s.
 Thelengthofthesemuscl efi
bresvaryfr
om 20µm (bloodvessels)to1000µm (
uter
us)whi
char
e
short
erthantheskel
etalmusclefi
bres
 Theyhaveonenucleus/fibre.
 The rearenosarcomeresandtheZl i
nesar
eabsent
 nomy ofi
bri
l
s-actinandmy osi
nf i
l
amentspresent
,butnotorganized
 Smoot hmusclescontai
nbot hactinandmy osinfi
lament
s,whichhavesimi l
archaract
eri
sti
csasin
skeletalmuscles.
 Theor gani
sati
onofmy ofi
lamentsi sdif
ferentinsmoothmuscl ecell
s.Theact infil
amentsare
attachedt ostruct
urescall
ed“ densebodies”,someofwhichareat t
achedt ocellmembr aneand
othersar edi
spersedinthesarcoplasm.Theyareheldi
nplacebythestructur
alprot
eins.
 Dispersedamongt heacti
nf i
lamentsareaf ew myosi
nf i
laments(ar ati
oofabout15: 1actin,

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my osin).
 Becauset heact i
nandmy osinf i
lament sarenotarrangedi nr egi
ster,thesmoot hmuscl ecel lsdo
notexhi bitstr
iati
ons.
 SRi sr udimentaryandTt ubul esar eabsent.
 Insmoot hmuscl ecel s,Ca2+ i
l nfl
uxi smai nl
yf r
om extracell
ularflui
dwhi chent ersthecellt hrough
2+
Ca channel st oiniti
atecont racti
on.Smoot hmuscl econt racti
oni sst ronglyinf l
uencedbyECF
calcium.
 Troponi ni s notpr esenti n smoot h muscles.Int he place oft roponi n,smoot h muscle cel l
s
hav eanot herregulator
yprot eincalledthecalmoduli
nwhi chr eactswithf ourCa2+i ons.
 TheCa2+bi ndst ocalmodul inandt heCa2+-calmodul
incompl exact i
vatesmy osinlightchainki nase,
whichphosphor y
latesmy osi nli
ghtchai n.Thisact
ivatesmy osinATPaseandact insli
desov erthe
my osinpr oducingcont r
action.
 Thei nf uxofCa2+i
l ntothesmoot hmuscl ecellandit
ssl owdi ffusi
onint hecy toplasm andsl owness
ofmy osinATPaseact i
vit
ymakest hesmoot hmusclecont ractionaslowerpr ocess.

Smoot
hmuscl
esi
nthebodyar
ecl
assi
fi
edi
ntot
wot
ypes

1) Mul
ti
uni
tSmoot
hMuscl
eFi
bres
 About1%oft hesmoot hmuscleoftheanimalbodyispr
esentasmulti
uni
tsmoot
hmuscle.
 Eachfibr
eoper atesindependentl
yofotherfi
bresandasingl
enerveendi
ngmayinner
vateeachmuscl
efibr
e
(asinskel
etalmuscl efi
bres).
 Theyarecontroll
edmai nlybynervesi
gnals
 Theydonotexhi bitspontaneouscont
ract
ions.
 Theyarefoundi nt hecil
iarymuscleofeye,i
ri
softheeye,nict
it
ati
ngmembraneandpil
oerect
ormuscl
esof
hair
s.
2) Uni
tar
yorSi
ngl
eUni
tSmoot
hMuscl
eFi
bres(
Viscer
alSmoot
hMuscl
eFi
bres)
 About99%oft hesmoot hmuscl esoft heani malbodyar eoftheuni t
arytype.
 Manymuscl efi
brescont r
actsimul t
aneousl yasasingleunit.
 Thist ypeoff i
bresi sfoundi nt hehol low vi
sceralorgansi ncludingdigestiv
e,urogeni t
al,r
espirat
oryand
vascularsy stems.
 Thesef ibresoccuri nbundles,theircellmembr anesar eadherentt oeachot heratmul t
iplepoi
ntsandhence,
forcegener atedinonemuscl ecellistr
ansmi tt
edtot henext.
 Thecel lmembr anesarejoinedby‘ gapj uncti
ons’t
hroughwhi chi onscanf l
owf reel
yfrom onecel ltot
henext,
sot hatact i
onpot enti
alcantravelfrom f i
bretofi
bre; t
hiscausessi multaneouscontractionofallthefi
bresi
n
thatunit.Thistypeofsmoot hmuscl ear rangementisknownas‘ syncyt
ium’.
Char
act
eri
sti
csofskel
etal
,car
diacandsmoot
hmuscl
es
Char
act
eri
sti
cs Skel
etal
muscl
e Car
diacmuscl
e Smoot
hmuscl
e

Locat
ion Attached to bones Foundonl
yinhear
t Foundinthewal l
sofbloodv essel
s
(skelet
on) and int he organs of digesti
ve,
respi
rat
ory,ur
inar
yandreproductiv
e
tract
s

Funct
ion Mov
ementofthebody Pumpi
ngofbl
ood Contr
olofbl ood vesseldiameter
;
- prevent
ion of tr
anspor
toff oodt hr
oughGIt ract
;
mov
ementofbody emptyi
ng ofbladder;alt
erati
on of
pupi
ldiameter;accommodat i
on in

7
PHYSI
OLOGYOFMUSCULARSYSTEM 2014-
15

ey
e;hai
rmov
ementet
c

Anatomical Verylarge,cyl
i
ndr
ical
, Short cel ls wi th Small
,spindl
eshapedcellsj
oinedt
o
descri
pti
on multi
nucleat
ed cells bl
unt, branched eachotherbygapjunct
ions
arr
anged i n par
all
el ends.Cellsjoinedto
bundl
es others by
i
ntercal
ated di scs
andgapj uncti
ons

Ini
ti
ati
on of Onl
ybyaner
vecel
l Spontaneous Some cont
racti
ons al
way
s
contr
acti
on (pacemaker cell
s), maint
ained.Modif
iedbyner
ves
modi f
iedbyner
ves

Vol
unt
ary Yes No No

Gapj
unct
ions No Yes Yes

Speed and Fast

-50 msec ( 0.05 Moderate-
150 msec Sl
ow-1-
3 sec. Sust
ainabl
e
sust
ainabi
l
ity sec);notsust
ainabl
e ( 0.
15 sec) ; Not indef
ini
tel
y
ofcontr
acti
on sustai
nabl
e

Fat
igue Vari
es widel
y Low; r
elaxat
ion Gener
all
ydoesnotf
ati
gue
dependingontypeof between
skel
etal muscl
e and contract
ions
workload reduces the
l
ikeli
hood

St
ri
ated Yes Yes No

Ca2+ bi
ndi
ng Tr
oponi
n Tr
oponi
i
n Cal
modul
i
n
pr
otei
n

MEMBRANEPOTENTI
AL

 Thei
nter
sti
ti
alf
lui
dandt
hei
ntr
acel
l
ularf
lui
dar
eel
ect
rol
yti
csol
uti
onscont
aini
ng147mmol
/L
posi t
iveions (cat
ions)outsi
dethe cel
land al
mostthe same concent
rat
ion ofnegat
iveions
(anions)about155mmol /Li
nsidethecell
.
 Thei nter
sti
ti
alflui
dhashigherconcentr
atonofNa+i
i ons(
142mmol /
L)andlowerconcent
rat
ionof
+
Ki ons(5mmol /L).

8
PHYSI
OLOGYOFMUSCULARSYSTEM 2014-
15

 Thei nt racell
ularfl
uidhasmor eofK+i ons(140mmol /L)andlessofNa+ions(14mmol /L).
 Thei ntracellul
araswel last heext r
acell
ularfl
uidsar eelectr
ical
l
yneutrali.
e.thet otalposi
ti
ve
char geofcat i
onsi nICFandnegat iv
echar geofani onsofI CFareequal,simil
arlyf
orECF,bot h
char gesar eequal
 The concent rati
on difference ofi ons across a selecti
vel
y permeable membr ane creat
es an
electrical potenti
aldiff
erence.
 Allthecel l
si nthebodymai ntai
nanel ectr
icalpotenti
aldiff
erencethatcanbemeasur edacross
theirpl asmamembr anewhi chiscall
ed“membr anepot enti
al”
.
 Thispot enti
aldi f
ferenceacr osst hecellmembr anemakest hepl asmamembr anea polari
zed
membr ane.
Basi
cMechani
smsofMembr
anePot
ent
ial
(pol
ari
zedmembr
ane)
 Thr
eemaj
orf
act
orscont
ri
but
etot
hemembr
anepot
ent
ial
.
1.Different i
alper meabi li
tyoft hemembr anetodif
fusi
onofions.Therest
ingmembranei
s50to100
ti
mesmor eper meabl et oK+i onst hant oNa+i
ons.Ther
efor
e,posi
ti
vel
ychargedK+i
onsar
eal
l
owed
todi f fuseoutoft hecel lthr ough non- gated
l
eakchannel sdownt heirconcent rati
on
gradi ent( i.e.fr om i nsi deoft hecel ltothe out
side).
 AsK+ mov esoutoft hecel l
,negat iv
e
char gesi ncr easei nsi det hecel lwi thless
posi t
iv ei onsi nsidet hanout sidet hecel l
. This
resulti nt he gener ation ofa negat iv
e
i
nt r
acel lular pot ent ial whi ch i n t urn
attr
act sK+ i nwar dt her ebyr educest he effl
ux
ofK+.
 Consequent l
yt heconcent r
ation-dr ienK+
v effl
ux
outoft hecel lat tainsequi li
brium wi thK+ inf
lux
i
nt ot hecel ldr iv en byel ect ri
calf orce. The
membr ane pot ent ial at whi ch t his
equi l
ibr i
um occur s i s known as
equi l
ibr i
um pot ent ial.
 The r est i
ng membr ane i s al most
compl et elyi mper meabl e /v erysl ightly
permeabl et oNa+ i onsandt heNa+ i ons
cannotdi ffusebacki nt ot hecel l( even
though concent rat ion gr adientf orNa+
towar dst hei nsideoft hecel lpul sNa+ i
l t
cannotent erint ot hecel l
,since, Na+l eakchannel
sarenotopen)
 Themagnitudeofequi
l
ibr
ium pot
ent
ialpr
oducedbydi
ff
usi
onofi
ndi
vi
duali
onscanbepr
edi
ctedbyt
he
Ner
nstequati
on.
Si
mil
arl
y,t
heequi
l
ibr
ium pot
ent
ial
forNa+i
s+60mVandf
orchl
ori
dei
on,
iti
s-70mV

Themagnitudeofmembranepotent
ialatanygiveninst
antdependsupont
hedi
str
ibut
ionofNa+,
K+
-
andClionsandpermeabil
i
tyofthemembr anetoeachoftheseions.
 Thi
scanbepr edi
ctedbyGoldmanconstant
-fi
eldequati
on.

