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Lec. 11

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Gener

alPhy
siol
ogy
Dr.I
ydenKamel

Ner
vesPhy
siol
ogy
Thehumancent ralner voussy stem (CNS)cont ainsabout( 100bi ll
ion)neur ons.I talsocont ains
10–50t imest hisnumberofgl i
alcell
s.Neuronscomei nmanydi ff
erentshapesandsi zesbuthav e
thesamepar ts; t
hecel lbody( soma)cont ai
nst henucl eusandi sthemet aboliccenteroft heneur on.
Neur onshav esev eralprocessescal leddendr i
test hatext endout wardf rom t hecel lbody .At y
pical
neuronal sohasal ongf i
brousaxont hatori
ginatesf rom asomewhatt hickenedar eaoft hecellbody,
theaxonhi l
lock.Thef ir
stpor ti
onoft heaxoni scal l
edt heiniti
alsegment .Theaxondi vi
desi nto
presy napticter minals,eachendi nginanumberofsy napticknobs( terminalbut tons)whi chcont ain
granul esorv esiclesf i
ll
edwi t
hneur ot
ransmitters.
Funct ionall
y,neur onsgener al
lyhav efourimportantzones:
1.Recept or(dendr it
ic)zone, wheremulti
plel ocal potenti
alchangesar eintegrated.
2. I mpul seor iginzonewher eactionpotentialsar egenerated(theinitialsegmentofaxon) .
3. I mpul seTr ansmi ssionzonewher eimpul sespr opagatedt othet erminal butt
ons.
4.Neur ot r
ansmi tt
erSecr eti
onzone, wheret het erminalbuttonsreleaset ransmi tt
ers.

Theaxonsofmanyneur onsaremy el
inat
edwi thasheathofmy el
i
n,apr otei
n–li
pidcomplexthati
s
wrappedaroundtheaxon.Intheper i
pheralnervoussy
stem,myeli
nformsbyaSchwanncel l
,whil
e
thoseinCNSar emyeli
natedbyOli
godendr ocy
tes.Themyeli
nsheathenvelopstheaxonexceptatit
s
endingandatthenodesofRanvier
.Notallneuronsaremyeli
nat
ed;somear eunmy el
inat
ed.

EXCITATION&CONDUCTI ON
Nerve cell
s haveal ow thresholdf orexcitati
on.The st i
mul us maybe el ect
rical,chemical,or
mechanical.Two t ypes ofphy sicochemicaldisturbances ( potenti
als)are pr
oduced:l ocal
,non-
propagated potenti
als known as el ect
rotonic potenti
al;and pr opagat
ed potential
s,the action
potenti
als(ner
vei mpulses).Thesear etheonl yelectr
icalresponsesofneur onsandot herexcit
able
ti
ssues,andt heyar ethemai nl anguageoft henervoussy stem.Theyar eduet ochangesi nt he
1
Gener
alPhy
siol
ogy
Dr.I
ydenKamel
conduct
ionofi
onsacr
osst
hecel
lmembr
anet
hati
spr
oducedbyal
ter
ati
onsi
nionchannel
s.

Inneurons,theconcent r
ationofK+i smuchhi gheri nsidethanout si
det hecel l
,whi lether everseis
thecasef orNa+.Ther efore,thereisast rongt endencyf orpot assium ionst odiffuseout wardt hrough
themembr aneandcar ryposi ti
veel ectri
calchar gest otheout side,t huscr eatingel ect
ro-positi
vit
y
outsidethemembr aneandel ectro-negati
vityinsi de.Thispot enti
aldi ff
erencebet weent heinsideand
outsideisabout( -94mi l
li
volts)inthemammal ianner vef i
ber.
Ont heotherhand, dif
fusionoft heposi t
ivelychar gedsodi um ionst othei nsi
decr eatesamembr ane
potenti
alwi t
hnegat ivi
tyout sideandposi ti
vityi nsidet hener vef i
ber,thispot ent i
alisabout( +61
mill
ivol
ts).Ther esultantoft heset wopot entialsi sknownasr estingmembr anepot ent
ialwhi chis
about(-70mv )because:
1.Themembr anei smor eper meabl etoK+t hant oNa+.
+ + +
2.TheNa–K pumpt ranspor tsthreeNa outoft hecel l
forev erytwoK+.
3.Thecel lmembr anei si mper meabl efort hel eakageofpr oteinmol eculesf rom insidet oward
outsi
de, whichar eani onint heirnature.

