1

LECTURE2
Functionaldescriptionofmuscl es.Mechanismof muscl e
contraction.
Role of skeletalmuscul ature.Mechani sm of muscul ar contracti on.
Electromechani csinterface.Ki nds( i
sometri c,i
sotonic,auxotonic)and
modes( singl
e,tetani c)ofmuscul arcontraction.Periodsofasi ngle
muscl econtraction.Termsofori gi
nofcompl eteandincompl etetetanus.
Optimumandpessi mum ( Vvedensky) .Rhythmiccontractionofmuscl esin
theorgani smoftheman. Workandforceofmuscl es.Fatigue. Physiology
featuresofasmoothmuscl es.
Lecturedemonstrati ons:thecomputermaki ngofani matedcartoon
is"Mechanismofmuscul arcontracti on".
Can not be any movi ng wi thout muscl e contracti on.Muscl e
contractionischangeofl engthandtoneofamuscl e.
KINDSOFMUSCLECONTRACTI ON
1.Isometri ccontracti on–i sthecontracti onthatgeneratef orce
(tensi
on)wi thoutshorteni ng( oneofthephasesofcardi accycl e;
orwhensomebodytri estoliftaveryheavyl oad).
2.Isotoniccontracti on–i sthecontracti onwithoutchangeofthe
muscletone.
3.Auxotoni c(mixed)contracti on–thecontracti onwhenthel ength
andthetonechange. Iti
sthemostspreadki ndofthecontracti on.

Muscl estructureandf i
laments
Eachmuscl ef i
beri smul tinucl
eateandbehavesasasi ngleunit.It
containsbundlesofmyof ibri
ls,surroundedbysarcopl asmicreticul
um.
Eachmyof ibrilcontainsinterdigi
tatingthickandthi nfil
amentsthat
arearrangedl ongitudinallyi
nsarcomeres. Repeati ngunitsofsarcomeres
accountf ortheuni quebandingpatterni nastri atedmuscl e.Asarcomere
runsfromZl i
netoZl ine.
Structure-f unctionalunitofamuscl econtracti onisasarcomer.
Z sarcomer Z

sarcopl
asma
Ca2+
Ca2+

1.Thickfil
aments
containmyosi
n.Eachmyosinmolecul
ehastwo"heads"attachedtoa
si
ngl
etail.Themyosi
nheadsbindATPandacti
nandareinvolvedi
ncross-
 2

bri
dgeformati on.
theyarepresenti ntheAbandi nthecenterofthesarcomere.
2.Thinf ilaments
containacti n, tropomyosi nandtroponi n.Troponi nistheregul atory
2+
proteinthatpermi tscross-bri dgeformati onwheni tbindsCa .
TheyareanchoredattheZl ines.
Theyarepresenti ntheI bands.
Theyinterdi gitatewi ththethi ckfil
amentsi naporti onoftheAband.
3.Ttubul es
areanextensi vetubul arnetwork, opentotheextracel l
ularspace,
whichcarrythedepol arizationtoth ecel li
nteri or.
Theyarel ocatedatthej unctionsofAbandsandI bands.
4.Sarcopl asmati creti culum( SR)
istheinternaltubul arstructurethati sthesi teofCa2+storageand
releaseitf orexci tation-contracti oncoupl ing.
Ithastermi nalci sternaethatmakei ntimatecontactwi ththeT
tubules.
Theirmembranecontai nsCa2+-ATP-ase( pumps) ,whichtransport
2+
Ca f rom i ntracel lular f luid into the SR i nteri
or,keepi ng
2+ 2+
intracellul ar[ Ca ]l ow.Wi thinthe SR,Ca i sboundl ooselyto
calsequestri n,toberel easedupondepol arizati on.

B.Stepsi nexci tation-contracti oncoupli

ngi nascel etalmuscl e
1.Acti on potenti al s in the muscl e cel l membrane i ni
tiate
depol arizationoftheTtubul es.
2.Depol arizationoftheTtubul esopensCa2+ channel sinthenearby
2+
SR,causi ngreleaseofCa f romth eSRi ntothei ntracel lularflui
d.
2+ -8
3.I ntracel lular[ Ca ]i ncreases f rom 1 0 mmol /literto 1 0-5
mmol /liter.
2+
4.Ca bi nds to troponi n C on the thi nf ilaments,causi ng a
conf ormati onalch angeintroponi n.
a.Tropomyosi nismovedoutofth ewaysothatthecross-bri dge
cyclecanbegi n.
b.Acti nandmyosi nhaveaf fini
tytoeachother. Theybi nd,theheads
ofthecross-bri dgespi vot, thethinandthi ckf il
amentssl i
deover
eachother, andATPi shydrol i
zed.
c.Subsequentl y,thecross-bri dgesbreakandanewmol eculeofATP
bindtothemyosi nheadsothatanewcycl ecanbegi n.
2+
d.Cross-bri dgecycl i
ngconti nuesasl ongasCa i sboundtotroponi n
C.
Sarcomercanbecomeshorterf or40%ofi tslength . About50strokes
takepl acef orit.
5.Rel axati on occurs when Ca2+ uptakes i nto the SR l ower the
 3

i
ntracell
ular[Ca2+].ATPisconsumedi ntheprocessofCa2+uptake(as
wellasduringthecross-bri dgecycle).Systemacti
n-myosinbecame
2+ .
i
nactive.
Forrel axationalsonecessaryionsMg andprotei nfactor
Marsha-Bendola.