9
PHYSI
OLOGYOFMUSCULARSYSTEM 2014-
15

Wher
eVi smembranepotent
ial
,PK,PNaandPClarepermeabil
i
tiesofthemembr oK+,Na+ and
anet
Cl
;br
acket
sindi
cat
econcentr
ati
onandi andorefersi
nsideandoutsi
deofcel
l.

2. TheNa+,K+pump.Cel l
membr aneshav eanener gy-r
equiri
ngpumpwhi chpumpsNa+ionsoutofthe
celandK+i
l onsintothecellagai nsttheirconcentr
ationgradi
ent.
 Thi spumpgener at
essomemembr anepot enti
al,becauseitpumps3Na+ i onsoutofthecellfor
+
every2K ionspumpedi ntot hecel l
,thusconcent r
atingposi
ti
velychar
gedionsout
sidecel
l.Hence,
thi
spumpi scal
ledas“electrogeni cpump”
3. Negati
velychargedani
onsaretr
appedinsidet hecell
.Manyi
ntr
acellul
aranionsar
elar
gemol ecules
andaretoolargetomov eout
war dsthr
ought heplasmamembr ane.Hence, t
heyaret
rappedwithin
thecel
landar eattr
actedtotheinnersurfaceoft hecel
lmembr anebyt heaccumulat
edpositive
cati
onsjustontheoutersi
deofthecell
.
 Theset
hreef
act
orsar
ethepr
imar
ycauseofmembr
anepot
ent
ial
.
+
 Thedi f
fusionpot enti
alcausedbyK al onewouldgiv
eamembr anepotentialofabout–86mV,al mostallof
+ +
thi
sbei ngdeter minedbyK di ff
usion[Na contri
but
esl i
ttl
etot herest
ingpot ent
ialsincethemembr aneis
+ +
almosti mpermeabl etoNa] .AtthisK dif
fusi
onpot ent
ialthereisaslightexcessofcat i
onsoutsi
deand
anionsi nsi
det hecel l
.Anaddi t
ional–4mVi scontr
ibut
edbyt heelectrogenicNa+-K+ pump,gi
vi
nganet
resti
ngmembr anepot enti
alof–90mV.
Rest
ingMembr
anePot
ent
ial
sofSkel
etal
Muscl
eFi
bres
 Dur
ingr
est
ingst
age,
themuscl
efi
breshav
eanel
ect
ri
calpot
ent
ialof-90mV(
rangef
rom -
75t
o-90
mV) .Thisindicates elect
ri
calpot
enti
alinsidethe fi
breis-
90mV mor
e negat
ivet
han t
he
potent
ial
out
sidethefi
bres(ECF)
.
 Resti
ngmembr anepotent
iali
sal
socaledasdi
l f
fusionpot
enti
al
.
Exci
tabi
l
ity
 I
tist heabi l
i
tyofanycel lortissuetoexhibi
tanelectr
ochemicalchange(acti
onpotenti
al)toastimul
us.
 Allthel iv
ingcellsar eexcitabletostimuli
.Howev er
,nerveandmuscl ecellsarehighl
yexci t
ablecel
lsthan
othercel l
s.
 Allactionpotenti
alsgener atedbyapar ti
cularcel
lhavesi
milarchar
acter
isti
cs
 I
nt hener vefibr
e,informationsaretransmittedasacti
onpotenti
alandinthemusclecell,theact
ionpotent
ial
s
i
niti
at econtract
ion.Endocr i
necellsthatproducepepti
dehor monesorcatechol
aminesal sohavetheabil
it
yto
gener ateacti
onpot enti
al

10
PHYSI
OLOGYOFMUSCULARSYSTEM 2014-
15

EVENTSOFACTI
ONPOTENTI
AL
Act
ionPot
ent
ial
ofNer
veandSkel
etal
Fibr
es

 Theact
ionpot
ent
iali
s causedbyasequence
ofchangesorev entsoccur r
inginthemembranepermeabil
itytoNa+andK+i ons.Whenanexci t
able
cell
issti
mul at
ed,actionpotential
occur
sinthecel
ls.
 Acti
onpot enti
ali
st herapidchangesoccur
ri
ngi nt
hemembr anepotenti
alf
rom itsnormalnegati
vi
ty
toposit
ivepotentialt
hatlastforaperi
odoffewmi l
li
seconds,whichthenreturnsbacktoit
sorigi
nal
rest
ingpotenti
allevel
.
 Acti
onpot enti
alaidsintr
ansmi ssi
onofimpulsest
hroughnerveandmuscl efibres.
 The f irstev entofact ion pot entiali s char acterised byr apidi ncr ease int he permeabi li
tyof
membr anet oNa+ i ons( 5000f olds)t ointerioroft hecel l
,t husgener atingmor eposi t
iveelectri
cal
potent i
al insideoft hecel l
; thisi scal ledthedepol arizationst age.
 Thiscausesmor eofNa+i onsf l
owi ngintot hecel lthanK+i onsout flow.Thepot ent i
ali
nsidet hecel l
becomes“ zer o”ori nsomecasesbecomesposi ti
v e( overshoot )
.
 Thisi sf ollowed by i nact i
vat on ofNa+ channel
i s( closur e)t hatoccurwi thin anotherf ew
+
mill
isecondsandt hemembr anebecomesi mper meabl et oNa i ons.
 Becauset hei nsideoft hecel l hasbecomeposi ti
ve, Na+i nfluxisl i
mi t
ed.
 Thev oltagegat edK+ channel sopenbutt hisi ssl owerandmor epr olonged.Thi salowsK+ i
l ons
outflowt ot heext er i
oroft hecel lmembr ane.
 Thepot entiali nsidecel li sr e- establishedt oi tsnor malr est i
ngl evel( -
90mV) .Thi sstagei scalled
ast her epol ar i
sationst age
 Higherconcent r
ationofK+ i onsi nt heext er i
oroft hecel ltowar dst heendoft heact ionpot ential
+
cont i
nuesf orashor tper iod.Thi sexcessi veout fl
owofK i onscar riesmor eofposi t
ivechar get o
outside oft he membr ane,t huscr eates mor enegat ivit
yi nside whi ch isr ef
erred to as hy per -
polarisedst at e(afterhy per pol ar i
zation).Att hisst ate, r
e-exci t
at i
onoft hecellwillnotoccur .
 The f inalev enti schar act erisedbyel ectrogeni cpumpmechani sm whi chaidsint het r
anspor tof
t eeNa+ i
hr onst ot heext er iorf orev erytwo K+ i onst oi nt eri
oroft hecel landcr eatet he normal
rest i
ngpot ent i
al( -
90 mV)ont heinsideoft hecel lmembr ane.
 Thewholeacti
onpotenti
all
astsabout1-2mi l
li
secondsinmostner
ves,butl
ongerinmanymusclecel
ls.
+ +
 ThoughNa andK i onsmov eacrossthecel
lmembr anedur
ingact
ionpotent
ial
,thenumberofi
onsinvol
ved
i
smi nut
erel
ati
vetothei
rtot
al numbers.
Fact
orCausi
ngt
heAct
ionPot
ent
ial
+
 Thecel
lmembr
aneoft
heexci
tabl
ecel
l
shav
eionchannel
sthatar
eper
meabl
etoK i
onsandt
hey