Measur
ementofmembr
anePot
ent
ial
:

Thecat hoderayoscil
l
oscope( CRO)isusedtomeasur etheelectr
icaleventsinl
ivi
ngtissuebecause
theel ectri
calevent
sinneur onsar erapi
d,bei
ngmeasur edinmi l
li
seconds( ms);andt hepotential
changesar esmall,
beingmeasur edinmill
i
volt
s( mV).Whentwoel ectr
odesar econnectedthrougha
suitableampl i
fi
ertoaCROoneel ectr
odeisplacedont hesur f
aceofasi ngl
eaxon,andt heother
electrodeisinsert
edintotheinteri
orofthecel
l,aconstantpotenti
aldif
ferenceisobserved,wit
ht he
i
nsidenegat i
verel
ati
vetotheout si
deofthecel
l atr
estwhichisusuall
yabout–70mV.
Act
ionPot
ent
ial
(ner
vei
mpul
se)
:
I
ftheaxonissti
mul at
edaser
iesofpot
enti
alchangesi
sobser
vedandknownast
heact
ionpot
ent
ial
.
Thesuccessi
vestagesoft
heacti
onpotent
ial
areasfol
l
ows:
1.st
imul
usar
ti
fact
:
Ir
regulardef
lect
ionoft hebasel
ineoccur
sduet
ocur
rentl
eakagef
rom t
hest
imul
ati
ng
elect
rodestotherecor
dingel
ect
rodes.
2.Rest
ingSt
age(
Lat
entper
iod)
:
Thisisther
esti
ngmembr anepotent
ialbefor
et heacti
onpot ent
ialbegins.Themembraneis
sai
dt obe“polar
ized”duringthi
sstagebecauseoft he( -
70milli
volt
s)negativ
emembr ane
potenti
alpr
esent
.Thisstagei sduetot heti
mer equiredfortheimpulset otr
avelal
ongthe
axonfrom t
hesit
eofst i
mul at
iont
other ecor
dingelectrodes.
3.Depol
ari
zat
ionSt
age:
Att hi
st i
me,themembr anesuddenlybecomesv erypermeabletosodium ionsthatdi
ffuseto
theinteri
oroft heaxonandt hepotenti
alri
singr api
dlyinthepositi
vedirecti
on.Thepointat
which thischange inrate occursiscall
ed the f i
ri
ng lev
elorsomet i
mest he t
hreshol
d.
Thereaft
er,the greatexcess ofpositi
ve sodium ions mov ing t
ot he insi
de causes the
membr anepot enti
alt
oactuall
y“ ov
ershoot
”bey ondt hezerolevelandtobecomesomewhat
positi
ve(+35mV) .
4.Repol
ari
zat
ionSt
age:
2
Gener
alPhy
siol
ogy
Dr.I
ydenKamel
Thesodi um channel
sbegint ocl
oseandthepot
assium channel
sopenmorethannor
mal.
Then,rapiddif
fusi
onofpot assi
um i
onstotheexter
iorre-
establ
i
shesthenor
malnegati
ve
rest
ingmembr anepotent
ial
.

Theshar priseandrapi
df allar
et hespikepotenti
aloftheneuron,andtheslowerfallattheendof
the process isthe af
ter-depolar
izat
ion.Afterreachi
ng the previ
ous r
esting l
evel,the t
raci
ng
overshoots sli
ght
lyint he hy perpol
arizi
ng di
recti
on tof or
m t he smallbutpr olonged aft
er-
hyperpolar
izat
ion.

Pr
oper
ti
esofAct
ionPot
ent
ial

1.Al
lorNoneLaw:

Two f act or
sar ei nvol
ved inpr oducing acti
onpot ent i
al;thei ntensi
tyofst i
mul usand i ts
duration.Oncet hresholdintensi
tyisreached,afullacti
onpot entialisproducedandanyf urther
i
ncreasesi nthei ntensi
tyofast i
mul uspr oducenoi ncrementi nt heactionpot enti
al,such
sti
mul ical l
edthresholdst i
mul i
.Howev ertheactionpot enti
alf ail
st ooccuri ft
hest imulusi s
subthr esholdinmagni tude,theref
oreact i
onpot entialobeyt heal l-
or-
nonel aw.Sl owlyrising
current sfailt
of i
ret hener vebecauset hener veadapt st ot heappl i
edst imulus,apr ocess
cal
ledadapt at
ion.