SI
NGLECONTRACTI
ON

Amuscl ereactstoani ndivi

dualstimul
uswi thasingl
econtracti
on
(onestimulus–onecontracti on)
.Singl
econtractioni
sdivi
dedi
ntothree
phases:
1.alatentperiodofcontraction(0,
01c)
;
2.contractionphase(0,
04c);
3.relaxati
onphase( 0,
05c)

1 2 3
0,
1c

Theintervalofti mef rom themomentofsti mulationuptothe

begi
nningofcontractionisthelatentorhiddenperi
odofcontracti
on.
Theintervalofti mef rom thebeginningofcontractionuptothe
maximalcontractioni
sthecontracti onphase.
Theintervaloftimefromthemomentofmaxi malcontracti
onupto
thetotalrelaxati
onistherelaxationphase.

SUMMATIONOFCONTRACTI
ONSANDTETANUS
Intenseandconti
nuouscontractionasaresultofrhythmicf
requent
stimulation isknownasatetaniccontracti
onortetanus.
Fortetaniccontractionintervalsbetweenthesti mulusmustbe
lessthanasi ngl
econtraction'
sdurati on(bef
oremuscl ehasrelaxed
afterthef i
rstcontracti
on).

Compl
eteandincompl
etetetanus.
I
fthesecondsti
mulusisappl
iedwhenthemusclehasal
readybegan
 4

torelaxafterthef i
rstcontracti
on,thepeakofthesecondcontraction
onthemyographicalcurvewil
lbeseparatedf romthatofthef i
rstbythe
sli
ghtsagofthecurve.Thisi
sincomplete(clonus)tetanus.
I
fthesecondsti mulusisappli
edbef orethef i
rstcontracti
onhas
reachedi
tspeak,thecompletetetanuswillbef ormed.

Mechanismoftetanus.
I
fthemuscl eisstimul
atedrepeatedly,thenmoreCa2+ isreleased
2+
from theSR,thereisagreaterriseinintracel
lul
ar[Ca ] ,andthereis
greatercross-bri
dgeformation.
Asaresul t,moretensi
oni sgenerated
bythemuscle(tetanus)
.

OptimumandPessi mumoff orceandf requencyofsti mulation.

Thesephenomenaweredi scoveredbyVvedensky.I twasf oundthat
theef fectoftwosuccessi vestimul iwasnotequaltotheari thmeti cal
sum ofi ndivi
dualcontracti onsatal l
.Itwas, asarul e,ei
therl argeror
smal l
erthanthatsum whi chindicatedthatthecapaci tyf oranew
contraction af ter each precedi ng i mpulse of exci tation changed
considerably.Itwasexpl ainedbyVvedenskyf romposi tionsofthetheory
aboutthephasesofexci tabili
ty.
If each next sti mulus wi llact at the supernormalphase of
excitabil
ityf romth epreviousexcitati on-ampl i
tudeofthetetanuswi l
l
bemore, thenexpected. Suchphenomenoni sknownas opti mum.
Ifeachnextsti mul uswi l
lactatthesubnormalphaseofexci tabil
ity
fromtheprevi ousexcitation-ampl itudeofthetetanuswi l
lbel ess,then
expected. Suchphenomenoni sknownas pessi mum.

Motoruni ts.
Eachmotornervef ibrei saprocessofthemotorcel lsofthe
anteriorhornsofthespi nalcordservesawhol egroupofmuscl efibres
ratherthanone.Thesegroupsareknownasmotoruni ts.Inmanthe
numberoff ibresmakingupauni tvari esbetween1 0and3000i nthe
dif
ferentmuscl es.Thesmal l
estnumberi scontai
nedinthemotoruni tsof
the fast muscl es with whi ch the most accurate movements are
performed.F or.
ex.
:intheocul armuscl esandthoseofthef i
ngersthe
motoruni tsarecomposedoftentotwenty-f ivemuscl ef i
bres.In
contrast,therelativel
ysl owmuscl esinvolvedinadjusti
ngbodyposture,
whichdonotrequi restri ctcontrol ,consistofmotoruni tscompri si
ng
between2000and3000f ibres.
 5

Al
lmusclesfibresofthesamemotoruni thaveequalexcitabi
lityand
contracttogether.Increasi
ngstrengthofstimulil
eadtoi ncreasingof
musclecontractionstrength( di
rectdependencebetweenthemi ni
mal
andmaximalthreshold).

nerve
f
ibre

Muscul
arf
ibres

Maxi
malthreshol
dismini
malstrengsofsti
mulati
on,
thatcauses
maximalcontracti
onofthemuscl
e.