11
PHYSI
OLOGYOFMUSCULARSYSTEM 2014-
15

arealway sopen-t hesear eknownasl eakchannelsandt heyar eresponsibl

eforest abl
ishi
ng
membr anepot ential
.Thecellmembr anesalsohaveNa+- K+ATPasepump
 Inaddit
iont othesel eakchannels,t
hecel lmembranesal socontainspecifi
cionchannel
swhi chare
permeablet ospeci f
icionslkeNa+,
i K+andCa2+.Thesechannel sar eguardedbygatesandopeni ng
andclosingoft hesegat esiscont r
olledbymembr anepot ent
ial
.Thesei onchannelsarecalledas
vol
tage-gatedchannel s.
 Both depolari
sation and repolari
sation ofthe nerve/muscle cellmembr ane duri
ng the acti
on
+
potenti
alisproducedbyt hevol
tage-gatedNa channel .
 Thev ol
tage-gatedK+ channelal soplaysani mport
antr oleinestabli
shi
ngt herepol
ari
sationofthe
membr ane.
Vol
tage-Gat
edChannel
soft
heCel
lMembr
ane
Vol
t edNa+channel
age-gat
 Havet
wogat
es-oneatt
heext
eri
orendwhi
chopenst
oout
sideoft
hecel
landr
efer
redt
oas
externalgat eoract ivationgat e.Ot hergat eisatt hei nteriorendandopenst oi nsideoft hecel land
referredt oasi nternalgat eorinact ivationgat e.
 Theact iv ati
ongat eiscl oseddur i
ngr estingst age( -
90mV) ,whi let hei nactiv
at i
ongat ei sopened.
Thispr ev ent sf reepassageofNa+i onsf rom out si
det oi nteri
oroft hecel l.
 Whent her est ingmembr anepot ent i
al( -90mV)dr opsorbecomesl essnegat iv e(-70t o50mV) ,
therei sasudden conf or mational changes i n t he act i
v ati
on gat e ( activated). Thi s
causesi ncr easedNa+i onper meabi li
tyasmuchas500t o5000f oldst oper mitf reepassageofNa+
i
onsi nt ot hecel lthrought hechannel sy stem.
 The dr op i nt he negat i
vit
yor gr adual i ncrease t o posi ti
vityf oll
owi ng t he openi ng oft he
activation gat e al socl oses t he inact ivati
on gat e.Howev er,thecl osur eoft hei nactivati
ongat es
i
scompar ativelyasl ow pr ocess.Thet i
mel apsebet weent heact ivati
onandi nact ivati
onoft he
channel scausest hepassageofNa+i onst ot heinterioroft hecel lforf ewmi l
liseconds.Att heend,
theinact i
v ati
ongat escl osesandnopassageofNa+i onst oinsideoft hecel lmembr ane.
 Dur i
ngr epol arisationst aget hei nactivationgat eiscl osed( inactivated) ,thuspr ev entsNa+i onent r
y
from out sidet ot hei nter i
oroft hecel l membr ane.
 Dur i
ngdepol ar i
sationst age,bot ht heact ivati
onandt hei nactivationgat esofNa+ ar eopened
+
(acti
v ated)andt heNa i onscanf reelypasst hrought hechannel sf rom t heext eriort othei nter
iorof
thecel l.

 Theinactivat
iongat
ewillnotr
eopenunt i
lthedist
urbedmembr anepotent
ialret
urnseit
hertoornear l
ytothe
ori
ginalresti
ngmembr anepot enti
allevel(-
70t o-
90mV).Therefor
e,withouttherepolar
isat
ionornear i
ng
+
compl eti
onofrepol
ari
sati
onofthef i
bre,i
tisnotpossi
blef
ortheNa channelt oopenagain.Thiscontr
ibut
es
tother ef
ract
oryper
iodintheexcit
ablecell
s.

12
PHYSI
OLOGYOFMUSCULARSYSTEM 2014-
15

Vol
tage-gat
edpot
assi
um channel
 Thi
schannelhasonl
yone gate
at t he i nterior oft he
membr ane.I t may ei ther close
oropent othei nterioroft he cell.
During r esting st age t his gate
i
scl osed( inactivated)and
K+ioni s pr evented f rom
passi ng t ot he ext er i
orof the
cell t hrought hischannel
system.
 Thesamechangei nr esting
potent ial( -
70mV—90mV) that
caused t he act i
vation of Na+
voltage- gated channel t o open
also causes K+ v oltage- gated
channelt oopen.Howev er
, the
openi ng ofK+ channeli s
slowerandt hisgat eopens only
when Na+ channel s get
i
nact ivat ed whi ch causes
decr easedent ryofNa+ i onsintot hecell
s,wher easK+ channelsacti
vat
ionall
owi easedK+ i
ncr ons
diff
usi on t oout sideoft hecells.
 Followi ngact i
onpot ential,t
hei nteri
orofthecellhasgai nedfewmor eNa+ionsandhasl ostf ewK+
+ +
i
ons.Wi thi
nf ewmsec, Na- K pumpr estor
esoriginali
ntracel
l
ulari
onconcentrati
ons
+
 Ifdepol ari
zat i
oni spr oducedr apidl
y,manyNa channel sopensi mult
aneouslyandthisov ercome
ther epol ari
sat i
onf orceandpr oducesdepolari
zat i
on.
 Ifdepol ari
zationi sinducedgr aduall
y,thegradualopeningofK+ channelsbalancestheopeni ngof
+
Na channel sandact ionpot ent i
aldoesnotoccur .Thi
sr esul
tsinaccommodat i
onofexcitablecel l
s
tost imul ati
on.

Rest
ingmembr
anepot
ent
ial
incar
diacmuscl
e:

 Cardi
acmuscl
e = -
85to-95mV
 Conducti
onsy
stem = -
90to-100mV
 Ventr
icl
emuscle = -
100to-105mV
 S.
A.node = -
50t
o-55mV
ACTI
ONPOTENTI
ALI
NCARDI
ACMUSCLE
 Cardiacmuscl ehasani nher entorint
ri
nsicproper
tyofgener at
ingi tsownact i
onpot enti
al
sr hyt
hmi
call
y,
i
ndependentofner vestimulati
on.Thisoccursinthepeacemakercel l
soft heS.A.node,AVnodeandPur kinje
fi
bres; SAdepolar
isesfast
ert hananyotherpart
softhehear tandist henor malpacemakeri nheart
.
 Thecar di
acmusclehassl owerbutpr ol
ongedacti
onpotentialthanskel et
almusclet hatlast
sfor150msecin
atri
aand300mseci nventri
cle.
 Incar diacmusclecel l
s,r
epol ari
sat
iondoesnotoccuri mmedi atelyaf t
erdepolari
sationbutt heposi
ti
v i
ty
remainsasapl at
eauneart hepeakoft hespikepotenti
al.Thispl ateaulast
sf oraf ew-hundredmsec.and
prolongst hecont
racti
onoft hecardi
acmuscl e.
ACTI
ONPOTENTI
ALI
NSMOOTHMUSCLES
 Thesmoot
hmuscl
eshav
elowerr
est
ingmembr
anepot
ent
ial
thansk
elet
alandcar
diacmuscl
es.

13
PHYSI
OLOGYOFMUSCULARSYSTEM 2014-
15

 Ther esti
ngpot entialofthesmoot hmuscl ecel
lsrangefrom –50mVt o-60mV.Theact ionpotential
i
sr egulat
edbyv olt
age- edCa2+channel
gat .
 Actionpot enti
alinv i
sceralsmoothmuscl eoccursintwof orms:
1)spi kepot enti
al
2)act ionpot enti
al wit
hplateau
 SpikePot ential
:Ty picalspikepotentialasi nskelet
almuscl eoccur si
nuni t
arysmoot hmuscles.
Theseact ionpot ential
scanbeel icit
edbyel ectri
calsti
mul at
ion,hormones,tr
ansmittersubstances
fr
om ner ves,byst r
etchorspont aneously.Ithasrel
ati
velyshor tdur
ati
on(5-10m sec)
 Actionpot enti
alwithpl at
eauoccur sinvisceral
smoot hmuscl ecell
ssimilart
ocardiaccells;
Rhy
thmi
cit
y(aut
omat
ici
tyorspont
aneousact
ionpot
ent
ial
s)
 Repet
it
ivedi
schar
geofi sesorr
mpul hyt
hmi
cit
ynor
mal
l
yoccur
sinhear
tmuscl
e(pacemakercel
l
s
ofSAnode),smoothmuscleandmanyneur onesofCNS.
 Toestabl
ishrhy
thmici
ty,
therest
ingmembr anepotent
iali
ntheseti
ssuesi
sonly-
55to-60mV( less
thanthenormalval
ueof-90mVi nskel
etalmuscle)whichisnotenought
oclosetheNa+ andCa2+
channel
s.
 Abnor
mal
ionconcent
rat
ionsi
next
racel
l
ularf
lui
d
+
 Reduct
ioni
nK concent
rat
ioni
nt shy
heECFi pokal
emi
aandi
tmakest
hemembr
anepot
ent
ial
mor
e
negat i
ve,andgr eaterintensi
tyofstimul usst rengthisrequir
edtostimulatemusclesandthepat i
ent
willsufferfrom muscl eweakness.Hy pokal emiaoccur sinhyperal
dosteronsi
m andincert
ainkidney
diseases
 ReducedCa2+concent rati
oni ntheECF, hy pocalcaemi amakest hecell
st ounder
goactionpotenti
al
easilywhi chcanl eadt omuscl espasmsandcr amps; hypocal
caemiaoccuri nvi
taminDdefici
ency,
hypopar athyroi
dism,mi lkfeverincows
 Hy pocalcemi aincowsbl ocksacetylcholiner eleasefrom synapsebet weenmuscleandner veand
causespar esiswher easi ndogsandhumans,t hisleadst ospontaneousdepolari
zat
ionofner ve
cellsthatproducesmuscul arspasm
MECHANI
SM OFMUSCLECONTRACTI
ON
1.Skel
etalmuscl
escont
racti
nresponset
ost
imul
ati
onbyCNS.Theef
fer
enti
mpul
sesf
ormuscl
e
cont ractionar ecar r
iedov ermot orner vest omuscl e
2.Anact ionpot ent i
altravelsal ongamot orner vetoi tsendingsi nthemuscl ef i
bre
3.Theendi ngsoft hener v esecr eteaneur otransmi t
t er–acet yl
choline(Ach).
4.Theacet ylcholineact sont hemuscl emembr anet oopenAch- gatedi onchannel s( l
igand-gated
channel s)ont hemuscl ef ibremembr ane
5.Achchannel sal lowNa+i onst of lowt ot heinter i
oroft hemuscl ef i
bremembr aneatt hepoi ntof
thener vet ermi nal.Thisi niti
atesanact ionpot entialinthemuscl ef i
bre
6.Theact ionpot ent i
altravelsal ongt hemuscl ef i
bremembr ane
7.Theact ionpot entialdepol ari
sest hemuscl ecel lmembr aneandal sot ravelsdeepl yintot he
muscl ecel lthr ought heT t ubul et ot het ermi nalci st
erna;thisr easesCa2+ i
el onsi ntot he
my of i
br i
ls
8.TheCa2+ i onsi niti
ateat t
ractionbet weenact inandmy osi
nf il
ament s,causi ngthem t osl ide
toget herwhi chi sthecont ractionpr ocess
9.Afteraf ractionofasecond, heCa2+i
t onsar er eupt akenintothesar coplasmi cret
iculum, stored
unt i
l t
heyar er eleasedf ornextcont ract i
on.
 Thispr ocessofanact i
onpot enti
alt r
av ellingalonganer vef i
brethatleadst ogener ationofact ion
potentialinmuscl ef i
brewhi chr eleasesCa2+ i ntot hesar coplasm andt heCa2+ initi
at esmuscl e
contractioni sref erredtoas“ excitation- cont r
actioncoupl i
ng”.

