BIO2A03

IntegrativePhysiologyofAnimals
Lecture20
March 1
st
, 2013 March1 ,2013
Dr.Nikol Piskuric
Propagation of action potentials Propagationofactionpotentials
Gradedpotentials
Actionpotential
WhenanAPisinitiatedattheaxonhillock(triggerzone),it
producesacurrentthatspreadstoadjacentareasofthe
membrane,causingawaveofdepolarization
Electrotonic conduction:passivespreadofvoltagechanges
alonganeuron(occursforGPsandAPs)
APpropagationinan
l d unmyelinated axon
Summation of GPs at the SummationofGPsatthe
axonhillockcreatesadepol
arization inthatregion
ThepolarityoftheV
m
reverses
insideofthecellbecomes
morepositivethattheoutside
of the cell ofthecell
The positive charges inside Thepositivechargesinside
thecellareattractedto
neighbouring negative
charges and current spreads charges,andcurrentspreads
Depolarizesadjacentregions
APpropagationinan
l d unmyelinated axon
Theneighbouring regionis
nowdepolarizedenoughto
generateanAP
Theprocesscontinuesdown
theaxon,aspositivecurrent
movesfromoneregionofthe g
axontothenext
Eventhoughpositivecurrent g p
spreadsinbothdirectionsfrom
thedepolarizedregion,theAP
cannot move backward because cannotmovebackwardbecause
thepreviousregionisinthe
absolute refractorystate
VoltagegatedchannelsinAPpropagation
t 1
APsarepropagated
whereas GPS are not
t =1
whereasGPSarenot
becauseAPscausethe
regenerativeopeningof
voltagegatedNa
+
channels g g
(thus,thereisnodecre
ment ofpositivecurrent
flowdowntheaxon)
t=2
APsarenotpropagated
backwardsbecausethe
previousregionisrefractory
(Na
+
channelsinactivated
andK
+
channelsopen)
t=3
Purves etal.2004
Mechanisms that increase AP conduction MechanismsthatincreaseAPconduction
1. Increaseaxondiameter
DecreasesresistanceoftheICFintheaxon
currentflowsalongthepathofleastresistance
2. Wraptheaxoninaninsulatinglayertopreventcurrent
leakage leakage
Increasesresistanceofcurrentflowacrosstheplasma
membrane
Accomplishedbymyelination
Thus,thefastestconductingaxonsarelargeandmyelinated
Myelinated axonspropagateAPsvia
l d saltatory conduction
Oligodendrocytes (CNS) APsareratherslow
andSchwanncells(PNS)
insulate>99%ofthe
distancealongtheaxon,
h d f
eventsinvolvingdiffusion
ofions.However,APscan
leapalmostinstant
l f d
Schwanncell
whereasNodesof
Ranvier occupy<1%of
thetotaldistance.
aneously fromonenode
tothenextbysaltatory
conduction(saltarein
Latin means leap)
NodeofRanvier
areexposedtotheECF;
voltagegatedNa
+
andK
+
channels are concentrated here
Myelin makes it much harder for
Latinmeans leap )
channelsareconcentratedhere
Myelinsheath/internode
Axon
Myelinmakesitmuchharderfor
currenttoleakoutthroughopen
ionchannels
APpropagationin
l d myelinated axons
APsaregeneratedthesame g
way,thoughthedepolarizing
currentflowsoverlonger
distances because of the distancesbecauseofthe
insulationprovidedby
myelin
NodesofRanvier are
strategically placed so that strategicallyplacedsothat
thereisenoughcurrent
remaining(i.e.alarge
enoughstimulus)tobring
thenextNodetothreshold
Why myelinate? Whymyelinate?
Onlyvertebrateshavemyelinated axons
Advantages include: Advantagesinclude:
1. Muchhigherconductionvelocity
2. Savesspaceaxonscanbemuchthinner (moreaxonsinagivenspace)
3. MetabolicallymuchcheaperAPsoccuronlyatnodes,thusonlyneed
voltagegatedNa
+
andK
+
channelsandNa
+
/K
+
ATPase atnodes
Squid giant axon No 500 25
Localanestheticsblock
l d h l voltagegatedNa+channels
Novocain,Xylocaine
blockvoltagegatedNa
+
channelsandthusAP
productioninthe
nerves that innervate nervesthatinnervate
thetooth
Prevents propagation of Preventspropagationof
painsignalstothebrain!