MUSCULARWORKANDSTRENGTH
Thestrengthofamuscl eismeasuredbywei ghi
ngthemaxi muml oad
itiscapableofli
fting.Thismaxi mumstrengthmaybeverygreat.
Themaxi mum l oadamuscl ei scapabl eofl i
ftingisdividedbythe
numberofsquarecenti metresi ni tsphysiologicalcross-section.Inthis
waytheabsol utemuscul arstrengthi scalcul ated.
Theworkofmuscl esi stheproductofthel oadliftedandthedegree
ofcontracti onandi sexpressedi nki l
ogramme-metresorgramme-
centimetres. A=mgh
Examples:absol.strengthi nkgpersquarecenti meters:ofthearm
flexor8,1
;ofthebicepsofthearm1 1,
4;th
etri cepsofthearm1 6,
8.
Inordertocomparethestrengthofdi fferentmuscl es,themaxi mum
loadamuscl eiscapabl eofl i
ftingisdividedbythenumberofsquare
centimetresinitsphysi ologicalcross-section.
Thegreaterthephysi ol
ogicalcross-secti onofamuscl e,i
.e.thesum
totalofthecross-secti onofal litsf i
bres, thegreaterthel oaditcan
li
ft.

F
ATI
GUE
Fatigueisatemporaryreducti onoftheworki ngcapacityofacel l
,
organororgani sm asawhol eresultingfrom prol ongedexertionand
passingafterarest.
Fatiguei
nmuscl eisbroughtaboutbytwopri ncipalcauses:
1.
One is the accumulation of metabolites i n the muscle during
contraction(inparticul
ar,oflacticacidthati sf ormedduri ngglycogen
breakdown) ,
whichproducesadepressi veeffectontheworki ngcapacity
 6

ofthemuscl ef i
bres.Partoftheseproducts, andpotassi umi ons,diffuse
from thef i
bresi ntotheperi cell
ularspaceandexert ani nhibitory
influenceonthecapaci tyofthemembranetogenerateacti onpotenti al
s.
2.Thesecondcauseofprogressi vef atiguei sgradualexhausti onof
theenergyreserveofthemuscl e.
Themyoneuralj unctionbecomesf ati guedmuchearl ierthanthe
muscl ef i
bres,andi nconsequence the muscl eisprotectedagai nst
exhausti on due to prol onged exertion by the bl ocking of i mpul se
transmi ssionfromnerve; andintheorganismaswhol ethenervecenters
tireevenearl ierthanthemyoneuraljunctions.
Thef ati
gueofnervecentersmaybecausedbyasharpf allinthe
reserves of synthesi zed mediator in the nerve endi ngs,di minished
sensi tivityofthepostsynapti cmembranetothemedi ator,orareducti on
ofitsenergyreserve.
Sechenovmadethetheoryabouttheacti verest.Sechenov( 1903)
firstshowedthatrecoveryoftheworki ngcapaci tyoff atiguedmuscl es
inthehumanarm af terprolongedexerti oni nlifti
ngal oad,markedl y
quickenedi fworkwasperf ormedwiththeotherarmduri ngtheperi odof
rest.

SMOOTHMUSCLES

I
nthehi gherani malandmansmoothorunstri atedmuscl esaref ound
inthevisceralorgans, bl
oodvessel s,andski
n,andarecapabl eofrel ativel
y
slowmovementsandprol ongedtoniccontraction.
Structure
- has thi ck and th inf il
aments,but they are not arranged i n
sarcomeres; theref ore,
theyappearhomogenousratherthanstri ated.
Typesofsmoothmuscl e
1.
Multi-unitsmoothmuscl e(eachf i
berhasindependentinnervation) .
-i spresenti nthei ris,
cill
iarymuscleofthel ens,
andvasef f erens;
- behavesl i
keseparatemotoruni ts;
- hasl i
ttl eornoel ectricalcoupli
ngbetweencel ls;
-i sdensel yinnervatedandcontracti onisunderneuralcontrol .