14
PHYSI
OLOGYOFMUSCULARSYSTEM 2014-
15

"
Wal
k-Al
ongTheor
y"ofCont
ract
ion(
Sli
dingMechani
sm ofCont
ract
ion)
 I
nther
elax
edst
ate,act
inf
il
ament
sfr
om t
wosuccessi
vez-
discsbar
elyov
erl
aponeanot
herandat
thesamet i
me, li
eadj acenttothemy osi nfi
lament s.
 Dur i
ngcont racti
on,act i
nf i
lamentssl ideov ert hemy osinf i
lament swi thshort
eningofsar comere.
Theact inf i
l
ament sar eact uall
ypulledi nwardt owar dtheM lineinar atchet-
li
kemannerbymy osi
n.
 Theact inandmy osinf i
lament sov erl
aponeanot hertoamaj orext ent.Thez-di
scsar ealsopull
ed
upt otheendsoft hemy osinfilament s.Thus,muscl econt racti
onoccur sbysl idingf i
l
ament
mechani sm.
 Thet roponin- t
ropomy osincompl exi nhibit
st heact i
vesitesont heact i
nf i
lamentsofrelaxedmuscle.
Hence, themy osi
nheadscannotat tachwi t
ht heactivesit
esofact intocausecont r
acti
on.
 Int hepr esenceofl argeamount sofCa2+ i ons,thei nhibi
toryef f
ectofT- Tcompl exont heactin
fi
lamenti sremov ed.
 WhenCa2+ i onsbi ndwi thtroponin-C,t hetr oponincompl exunder goesaconf ormationalchange;
thismov esthet ropomy osinmol eculesdeeperi ntothegr oovebet weent wostandsofact instr
ands.
Thisef fect“ uncov ers”t heact i
vesi tesofact in,hencet hemy osinheadat tacheswi ththeactive
sit
est ocausecont raction.
I
nter
act
ionbet
weenact
inandmy
osi
nfi
l
ament
s

 Assoonast heacti
nf i
lamentbecomesact i
v at
edbyt heCa2+ions,theheadsofthecr
ossbridgeof
the myosi
nf il
aments ar ei
mmedi atelyat
tract
edt ot he act
ivesit
esofthe act
infil
aments.This
causessli
dingmov ementofact inf i
lamentov erthemy osinfi
lamentwhichcanbeexpl ai
nedby
“Walkal
ong”theor
y( “
ratchet
”theory)ofcontracti
on.
1. ATPmol eculei spr esentboundwi t
hheadsoft hecr oss-bri
dgesofmy osinmol ecul
e.My osinheadhas
ATPaseact ivi
tywhichhy drol
ysesATPt oADPandi norganicphosphate(Pi
).Theheadext endsperpendi
cular
totheact i
nf i
lamentbutdoesnotbi ndwi thactin.
2. Thetroponi n-tropomy osincompl exbindswi thCa2+ions.Act i
vesit
esontheact i
nfi
lamentsareuncovered.
3. Thebondbet weent heheadoft hecr oss-bri
dgesandt heact ivesit
esoftheact i
ncauseaconf or
mat i
onal
changei nt hehead.Theheadt il
ttowar dsthear m ofthecross-bri
dge.Thi
stil
tingoftheheadisknownast he
“power-stroke”;thispul l
st heact i
nfilamentwhi chslideov erthemy osi
nf i
lament.Ener
gyr equi
redfort he
power-strokeispr ovidedbyspl it
ti
ngt heATPat tachedwi t
htheheadoft hecrossbri
dges.
4. Onc etheheadoft hecr oss-bri
dgei st i
lted,theADPandPiat tachedtot heheadar ereleased.When
theADPi sr el
eased,anew mol eculeofATPbi ndswi t
ht heheadofcr oss-
bri
dge.Thisbindingof
ATPtocr oss-br
idgeheaddet achest heheadf rom theact i
vesite.
5.Thecrossbr i
dgeheadboundwi thnewATPbegi nt henextcy cl
eleadi
ngtoanotherpower-
stroke,i.
e.
thedetachedheadi s‘cockedt oi t
sper pendicularposi t
ionreadytobegi nanotherpower-str
oke
cycl
eagain.Dur i
ngr apidcont racti
onofskel etalmuscl e,eachmy osi
nheadunder goes5cycles/sec.
Eachmy osinmol eculehy drolyses5-10ATPmol eculesev erysec.
6.Whent hecockedheadwi thitsst or
edener gybi ndswi t
hanewact i
vesit
eont heacti
nfil
ament ,it
becomes‘ uncocked’ andr epeatst hepower -
strokeandt heact i
nfil
amentispull
edtoonemor estep.
 Simil
arrepeatedbackandf ort
hmov ement soft
he cross br
idgemakes theheadswalkalongt
heacti
n
fi
lamentstepbystepthuspullt
heactinfi
lamentt
owardsthecentr
eofthesarcomer
e,unt
ilt
hez-membr
ane
reachest
heendsoft hemyosinfi
laments.
El
ect
romy
ogr
am
 Whenanact
ionpot
ent
ialspr
eadsal
ongamuscl
efi
bre,asmal
lpar
toft
heel
ect
ri
calcur
rent
gener
atedspreadsawayf rom thefibr
ewhi chcanreachtheov erl
yingski
n.El
ectr
odesplacedonthe
ski
norinsert
edi nt
ot hemusclef i
brecanr ecor
dtheelectr
icalpotenti
alwhenthemuscl
econt r
acts.
Suchameasur ementi scal
ledelectr
omy ogram.
 EMGhel pstoidenti
fywhetheraweaknessorpar al
ysi
si namuscl eisduetodiseasei
nt heskel
etal
muscle,neur
omuscul arj
uncti
on, motorneuronorCNS.

15
PHYSI
OLOGYOFMUSCULARSYSTEM 2014-
15

ENERGYFORMUSCLEACTI VITY
 Energyi
sr equi
redbythemuscl efort
hefol
lowingf unct
ions
 Energyfrom thehydroly
sisofATPisconvertedtomechanicalworkt
hatcausesmy
osi
nheadt
obend
 ATPi srequir
edforbreakingthemyosi
nheadat t
achmentwithact
infi
lament
 Energyisrequir
edtor emoveCa2+fr
om cytosol t
otermi
nat
econt r
act
ion
+ +
 Tomai ntai
nNa- K pump
 Theimmedi at
esour ceofenergyformuscl
econt racti
onisprov i
dedbyATP.
 Thesarcoplasm containsabout4mM ofATP,whi chissuffici
enttomaintai
ncont
ract
ionf
or1-
2seconds.
OncetheATPi sbrokeni nt
oADPandener gy(Pi),t
heADPi srephosphor
ylat
edt
oform anewATPwit
hina
fr
acti
onofasecondi nthemuscle.
 Thereareseveral
sour cesofener
gyforthi
sr ephosphory
lat
ion.
 Fi
rstsourceofenergyi st
hephosphocreat
ine.
Cr
eat
inePhosphat
e(Phosphocr
eat
ine)

 It
sconcent rationint hemuscl efibreis4t o8t imesmor et hanATP.Si milartoATP, eachmolecul eofcreati
ne
phosphat ecar ri
esahi gh-energyphosphat ebond,butwi thl i
ttl
emor eenergy(9500cal )whil
ecompar edwith
ATP( 8000cal )perbond.
 Thecr eat i
nephosphat ei sinstantlycleav edandr el
eases~Pi ,whichbi ndswi thADP der i
v edfrom ATP
breakdownt or econst itut
eanew ATP.Theenzy mecr eatinekinase( phosphokinaseorphosphocr eati
ne
kinase)cat alyzest hisreaction.
 Thecr eatinephosphat ei nthemuscl eisreferredtoasATPspar erorenergybufferofthemuscl e.
 Thecombi nedener gyofbot ht hest oredATPandt hecr eati
nephosphat eint hemuscl ecausesmaxi mal
contractionf ornotmor ethan5- 8seconds.
 Duringmuscl econt r
action,concent rati
onofcr eat
inephosphat edecreaseswhi chmustber egeneratedto
sustaincont ract i
on.Thi soccur sthroughcel l
ularoxidatonbygl
i ycol
ysiswhenmuscl econtract i
onproceeds
forupt o2mi n.
 Forsust ainedcont racti
onbey ond2mi nutes,oxidat
ivemet abolism i
susedt oprovideenergy.