Chapter8:
SynapticTransmissionandNeuralIntegration
2typesofsynapses
ELECTRICAL CHEMICAL
Purves etal.2004
Electrical synapses Electricalsynapses
Neuronneuron&neuronglia
Directcytoplasmic connections
betweenpre &postsynaptic
cells allow electrical (& chemical) cellsallowelectrical(&chemical)
signalstobetransmittedfrom
oneneurontoanother
Occursviagapjunctions
Somearealwaysopen,othersare
gated
Commonincardiac&smooth
muscle;uncommoninnervous
system
Electrical synapses Electricalsynapses
Advantages:
Allowrapidcommunication(0.2msec)
Synchronize activity of connected cells Synchronizeactivityofconnectedcells
Canbeexcitatoryorinhibitoryatthe
samesynapse
Disadvantages
Oftenbidirectional communication
Nocapacityformodulationor
amplificationofthesignal(signalin
presynapticcell=signalin
postsynaptic cell) postsynapticcell)
Chemical synapses Chemicalsynapses
ArrivalofanAPinthepresynaptic
terminal leads to neurotransmitter (NT) terminalleadstoneurotransmitter (NT)
releaseintothesynapticcleft
NTbindstoreceptorsonthepostsynaptic p p y p
membraneandcausesaresponse
Canbeneuronneuronorneuroneffector
ll ( l l d) cell(e.g.muscleorgland)
Advantages:
Unidirectional Unidirectional
Facilitateintegration
Disadvantages:
Relativelyslow(2msec)
Highcost
Functional anatomy of chemical synapses Functionalanatomyofchemicalsynapses
Spacebetweenpre andpost
synapticterminals=synaptic
cleft (2050nmwide)
Mostcommontype
Chemicalsynapseatrest
Axonterminal
Presynaptic
neuron
(V
m
~75mV)
[Ca
2+
]
o
=10
3
M
Voltagegated
Synapticvesicles
[Ca
2+
]
i
=10
7
M
Ca
2+
channel
(VGCC)
Neurotransmitter
Reuptakemolecule
Synapticcleft
Plasmamembrane
Enzyme Receptor
Postsynapticneuron
Chemicalsynaptic
i i
1 AP arrives at terminal
Action
potential
1
transmission
1. AParrivesatterminal
2. VGCCsopen
Ca
2+
3
7
2 8
Q.WhichwaydoCa
2+
ionsmove,and
why?
V = 75 mV
4
6
V
m
=75mV
E
ion
=61mV log[ion]
out
z[ion]
in
5
E
Ca
=61mV log[10
3
M]
out
+2[10
7
M]
in
= +122 mV
Activepresynapticneuron
Responseofcell
=+122mV
Chemicalsynaptic
i i
1. AParrivesatterminal
Action
potential
1
transmission
2. VGCCsopen
3. Ca
2+
entrytriggersvesicle
dockingandsecretion;this g ;
involvesSNAREproteins
(accountsfor>90%ofsynaptic
delay)
Ca
2+
3
7
2 8
4. NTdiffuses(<10%ofsynaptic
delay)andbindstoreceptor
5. Responseincell
4
6
p
OftenincludeschangesinG
K
,G
Na
orG
Cl
GP
~0 55 msec from time of AP
5
0.5 5msec fromtimeofAP
arrivalatterminaltotimeof
postsynapticresponse
Activepresynapticneuron
Responseofcell
Terminatingchemical
i i i
Action
potential
1
synaptictransmission
Response is terminated by Responseisterminatedby
removalofNTfromcleft
6. Degradationbyenzymes
(multiple locations)
Ca
2+
3
7
2 8
(multiplelocations)
7. NTreuptakeintopresynaptic
terminal
Degradedorrecycled
4
6
8. NTdiffusionoutofsynapticcleft
9. NTreuptakeandmetabolismby
surroundingglial cells
5
Alloftheabovedecrease
receptoroccupancyandthus
Activepresynapticneuron
Responseofcell decreasepostsynapticresponse