2.
Uni
tary(single-unit)smoothmuscl e
-i s the most common type and i s present i
n the uretrus,
gastrointestinaltract,ureter,andbl
adder;
-i sspontaneousl yactive( slow waves)andexhibits"pacemaker"
activity,whi
chi smodulatedbyhormonesandneurotransmi tters;
- hasahi ghdegreeofel ectricalcoupl
ingbetweencell
s" syncytial
structure"sothatcoordi natedcontractionoftheorgancan
occur( e.
g.bladder)
.
 7

3.
Vascularsmoothmuscle
hasproperti
esofbothmul
ti-uni
tandsi
ngl
e-uni
tsmoothmuscl
e.

Physiologicalcharacteri sticsofsmoothmuscl es
1.
Thepl astici tyofsmoothmuscl eiscapaci tytoretai nitsl ength
uponextenti onwi thoutachangei ntensi onunti lactivecontracti onis
inducedbysti mul ation.
Plasticityisaveryi mportantpropertyf orthenormalf unctioningof
thesmoothmuscl esofthewal l
sofhol low organs, forexampl e,ofthe
urinarybl adder; asaconsequenceofthepl asticityofi tswal lspressure
changesi nsidei tarerel ati
velysl i
ghtwhatevertheamountofuri neit
contain.
2.Excitabi l
ityandsti mulationofsmoothmuscl es.
Smooth muscl es are less exci table than skel etalones;thei r
stimulationthreshol dsarehi gherandthechronaxi ei
sl onger.Theacti on
potentialsofmostsmoothmuscl efibresareofsmal lampl i
tude( ofthe
orderof60mVasagai nst1 20mVi nskeletalfibres)andofl ongdurati on,
it
certainmuscl es,between1and3seconds.
Theref ractoryperi odisveryl ongandi tl aststhroughoutthatof
theacti onpotenti al,i
.e.Foroneto3seconds.Thespeedofi mpulse
transmi ssionvari eswi thdi fferentf ibresf rom af ew mi ll
imetresto
severalcenti metrespersecond.

-60

-80
0 1 2 3 4
seconds

Contracti
onofsmoothmuscl
e

-Themechani
sm ofexci
tati
on-contracti
oncoupl
ingi
sdi
fferent
 8

fromthatwhi choccursi nskeletalmuscl e.

Asi ngl
esti mulusofgratestrengthcangi verisetothecontraction
ofasmoothmuscl e.Thel atentperi odofthi ssi ngl
econtractionis
considerablyl ongerthanthatofskel etalmuscl e,beingasmuch ,for
exampl e,as0,
25to1 ,0secondinthei ntesti nalmusculatureofarabbi
t.
Thecontracti oni tselfi
sal sol ong,lastinguptof i
vesecondsinthe
stomachofarabbi t,andonemi nuteormorei nthatofaf rog.The
relaxationphasef ollowingcontracti onisparticularlyslow.

Stepsi
nexci
tati
on-contracti
oncoupl
ingi
nsmoothmuscl
e

- Therei snotroponi n;insteadCaregul atesmyosi nonthethi ck

filaments.
1.Depol arizationofthecel lmembraneopensvol tage-dependent
2+ 2+
Ca channel sandCa f lowsi ntothecel ldowni tsel ectrochemical
2+
gradi ent, i
ncreasingthei ntracellular[Ca ] .
2+
2.TheCa thatentersacrossthecel lmembranemaycauserel ease
2+ 2+ 2+
ofaddi tionalCa f romtheSRth roughCa –gatedCa channel s.
2+
Hormonesandneurotransmi ttersal sorel easeCa f rom theSR
2+
throughV-gatedCa channel s.
2+
3.Intracel lular[Ca ]i ncreases.
2+
4.Ca bi ndstocal modul i
nandtheC a2+-cal modulincompl exbi
ndsto
andacti vatesmyosi nlight-chainkinase.Whenacti vated,myosin
light-chai nkinasephosphoryl atesmyosi nandal l
owsi ttobindto
acti n.Shorteningf ollows.
5.Dephosphoryl ationofmyosi nproducesrel axation.

Automati smoftheunstri atedmuscl es.

Unstriatedmuscl esofthestomach,i ntestine,gall-blader,ureter
andothers haveacapaci tyforspontaneous,automaticacti vity.
Thisautomati sm i sofmyogeni cori gi
n.Thef oll
owi ngspontaneous
variationsi nmembranepotenti al
saredi sti
nguishedinunstri atedmuscle
fibres:1.slow depolari
zati onwaveswi thacycl eoftheorderofseveral
mi nutesandanampl itudearound20mV;2.F astsl i
ghtvari ati
onsof
potenti alproceedi ng the appearance ofacti onpotenti als;3.Action
potenti als.
Anotherf eatureofthesemuscl esistheirhighsensitivitytocertain
chemi calagents,i nparti cul
artotheacetyl choli
ne,secretedi nthe
endings ofthe parasympatheti c nerve f i
bres,to the noradrenal i
ne
producedbythemedul laoftheadrenalgl andsandtheendi ngsofthe
sympatheti c nerve f ibres,and to a number of other substances
(histami ne,serotonin)
.