Gl
ycolysis
 Themuscl eglycogen(1%ofmuscl
e)i
senzy mat i
cal
lybr
okendownt oglucosewhi
chundergoesgl
ycol
ysi
sto
produce2mol eculesofATP and2mol eculesofpy ruv
icacidorl act
icaci
d;theli
berat
edenergyfrom
glycol
ysisisusedtoconver
tADPtoATPandr e-synt
hesi
sephosphocreati
ne.
 Thegl ycol
yti
ccy cl
eoccursver
yrapi
dlyandev enintheabsenceofO2,butendpr oductsli
kelact
icaci
d
accumul at
esinthemuscle.

16
PHYSI
OLOGYOFMUSCULARSYSTEM 2014-
15

 Thegl
ycol
ysi
scansust
aincont
ract
ionf
orabout1t
o2mi
nut
es.
Oxi
dat
ionofNut
ri
ent
s
 Themaj orsour
ceofener
gyf ormuscl
econt
ract
ioni
soxidati
vemetabol
ism occur
ri
nginmit
ochondr
iainthe
presenceofO2.
 Mor ethan95%oftheenergyusedbythemusclesf
orsustai
ned,l
ong-t
erm contr
act
ioni
sderi
vedfrom t
his
source.
Ri
gorMor
ti
s
2+
 Immedi at elyf ol
lowi ngcont r
action,thecy t
oplasmi cCa i spumpedbacki nt
oSR.I ft
hispumpi ngisinhibi
ted,
rel
axationofmuscl ecannotoccurev ent hought her
ei snoact ionpot ential.Muscler emai nsinsust ai
ned
contr
act ion( i.e.cont racti
onwi thoutt hepr esenceofact i
onpot ential)whichisknownascont racture.
 Whenmuscl ef ibresar ecompl etelydepl etedofATPandcr eatinephosphat e,t heydevelopast at
eofr igi
dity
cal
ledr i
gor .
 Ther i
gort hatdev elops5t o6haf terdeat hi scal ed r
l igormor ti
s.Iti schar act
erisedbyt hef ai
lureof
separationoft hecr oss-bri
dgesf r
om t heact inf i
lamentsoft hecont r
actedmuscl eduet olackofATP.The
l
ossofcel lfunct i
onandf ai
lureofr eplenishingtheATPaf terdeat har etheprincipalcausesofr i
gormor t
is.
2+
 Ca pumpsr unoutofATP
2+
 Ca c annotber emov edf rom cy t
opl asm
 r emai nsi ncont inuouscont raction
 e v ent ual lyt i
ssuesbr eakdown
 Themuscl er emai nsi nr i
gorasl ongast hecont ractil
eproteinsar eintact.
 Theput rificationofdecomposi t
ioni susual lycausedbyaut olysi
sbyt heenzy mesoft hel ysosomes, whichare
rel
easedi nabout15t o25hour saf terdeat h.Theydenat ureordest r
oyt heprotei
nsandt hedeadbody
becomesf laccid.
Heatpr
oduct
ioni
nmuscl
e
 Whenamuscl econtracts,worki
sperformedandenergyisuti
li
zed.
 Thegreatertheamountofwor kperformedbythemuscl e,great
ert
heamountofATPthati
scl
eav
edand
l
argertheamountofheatr el
eased.Thisi
scal
ledFenneffect.
 Ofthetotalenergyli
beratedfrom ATP,themuscleut
ili
ses40t o50%ener
gyi
nperfor
mingwor
kandther
est
i
sliber
atedasheat .Thishelpstomaintai
nbodytemperature.
 Heatpr
oducedi
nthemuscl
eiscl
assi
fi
edas:
 Rest i
ngheat :Heatproducedi nmuscl eatr est.I
tindicatesheatreleasedbybasal met abolicprocesses.
 Initi
alheat :I
tisproducedbot hati ni
ti
ationanddur i
ngt hecourseofmuscl econt raction.Itconsistsof
o Act i
v at
ionheat :Heatpr oducedi namuscl ewheni tbeginscont racti
onpr ocess.I tisduet oel ect
ri
cal
ev entsandmechani cal acti
onsassoci atedwi t
hi ni
tiati
onofcont racti
on
o Shor teningheat:Heatl i
berat edbyslidingprocess; whenmuscl el if
tsloadanddoesext ernalwork
 Mai ntenanceheat :Heatl i
beratedi namuscl ethati sst i
mulatedbutdoesnotl eadt ophy si
calwor k
(isomet ri
ccont racti
on) ;iti spr oport
ionalto dur ation ofcont r
act i
on and tension dev el
oped during
cont racti
on.
 Rec overyheat :Heatliberatedaf tercont r
acti
onofmuscl efibr
e.Iti sduet opumpi ngofCa2+ i onsback
i
nt ot het ubul
esandr esy nthesisofcr eati
nephosphat eandATPandot herpr ocessest hatrestorethe
muscl etoprecontractionstat e..
TYPESOFWHOLEMUSCLECONTRACTI
ON
 Mus cl
econt
ract
ioni
spri
nci
pal
lyoft
wot
ypes.
1) I
somet
ri
ccont
ract
ion
 Musclecontract
ionwit
houtshorteni
nginlength.Whenananimaltr
iest
opushaheav
yobjectwit
houtmovi
ng
i
t,concer
nedmuscl esgeneratef or
cewi thoutshor
teni
ng.Tostudythei
sometr
iccont
racti
on,themuscl
e
hastobefixedbetweenimmov ablepoi
nts.
2) I
sot
oni
ccont
ract
ion

17
PHYSI
OLOGYOFMUSCULARSYSTEM 2014-
15

 Whenamuscl econtr
actsandshort
ensinlengt
h,butt
hetensi
onont
hemuscl
eremaini
ngconst
anti
tiscal
l
ed
i
sot
oniccont r
acti
on.Thisty
peofcontracti
onoccurwhi
lel
if
ti
ngawei
ghtormovi
ngalimb
 Tostudyisotoni
ccont r
acti
on,themuscleisatt
achedbyoneendtoanimmovabl
epointandtheot
hertoa
movablepoint(l
everormov abl
eweight)
.
 Char
act
eri
sti
csofi
somet
ri
candi
sot
oni
ccont
ract
ionsofskel
etal
muscl
e
I
somet
ri
cCont
ract
ion I
sot
oni
cCont
ract
ion
1 Cont
ract
ion bet
ween t
wo f
ixed
Cont
ract
ion bet
ween one f
ixed and t
he
poi
nts ot
hermov
eabl
epoi
nts

2 Nosl
i
dingmov
ement
snoshor
teni
ng
Showssl
i
dingmov
ement
sshor
teni
ngi
n
i
nlengt
h l
engt
h

3 Manif
est
ation of the process of
Mani
fest
ati
on of t
he pr
ocess of
cont
ract
ion l
astthroughoutthe perod of c
i ont
ract
ion l astlongerev
enafterthe
cont
ract
ion complet
ionofcontr
acti
onofthemuscl
e

4 Nowor
kper
for
mance
Wor
kisper
for
med

5 Lessener
gyr
elease,
lowFennef
fect
"
Mor
eener
gyr
elease,
high"
Fennef
fect
"

Cont
ract
ionsofMuscl
eint
heBody

 I
nthebody ,mostofthemuscul
arcontr
act
ionsareamixt
ureofbothisometr
icandisotoni
ccont
racti
ons.
 Duri
ngstandingpostur
ethequadri
cepsshowsisometr
iccont
ract
ion,thust
ightenthekneejoi
nttokeepthe
l
egsstif
f.
 Contr
acti
onofthebicepsi
sisot
onicwhenthehandli
ft
stheweight.
 Alt
ernateisometri
candi sotoni
ccont r
actionsofthequadr i
cepsandgast rocnemius(calfmuscle)
helptoeff
ectrunni
ngorwalki
ng.Thequadr
icepscont
ract
si sometr
ical
l
ywhent hef
oothi
tthe
ground,
whil
ethecal
fmuscl
econtr
act
sisot
oni
call
ywhenthefootisl
if
tedofft
hegr
ound.
Ser
iesEl
ast
icComponentofaMuscl
e

 Themuscl efunct
ional
l
yconsi stsofcontracti
l
eel ement sarrangedinparal
lelwiththeelasti
ccomponents
(sarcol
emma,connecti
vet i
ssueet c)andinser i
eswi thasecondsetofel asticcomponent scaledser
l i
es
elasti
ccomponentsv i
z.tendons,connect i
vet i
ssuel inki
ng musclef i
brest otendonsand cr ossbri
dge
elements.
 TheSECcomponent sstret
chsl i
ghtl
yastensionincreases.Consequentl
y,t
hecont r
acti
leunitmustshor
tenan
extra3-5%tomakeupf orthest ret
choftheseel ement s.Theseelementsofthemuscl ethatstr
etchduri
ng
contract
i eknownasser
onar i
esel ast
iccomponent softhemuscl e.
Ty
pesofMuscl
efi
bres
 Basedont
hev
eloci
tyofcont
ract
ion,
themuscl
efi
bresar
ecl
assi
fi
edas
 slow(toni
c)orTypeIfi
bres
 fastorphasi
c(twit
ch)orTypeI
Ifi
bres
 Ev
erymuscleinthebodyiscomposedofslowandf
astmuscl
efi
bres.
 Compar
isonbet
weensl
owandf
astmuscl
efi
bres
Sl
owMuscl
eFi
bres FastMuscl
eFi
bres

18
PHYSI
OLOGYOFMUSCULARSYSTEM 2014-
15

1. Smal
l
erf
ibr
es Muchl
argerf
ibr
es

2. I
nner
vat
edbysmal
lner
vef
ibr
es I
nnerv
atedbycompar
ati
vel
ylar
gener
ve
f
ibr
es

3. Hav e extensi
ve blood suppl
y,hence Havelessbl
oodsupply,hencecall
edas
referr
ed as " Red muscle"
, shows "
White muscle", shows qui ck and
prolongedper formanceofwor k(
Long r
epeti
ti
vecont
racti
ons(Shortcont
ract
ion
contracti
ont i
me) . t
ime).