2.Ref
ract
oryPer
iod:

Dur
ingtheacti
onpotent
ialt
heneuronisnotrespondt
of ur
thersti
mul
ati
on.Thi
sref
ract
ory
per
iodi
sdiv
idedint
oanabsolut
eref
ract
oryper
iod(ARP)
,corr
espondi
ngt
otheperi
odfr
om the

3
Gener
alPhy
siol
ogy
Dr.I
ydenKamel
ti
met hef ir
inglevelisreachedunt ilr
epolari
zat
ionisaboutone-thi
rdcomplete,andar elat
ive
ref
ractoryper i
od(RRP) ,andlastingfr
om t hi
spointtothestar
tofaf t
er-
depol
arizati
on.During
theabsoluter efr
actoryperiod,nost i
mulus,nomat terhow str
ong,wi l
lexci
tet henerve,but
duri
ngt herelati
verefr
actoryperiod,st
rongerthannormalsti
mulicancauseexcitati
on.
3.PropagationofAct i
onPot enti
al:

Thenerv
ecellmembraneispol
ari
zedatrest,wi
thposit
ivechar
gesal
ongtheout si
deofthe
membraneandnegat
ivechar
gesalongt
hei nsi
de.Dur
ingtheacti
onpotent
ial
,thi
spolar
it
yis
abol
i
shedandfora

bri
efper iodisact uallyreversed.Suchel ectrotonicdepolarizati
oni nit
iat
esal ocalr esponse,
andwhent hefi
ringl evelisreached, apr
opagat edr esponseoccur st hatintur
ndepol arizest he
membr aneinfrontofi t
.Anaxoncanconducti nei t
herdirection.Whenanact i
onpot entialis
i
niti
atedi nthemi ddl eofi t
,twoi mpulsesar et ravel
inginopposi tedi r
ecti
ons,impul sespass
al
ongaxonst ot heirtermi nati
onar ecalledor thodromic,whi lethosepassi ntheopposi te
di
rectionarecalledant i
dromi c.Becausesy napsesper mitconduct ioninonedi r
ectiononl y ,an
anti
dromi cimpulsewi llfai
ltopasst hefir
stsy napset heyencount eranddi eoutatthatpoi nt .
4.TypesofI mpulseConduct ion:

Thedi stri
buti
onofionchannelsal
ongt heaxonplaysakeyroleintheini
ti
at i
onandregulat
ion
oftheact i
onpotenti
al.ThenumberofNa+channel spersquaremi cr
omet erofmembr anein
my el
inatedmammal ianneuronshasbeenest i
matedt obe50–75int hecellbody
,350–500i n
theinit
ialsegment,lessthan25ont hesurfaceoft hemyeli
n,2000–12,000att henodesof
Ranvier,and20–75att heaxont er
mi nal
s.Alongt heaxonsofunmy el
inatedneurons,the
numberi sabout110.Therearetwotypesofimpulseconducti
on:

1)Cont
inuousConduct
ion

I
toccursinunmyel
inat
edaxons,inwhichcircul
arcur
rentf
low ofact
ionpot
ent
ial
pr
opagat
edal
ongall
pointsofaxonmembrane.

2)Sal
tat
oryConduct
ion

I
t occurs i
n
my eli
nated axons,
i
n whichcur rentflow
j
umps from onenodeof
Ranv i
ertot henext.
Itisa rapidprocesst hat
all
ows my eli
nated axons
to conductup t o 50
ti
mes faster t han t he
fastest
unmy eli
nated
f
iber
s.

4
Gener
alPhy
siol
ogy
Dr.I
ydenKamel

5.Vel
oci
tyofConduct
ion:

Sev
eral
fact
orsar
einv
olv
edi
ndet
ermi
ningt
hev
eloci
tyofner
vei
mpul
seal
ongt
hener
vef
iber
s:

1)My
eli
nat
ion,
myel
i
nat
edf
iber
sar
efast
ert
hanunmy
eli
nat
ed.

2)Di
amet
er,
thi
ckf
iber
sar
efast
ert
hant
hin.

3)Spi
keDur
ati
on,
fiber
swi
thl
essspi
kedur
ati
onar
efast
ert
hant
hosewi
thl
ongdur
ati
on.

4)Absol
uteRef
ract
oryPer
iod,
fiber
swi
thl
essARPar
efast
ert
hant
hosewi
thl
ongARP.

I
ngener
alner
vef
iber
scanbecl
assi
fi
edi
ntot
hreemaj
ort
ypes:

Ty
pe-
Aner
vef
iber
s:

Theyaremy el
i
natedfi
berswi
thl
argedi
ameterandless(
spikeandARPdurati
on)
,ther
efor
e
theyarefastfiber
swi t
hav el
ocit
yabout15-120m/ s(e.g.somat
ictouch& pressur
e
sensat
ion)
.