4. I
ncr
easednumberofsar
cosomes Fewersar
cosomes

5. Muscle met aboli

sm by aer obi
c orGl ycolysi
s or anaer obi
c t ype of
oxi
dati
on ofgl ucose,fat
ty acids and metaboli
sm liberat
ing 2 molecul
es of
aminoacids,causest herel
easeof38 l act
icacidand2mol ecul
esofATP.
6. Lar
ge amount of my ogl
obin i n Lackofmy ogl
obin.
molecul
esofATP.
sar
coplasm.

7. Lesssar copl
amicr eti
culum and l
ess Extensive sarcoplasmicr eti
cul
um and
Caionr el
ease. rapidCai onrelease.
8. Hasl argemot orunits(Moremuscl e Hasf ewmot orunits(l
essmuscl efibr
es/
fi
bres/ neuron), nofine degree of neuron),higherdegreeoffinecont r
ol.
9. Ad
coa
np
trt
e
od f
l
. or prolonged, conti
nued Adapt edf orrapidandpower f
ulmuscl e
muscleact i
vit
y– l i
kesuppor tofthe contract
ions–l i
kejumping,running
bodyagainstgravi
ty
 Basedonst
ruct
uralandf
unct
ionalchar
act
eri
sti
cs,
skel
etalmuscl
efi
brescanbecl
assi
fi
edi
ntot
wo
ty
pes–
 slow- twi
t ch(t
y pe-)oxi
I dati
vef i
bres
 fast-twit
ch( type-I)gl
I ycoly
ticfi
bres
 TypeIfibres,alsocal edsl
l owt wit
chorsl owoxi
dati
vefi
bres(redmuscl es)containl argeamounts
ofmy oglobin,manymi tochondr i
aandmanybl oodcapil
l
aries.Thesef i
bresarer ed,spl i
tATPata
sl
owr ate,producel owamountoff orce,haveaslowcontract
ionveloci
ty,ver
yr esistanttofati
gue
andhav eahi ghcapacitytogener ateATPbyoxidati
vemetabolicprocesses.Suchf ibresarefound
i
nlargenumber sint hepost ur
almusclesoftheneck.
 Fasttwit
ch–t ypeII-fi
bresproducehi
ghamountoffor
ce,cont
ractver
yquickl
y,andcont
ainl
ess
my ogl
obinandfewmi t
ochondri
a.Thefastt
wit
chf
ibr
es(whit
emuscle)aref
urthergr
oupedi
ntot
wo
subgroups-TypeII
AandI IBf
ibres.
 Ty
peI IAFi bres
o Thesef i
bres,al
socal edf
l asttwitchorf astoxidati
veglycoly
ticf
ibr
es( FOG,)haveav eryhi
ghcapaci t
y
forgener ati
ngATPbyoxi dativ
emet abol
icpr ocesses,spli
tATPatav eryrapi
dr ate,haveaf ast
contractionv el
ocit
yandar eresi
st antt ofati
gue.
 Ty
peI IBFi bres
o Thesef ibres,also cal
led fastt wi tchorf astglycolyt
ic(FG)f i
bres,containrelati
vel
yf ew blood
capill
ariesandl ar
geamount sgl y cogen.Ty peI IB f i
bresproducet hehighestamountoff orce,
gener ate ATP by anaer obic gly colyti
c processes,notabl et o supply skel
etalmuscl ef i
br es
continuousl ywithsuffi
cientATP,f at i
gueeasi l
y,spli
tATPataf astrateandhav eaf astcontr
act ion
velocity.

19
PHYSI
OLOGYOFMUSCULARSYSTEM 2014-
15

o Suchf
ibr
esarefoundinlar
genumber
sint
hemuscl
esoft
hear
ms.
o TypeI
IBfibr
esareabsenti
ncani
nes.
 Mostmusclesi
nt hebodyar eamixtureofsl
owandf astf
ibres
 Ty
peIfi
bresdevelopl owforcebutprovi
desustai
nedcontracti
onforlongerper
iodsandhel
pinl
ongdi
stance
movementsandpost ure.
 Ty
peIIf
ibr
esdev elophigherfor
cebuteasil
yfat
igabl
eandhel pinf
astermov ementsf
orshor
tdur
ati
on.
Mot
orUni
tsoft
heMuscl
e
 Eachmot orneuronleav
esthev ent
ralhornofthespinalcordandinnervatesmanymuscl ef
ibr
es.
 Allthemuscl efi
bresinamuscl ethatareinnervatedbyasi nglemot orneuron(allt
hemusclefibr
essuppl
i
ed
byasi nglemotorneuron)arecaledasmot
l orunit
.
 Ther ati
oofmot ornervetot henumberofmuscl efibresinner
vat edinagi venskelet
almuscleiscal
l
ed
i
nner vat
ionrati
o.
 Thenumberofmuscl efi
brespermot orunitdif
fersaspert hef unct i
onforwhichthemuscleisusedinthe
body .
 Smal lmuscleswhichactrapidlyandwhi chrequir
ef i
nercontrolofmov ementhaveaninnerv
ati
onrati
oof1:2
to1:4.
 Largemuscl esthatdonotrequirefi
nercontrolhaveani nnerv
ationrat i
oof1:800ormore.

MECHANICAL
CHARACTERISTI
CS OF
MUSCLE CONTRACTION
St
imul
us

 I
tisanexter
nalagent
,whenappl
i
edt
oanexci
tabl
eti
ssuepr
ovokechangei
nmembr
anepot
ent
ialor
avisi
bler
esponse.
Ty
pesofSt
imul
i(Basedonst
rengt
h)

 Subt hr
eshold:(
submi ni
mal )l
owi ntensi
ty,
unabl
etoproduceavisi
bleresponse
 Threshold:minimum i ntensi
ty,st
rongenoughtoini
ti
ateanacti
onpot ent
ial
 Submaxi mal:higherthresholdstrength
 Maximal: hi
ghestthresholdstrength
 Supramaxi mal:destructiv
etot i
ssues
Cl
assi
fi
cat
ionofSt
imul
us
Mechani
cal
:pai
n,pr
essur
e,andt Chemi
ouch; cal
:aci
d,al
kal
i
;Ther
mal
:war
mth,
col
d

Osmot
ic:Hy
per
toni
csol
uti
on,
hypot
oni
csol
uti
on

Al
lornonel
aw
 Whenamuscl efi
breissti
mulatedbymi ni
malormaxi
malef
fecti
vest
imul
i
,thewhol
efibr
econt
ract
stothe
maximum ori
twil
lnotcont
ractatal
l.
 Inskelet
almuscl
es,thi
slaw isappli
cabletosi
ngl
emotorunit(mot
ornerveandal
lthemusclefi
bresi
t
suppl
ies)
.

20
PHYSI
OLOGYOFMUSCULARSYSTEM 2014-
15

 Whol
ecar
diacmuscl
eact
sinal
lornonemannerduet
othepr
esenceoff
unct
ional
syncy
tium
Si
mpl
eMuscl
eCont
ract
ion:
(SI
MPLETWI
TCH)

 Appli
cat
ionofsingl
ethresholdl
evelofsti
mul
us(el
ectr
icalst
imul
ati
on)causesasi
ngl
emusclecontr
act
ion
fol
lowedbyrel
axati
on.Thetimetakenfort
hesi
mpletwi
tchis100-120msec.Thesi
ngl
emuscl
etwit
chhas3
phases.
a) Latentperi
od
 Iti
st heper
iodbetweent heappli
cati
onofsti
mulusandbegi
nni
ngofcont
racti
on(
about10msec).Iti
sdueto
thetimeitt
akesfrom thearr
ivalofacti
onpot
enti
alatTt
ubul
esuptothebuidupofCa2+concent
l rati
oni
nthe
cytosolt
oal ev
elenought oi
niti
ateacontr
act
ion
b) Per
iodofcont
ract
ion
 Thi
speriodi
ndi
cat
estheact
ual
shor
teni
ng(
about40msec)
.
c) Per
iodofr
elaxati
on
 Iti
stheperi
odbet weent
hepoi
ntofmaxi
mum cont
ract
ionandt
heper
iodofcompl
eter
elaxat
ion(
about
50msec).
Ref
ract
oryPer
iod
 I
tisabr
iefper
ioddur
ingwhi
chamuscl
eunder
goi
ngcont
ract
ionf
oraf
ir
stst
imul
usi
sunabl
eto
r
espondt
oasecondst
imul
us.I
tisoft
wot
ypes.
Absol
uter
efr
act
oryper
iod

 Iti
stheti
mei nt
ervalbetweentwoimpulses(
lesst
han3msec)duringwhichthemuscl
ecannotbe
sti
mulat
edevenbyst rongersti
mul
i.Thi
sisduetother
isi
ngphaseoftheacti
onpot
entalwhenNa+
i
i
onconductancetotheinteri
oroft
hecelli
shi
gh.
Rel
ati
ver
efr
act
oryper
iod

 I
tist heperi
odofr educedexcit
abil
it
ywhichrequi
resincr
easedint
ensi
tyofsecondst i
mulati
on
togenerat
eanotheracti
onpotenti
al.Iti
s char
acteri
sedby af
terdepol
ari
zat
ion stageofact i
on
potent
ialdur
ingwhichtheK+ i
onper meabi
li
tyi
sv er ghandNa+ channel
yhi sarer ecover
ingtheir

21
PHYSI
OLOGYOFMUSCULARSYSTEM 2014-
15

exci
tabi
l
ity
.
Tr
eppe:
(St
air
casephenomenon)

 Whenast i
mulusofconst antst
rengthanddur ati
oni sr
epeated(belowtetanisi
ngfrequency)onceor
twicepersecond,itcausesi ncreasedcont r
acti
onsdur ingthefi
rstfew sti
mul at
ions,whichfi
nally
reachaconst antr esponse.Thisi sduet odev elopmentofphy sicalandchemi calchangesi.
e.
reducedv i
scosit
yoft hesarcoplasm,increasedheatpr oducti
on,i
ncreasedCa2+ ionav ai
l
abil
it
yfor
bindingwithtroponindur i
ngfir
stf ew contract
ions;thesechangesar er efer
redt oasbenef i
cial
effects.
Summat
ionofMuscl
eCont
ract
ion

 Summationistheaddedef
fectofindi
vi
dualmuscl
etwi
tchest
ogetst
rongandpower
fulmuscl
e
cont
ract
ion.
 Summationoccur
sintwodi
ff
erentways
1) Byi
ncreasi
ngthenumberofmot orunitst
ocontractsimult
aneousl
y.
2) Byi
ncreasi
ngtherapidi
tyofcontr
acti
onofindi
vi
dual motoruni
tofacontracti
ngmuscle.
 Thefor
meri sref
err
edt oasmul t
i-
mot oruni
torspat i
alsummat ionand t
hel at
teras wave(
or)t
empor
al
summati
on.
Mul
ti
mot
oruni
tsummat
ion(
Spat
ial
summat
ion)

 Thest r
engt hortheforceofcont racti
onofamuscl eincreasespr ogressi
vel
ywi thincreasi
ngnumberof
contr
actingmotoruni
tsbyincreasi
ngt hestr
engthofst
imulati
on.
 Thestrengthofresul
ti
ngcontracti
onsi naskelet
almuscl
eispr oport
ionaltothesizeorthenumberofmotor
uni
tscont ract
edatt hetime,whi chi ntur
ni sdir
ectl
ypr oport
ionaltot hestrengthoft hesti
mulus;i
.e.
i
ncreasingthestr
engthofsti
mulusi ncr
easesthenumberofmot orunitsundergoi
ngcont r
action
Wav
esummat
ion(
Tempor
alsummat
ion)

 Byincr
easingthefr
equencyofsti
mulat
iontomot
oruni
ts,successi
vest
imul
iwi
l
lst
imul
ateaddi
ti
onalmot
or
uni
tsduri
ngitscont
ract
ionphase.
 Whe na mot orunitisexci ted r
epeatedl
ybya rapidsuccessionofweakstimuli
,itmayev oke
str
ongercontr
acti
on.
 Applyi
ng st
imuliinr api
d succession to a muscleincr
eases the excit
atory pot
ent
ialofthe
postsy
napti
cnerveending. Thi
si ncreasedexci
tat
orypotent
iali
softenr ef
erredtoasexcitat
ory
postsy
napti
cpotenti
al(EPSP)
.
 The second st imulus arr
ives bef
ore t he compl
eti
onof t
hefi
rst contraction which init
iat
es
thesecondcont
racti
on.Thi
sgivesgr
eaterst
rengt
horfor
ce,aswel
lasdurati
ontot healreadypr
ogressi
ng
cont
raction.

22
PHYSI
OLOGYOFMUSCULARSYSTEM 2014-
15

Fact
orsI
nfl
uenci
ngt
heSt
rengt
hofCont
ract
ion
 Si zeandt henumberofmot oruni tstimulated.
 Rapi ditywi thwhi chi ndiv
idualmot orunitofamuscl eisst
imul
ated.
 St ar l
ing'slaw:
 Thefirstf actorisgov ernedbyt hestr engthofst i
mulus.Thesecondi smostlybytherat
eofef f
ect
ivesti
muli
.
Thelasti sbyt hei niti
al l
engthoft hemuscl efi
bre.
 Incardiacmuscl e, wav elsummat ioni snotpossi blebecauseofpr
olongedactionpot
enti
al (
250msec.)al
most
compl etelyov erl
apswi thlongt wi
tch( 300m sec. ).
 Insmoot hmuscl e,theact ionpotentialisrelat
ivelyshort(
5-10msec.),hencenooverl
appingoftwi
tches,t
hus
summat ionispossi bl
e.
St
arl
i
ng'
sLawofMuscl
eCont
ract
ion
 Theener
gyofcont
racti
onofthemusclei
sdir
ect
lyproport
ional
totheini
ti
all
engt
hoft
hefi
bre.
 Thi
slawcanbeprovedbytheloadandaf
terl
oadexperi
mentont hegast
rocnemi
usmuscl
eofthef
rog.
Tet
ani
ccont
ract
ion
 I
tisthefusi
onofsuccessivet
witcheswhent hefr
equencyofstimulii
sgi
venatar
api
dratei
.e.whenaser
ies
ofacti
onpotent
ial
sisappl
iedrapi
dly,
resul
tsinpersi
stentcont
ract
ionoft
hemuscl
efi
bre
 Twotypesoftet
aniccontr
acti
onscanbepr oduced
I
ncompl
etet
etanus
 Whenthefrequencyofsti
mulati
onisr
elat
ivel
ylowi.
e.t
hesubsequentst
imul
ifal
ldur
ingt
her
elaxati
onphase
ofthepr
ecedingcontr
acti
on,ther
ewil
lbei ncompl
etef
usi
onofcont r
acti
onswhichi
sknownasi ncompl
ete
tet cl
anus( oniccont
ract
ion).
Compl
etet
etanus
 Whent hefr
equencyofst i
mul at
ionisv eryrapid,i
.e.subsequentsti
mulifal
ldur
ingthecont racti
onphaseof
theprecedingcontracti
on,therewillbecompl etef usi
onofmuscl etwi
tchesint
oonepr olongedcont r
act
ion
cal
ledascompl etetetanus(toni
ccont r
action).
 Cardi
ac musclef ibres do notf uncti
onallytetanise,si
nce,mechanicalsummat i
on i
s notpossi bl
e due
toprolongedacti
onpot enti
al(250msec) ,whichalmostcompl etel
yover
lapswithlongcontraction(300msec)
durat
ion.
Muscl
eFat
igue
 Fati
guei sthedecr easei
nt heworki
ngcapaci
tyofamuscl eortir
ednessofthemusclewheni tiscont
inuousl
y
sti
mul ated.
 Itischar acteri
sedbydi mini
shedforceofcontracti
on,increasedlatentper
iodandcont r
actionperi
odand
prolongedr elaxat
ionperi
od.Therel
axati
onisincompletei.e.
,themuscl ei
sinastateofcont r
actur
e.Att hi
s
staget hemuscl ecannotbeexcit
ednordoanymor ework.
 Inthelocomot orsyst
em ofskel
etal
muscles,
therearethreesites,whi
chareeasil
yfat
igued.
 Sy
napsesofCNS.
 Neur
omuscul
arj
unct
ion.
 Themuscl
e.
Causesoff
ati
gue
 Conti
nuousst
imul
ati
onoft hener
v efi
breorcont
inuousst
imulat
ionoft
hemuscl
ethatt
oounderischemia
(r
educedbl
oodsuppl
y)causes,
1.Lackofenergysources:Depleti
onofO2f r
om my ogl
obin;depl
eti
onofmuscl
eglycogen;
depl et
ion
ofATP

23
PHYSI
OLOGYOFMUSCULARSYSTEM 2014-
15

2.Exhaust i
on of neur ot ransmi tt
ers: Pr olongedmuscl e act ivit
ydi mi ni
shes neur ot
ransmi t
ter
acet ylcholi
neconcent rationi ntheneur omuscul arjunction,t her ebyr educest hetransmissionof
thei mpul sest omuscl ewhi chcausesdi minishedmuscl econt racti
on.
3.Accumul at
ionofmet abol icendpr oductssuchasl acticaci d:Inoxy genl ack,themuscl eglycogen
isbr okendownbyanaer obi cglycolysi
st ogi vepyruvicaci dwhi chi st henreducedt olacti
caci d
byt heact i
ono flacticdehy drogenase.
 Buti nani ntactmuscl et hebl oodsuppl yminimi zestheabov eef fectsbycont i
nuouslysuppl yi
ngthe
O2andnut r
ientstoaf unct i
oni ngmuscl e.
 Av er
yl argedr opi nATPl eveli nmuscl ewoul dmakei tnotpossi blet obr eakt hebondbet ween
my osinheadsandact inandt hemuscl ewoul dunder gor i
gor .Inpr actice,whent hemuscl eperforms
maxi mal l
yf oralongert i
me, t
heper formanceoft hemuscl edecl ineswi t
hdecl ineinATPpr oducti
on
andhence,t hemuscl ewoul dst il
lhav eadequat eATP l ev elwhent hemuscl ehasunder gone
exhaust i
on.Thusf atiguei sasaf et
ymechani sm topreventset ti
ngi nofr i
gor
Myogl obinur ia(Monday -mor ningdi sease)inhor ses:Whenawel l-
fedhor sei ssubj ect
edt ostrenuous
phy
sicalact i
v i
tyaftersev eralday sofcompl eter est,my oglobi nuriaoccur s.Dur ingheav yexer ci
se,
muscl ecel lmembr anesar edamagedandcel lcont ent
sar epar tlyreleasedi ntoblood.My ogl
obinbeing
smallerinsi zei sfil
teredt hroughgl omerulusandappear sinur inei mpar t
ingadeepr edorbr ownishred
col
ourt our i
ne–my oglobi nuria.