Ty
pe-
Bner
vef
iber
s:

Theyaremy el
inat
edfiber
swi t
hsmallerdi
ameterandr
elat
ivel
ylonger(
spi
keandARP
dur
ati
on),ther
eforetheyaremoder atefi
ber
swi thav el
ocit
yabout3-15m/ s(
e.g.
vi
scer
altouch&pressuresensati
on)
.

Ty
pe-
Cner
vef
iber
s:

They are unmyeli


nated f
iber
s wit
h small
estdi
ameterand l
argest(spi
ke & ARP
durat
ion)
,theref
oret heyareslow fi
ber
swi t
h avel
oci
tylessthan 3 m/s(pai
n&
temperat
uresensati
on).
5
Gener
alPhy
siol
ogy
Dr.I
ydenKamel
Inadditi
ont ov ari
ationsinspeedofconduct i
on,thevariousclassesoff i
bersinperi
pheral
nerves dif
feri nt heirsensit
ivi
tyt o hypoxia,pressur
e and anest het
ics.Localanesthetics
depress transmi ssion i
nt he group(C)f i
bers beforetheyaf fectgr oup (A)touch fi
ber s.
Conv er
sely,pressur eonaner vecancausel ossofconduct i
oninlarge-diametermotor
,touch,
andpr essuref iberswhi l
epainsensat i
onr emainsr el
ati
velyint
act.Pat t
ernsoft hi
stypear e
somet i
messeeni ni ndi
vi
dual
swhosl eepwi ththeirar
msundert heirheadsf orl
ongper i
ods,
causingcompr essionofthenervesinthear ms( Sundaymor ni
ngparalysis).

6.CompoundAct
ionPot
ent
ial

Peripheralnervesi nmammal saremadeupofmanyaxonsboundt ogetherinaf ibr


ous
env el
opecal l
edt heepineurium.Thet hr
eshol
dsofthei ndi
vidualaxonsinthener veandt hei
r
distancefrom thest i
mulati
ngel ect
rodesvary.Whent hest i
mul iareofthresholdi ntensi
ty,
axonswi thlowt hreshol
dsf i
reandasmal lpot
enti
alchangeisobser ved.Astheintensityofthe
stimulat
ingcur r
enti sincreased,theaxonswithhigherthresholdsar eal
sodi scharged.The
stimulusthatproducesexci tat
ionofallt
heaxonsiscal
ledthemaxi malst
imulus.

Sy
napt
icTr
ansmi
ssi
on
Impulsesar etransmi ttedf r
om onener vecelltoanot hercellatsy napses.Thesear et hejunctions
wher etheaxonofonecel l(presynapticcell
)terminatesont hedendr i
tes,soma,oraxonofanot her
neuronormuscl eorgl andcel l(postsynapti
ccell
).Transmissionofi mpulsesacr osssy napsescanbe
classi
fied int
ot wo t y pes;el ect
ricaland chemi caltransmi ssion.I n electri
calsy napses,t he
membr anesofthepr esy napticandpost synapti
cneur onscomecl oset ogether,andgapj unctionform
betweent hecel l
st hroughwhi chi onspass. Themostsy napti
cj unctionsischemi calinwhi cht he
i
mpul sei nthepr esynapt i
caxoncausessecr et
ionofaneur otransmi tt
ersuchasacet yl
choline.This
chemi calmediatorbindst oreceptorsont hesurfaceofthepost synapticcell
,andt hi
st ri
ggersev ents
thatopenorcl osechannel sint hemembr aneoft hepost synapticcel l
.Theef fectsofdi schargeat
i
ndi v
idualsynapticendi ngscanbeexci tat
oryorinhibit
ory:
1. Exci tatorySynapses:
Inwhi cht heneurot
ransmit
terr
eleasedbypresy
napticmembr
aneopenssodi um channel sinthe
postsy napticmembr aneandproduceact
ionpotent
ial.
2.Inhibitory Sy napses:
In whi ch t he neurotr
ansmi tt
ers
released f rom presynaptic membr ane
opens potassium or chl ori
ne
channel s that maket hepot enti
alofpost
synapt ic membr anef arawayf r
om
fir
ingl eveland fai
lt o pr oduce act ion
pot ential
.

6
Gener
alPhy
siol
ogy
Dr.I
ydenKamel

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