NEUROMUSCULARJUNCTI
ON
 Iti
sthej unct ionbet weent heter minal branchofamot orneuronandskel etalmuscl ecel l
 Themot orneur onbr anchesati tsendandeachbr anchcomesi ntoacl oseopposi ti
onbutwi t
houtcont acting
theskel etal muscl ecel l
ataspeci alisedar eacalledt heneur omuscul arjunct ionormot orendpl at
e.
 Theaxonandmuscl emembr anesar esepar atedf rom eachot herbyanar r
ow spaceof50nm wi decal led
synapticcl ef t.
 Thesy napsehasapr esynapt i
csi de, hesy
t napticcl ef tandapost synapt i
csi de.
 Thepr esy napt i
cpar toft hesy napsei sf or medbyt het erminalpor tionoft hemot orneur on;theaxonoft his
neuronext endsf r
om t heCNSt ot hemuscl ecell.
 Thecy topl asm oft r
ansmi t
ti
ngendoft heaxoncont ainsal argenumberofmembr ane-boundv esi cl
escal led
synapticv esi cleswhi chcont aint heexci tatorychemi calneur otransmi ttersubst ance–acet yl
chol ine.Each
vesi
cle cont ains about5000 t ransmi ttermol ecul es.These v esicl
es ar e clustered att he pr esynapt i
c
membr aneadj acentt othepost synapt icjunct i
onal f
ol ds.
 Thepr esy napt ict er
mi nalalsocont ainsmanymi tochondr iar equiredf ort hesy nthesisofneur otransmi tt
er
substance.
 Acetyl
chol i
nei ssy nthesisedi nthecy toplasm oft hener vetermi nalandst oredint hesy napticvesicles.
 Thesy napt iccl ef
tisf i
ll
edwi thext racellularfluidandspongyr et
icularf il
ament scal l
edbasall ami na.The
synapticcl ef tcont ainsanenzy me, acety l
chol i
nest erasewhi chi scapabl eofdest r
oy i
ngAch.
 Thepost sy napt i
cmembr ane( cellmembr aneofmuscl ecel l
)_hasaser iesofi nvaginationscalledj unctional
fol
dst hati ncr easet hesur f
acear eaoft hepost synapt icmembr ane.
 Acetyl
chol i
ner ecept orsar elocatedont hepost sy napt i
cmembr aneatt heent r
anceoft hesejunctional folds.
 Becauset heneur otransmi tt
eri sf oundonl yont hepr esynapticner vet ermi naloft hesy napse,t ransmi ssion
canoccurf rom ner v
et omuscl eonl y,andnoti nt her eversedirect i
on.

24
PHYSI
OLOGYOFMUSCULARSYSTEM 2014-
15

Tr
ansmi
ssi
onofI
mpul
seacr
osst
heNeur
omuscul
arJunct
ion
 Thef unctionoftheneuromuscul arsy napseistot ransmitanact i
onpotenti
alunidi
rect
ional
lyfr
om t hemotor
nervetot heskelet
almusclef i
bre.
 Whent heact i
onpot ent
ialreachest hener v
et er
mi nal,i
ttri
ggerstherel
easeoft heneurotr
ansmitterfr
om t
he
presynapticter
mi nal
;ther el
easedt r
ansmi t
terbindswi thther ecept
orsfoundont hepostsynapti
cmuscl e
membr aneproducinganact i
onpot entialal
ongthemuscl ecell
.
 Ana ctionpot enti
alarisesonamot orner veati t
si ni
ti
alaxonpar twhicht henspr eadsal ongthe
enti
reaxonandar r
ivesatthepr esy napticaxontermi nal
.
 Whent heactionpot enti
alreachest hepresynapti
ct erminal,
thedepol ar
isatonopensv
i ol
tage-gat
ed
Ca2+channels;andext r
acell
ularCa2+di f
fusesint
ot hepr esy
napticnervecy t
oplasm.
 Theincreasedi ntr
acell arCa2+ l
ul ev elcausesthesy napti
cv esi
clest ofusewi ththepresynapti
c
membr ane;t
hef usedsy napticvesi cleopensandr eleasestheacety l
choli
neintothesynapti
ccleft
.
 Aft
erther el
easeofAch,t hev esicleent ersbackintot hepresynapticcytoplasm t
ober efi
ll
edwi t
h
newAchf rom cy t
oplasm.
 Ther eleasedacet yl
chol i
nedi ffusesacr osst hesy napt i
cclef t,reachest hepost sy napti
cmembr ane,
andbi ndswi t
hacet y
lcholine- specificrecept or s.Thisbi ndingofacet ylcholineneur otr
ansmitterwi t
h
receptorcausesopeni ngofl igand-gat edi onchannel si nthepost synapticmembr ane.
 Sodium i onsdi ffusei ntot hemuscl ecel lthr ought hel i
gand- gatedi onchannel sandcausesl ocal
potentialchangeatpost sy napt i
cmuscl ecellmembr anecal edendpl
l atepot ent
ial.
 Thisendpl atepot ent i
al( depol arisation)opensv oltage-gat edNa+ channel satt hepostsy napti
c
membr aneandl eadst ot hegener at i
onofanact ionpot ential ont hemuscl ecellmembr ane.
 Thebi ndingofacet ylcholinewi t hit
sr ecept orr emai nsf orav erybr i
efper iod.
 Theacet y l
cholineisr apidlyr emov edf r om ther eceptor sbyt womeans.
1)Asmal lamountofAchdi ff
usesoutoft hesy napt i
cclef tandi snol ongerav ail
ablet oactont he
muscl e
2)The Ach i s dest royed by an enzy me acet ylcholi
nest erase locat ed on t he postsy napti
c
membr ane.Thi senzy mecl eav est heacet y lcholi
nei ntoacet ateandchol i
nemol ecules.
Abnor
malsi
gnalt
ransmi
ssi
onacr
ossneur
omuscul
arsy
napse

 Organophosphor usi nsecti

cidesandphy sosti
gmineinhibi
tt heactionofacet yl
choli
nester
ase.A
singlener vest imuluscanl ead to severalimpulsesand themuscl esaret hr
owni ntot et
anic
conv ulsi
ons.
 D-tubocur ari
ne,t heact i
vecomponentofcur ar
e,a plant
-derived ar
row poison used bySout h
Amer i
canI ndi
ansandt oxi
nf rom cobravenom, bl
ockstheeffectofacet y
lchol
inebybindingtoAch
recept ors.Skeletalmuscl es ofaf fect
ed animals are paraly
sed and t he ani
malmaydy e of
respiratoryarrestduet oparalysi
sofrespirat
orymuscles.

25
PHYSI
OLOGYOFMUSCULARSYSTEM 2014-
15

 Bot ul
i
num t oxi
nreleasedbysoi lbacteraCl
i ostri
dium botuli
num underanaer obiccondit
ionblocks
Achr eleaseandcausesbot ul
ism –af orm off oodpoisoning.Unhygieniccondit
ioninpreparati
onof
foodpr oducesbot uli
sm.Itcanpr oduceskel etalmusclepar al
ysisanddeat hduet or espir
ator
y
arrest.Catanddogar eresi
stanttobot ul
ism butr uminant
sar eaff
ected
 My astheni agravi
sisaneur omuscul ardisorderinwhi chautoanti
bodi esareproducedagai nstAch
receptor s.Hence,neuromusculart ransmissioni simpair
edandl eadst oparalysi
s.Thecondi ti
on
canbet reatedwithacetyl
choli
esteraseinhibi
tors
Ef
fectofTemper
atur
eint
heMuscl
e
 Ani ncreaseoft emper atureshortenst helatentandcont racti
onperiods.Thegr adientofcontracti
oni
s
i
ncreasedandhei ghtdecreased.Relaxati
onisnotmuchaf f
ected.
 A decr easeoft emper atur
ehast heopposi teef fect– latentandcont racti
onper iodar eincr
easedand
rel
axationperiodisprolonged.
 Whent emperatur
eexceedsaf ewdegr eesabov e40 C,i
rr
eversibl
echangest akeplaceint hesar
coplasm.
 Therei sami nimum t emperatureformuscl econt ract
ion,below whi
cht hemuscl ef i
br esfai
ltorespondt
o
sti
mul ati
on;butthelossofexci t
ati
oni srever
sible.Themi ni
mum t emperat
ur eisnearzero.

26