1

LECTURE3
Generalpri
nci
plesofthebi ologi
caladjusting.
Nervouscontrol
.
Reflex.Refl
exarc. Functionalsystemasuni versalpri
ncipl
eof
regulation.
Lawsof exci tationconductiononnervousf i
bres.
Reflex as elementary reaction of organi
sm.Ref l
ex arc,its functionaland
structuralorganizati
oni
nobedi encetoaneurontheory.F
eed-back,i
tsimportance.
Functionalsystem asuni versalprincipleofregulation(P.К.Anokhin).Reverse
afferentationassystem-maki ngprinci
ple.Cl
assi
ficati
onofnervousf ibresandlaws
ofanexcitationconducti
ononthem.

Thebasi cstructuralel ementofthenervoussystem i sthenerve

cellorneurone. Eachneuroneconsi stsofasoma, orbody. Andprocesses,
whicharedi videdintoaxonsanddendri tes.Anaxoni sal ongprocesswhose
functioni stoconveyi mpul sesf rom thecel lbodytoothercel lsorto
peripheralorgans.
Dendritesarenumerousarborescentprocesseswhosefuncti onis
the recepti on of i mpulses arri ving f rom other neurones and thei r
conductiontothebodyofthenervecel l.
Thestructureofnervef i
bresandMECHANI SM OFANERVEI MPULSE
CONDUCTION
Nervef ibresaretheprocessesofnervecel ls.Theyaredi videdinto
myel i
natedandunmyel inatedf ibres. Themyel i
nsheathi sinterruptedby
annularconstri ctions;theporti onsoftheaxi s-cylindernotcoveredby
thesheath( 1
-2mi cron)areknownasRanvi er'snodes. Myel i
nhasatrophi c
functionandhasaveryhi ghelectri calresi stance.Ranvi er'
snodesi sof
great importance i n the mechani sm by whi ch nerve i mpulses are
transmi tted.
Conducti onofexci tation. I
nmyel i
natedf ibrestheacti onpotential
canspreadonl ybyj umpingoverporti oncoatedwi ththei nsulti
ngmyel i
n
sheath–f romonenodeofRanvi ertoanother. Thenodes' smembranehasa
+
great concentrati onofNa- channel s.Suchmechani sm i sknownas
saltotoryor“ jumpi ng”.

-
+ +
+
- -

TheRanvi
ernode

I
nunmyel
inatednervef
ibresexci
tati
oni
stransmi
ttedconti
nuousl
y
 2

al
lal
ongthemembranef
romoneexci
tedporti
ontothenext.

Classi
ficationofnervefi
bres(af
terErl angerandGasser1937).
Theclassificati
oni sbasedonthef ibres’sdiameter,presenceor
absenceofmyel ineandveloci
tyofthenervousi mpulseconducti
on.

Generalf
ibertype Sensoryf
ibertype Di
ameter
Conducti
onveloci
ty

(microns)
A-( al
pha) I
amuscl
espi
ndl
eaf
ferents 1
2-22
70-120m/ s
Large-motoneurons I
bGol
gitendonorgans

A-( beta) II
Secondaryafferentsofmuscl e 8-12
40-70m/ s
Touch, pressure spindles;touchandpressure
A- (gamma)
4-8 15-40m/ s
-Motoneuronsto
muscl espi ndles
(intraf usalf i
bers)
A-( delta) I
IISensoryfibersf rom
1-4 5-15m/ s
Sensoryf ibersfrom visceroreceptorstemperature
visceroreceptors, andpain
temperatureand pai n
B
1-3 3-18m/ s
Pregangl ionicautonomi
c
fibers
C (
unmyelinated)
0,5-1 0,5-3m/ s
Slow pai n; postgangl
ionic I
V Pai n and temperature f ibres
(slowest)

ThelawsoftheExcitationConductiononnervousfibres.
1.
Thelaw ofanatomi calandphysi ol
ogi
calcontinuityofanerve.
I
mpulseconducti
onisonl
ypossi bl
eifthephysiologi
calcontinui
tyofnerve
fi
bresi
sunbroken.
2.Thelaw oftwo-wayconducti on.Whenasti mulusisappli
edinto
themiddleofanervef ibre,excitationisconductedal ongitinboth
 3

directions.
3.Thel aw ofisol atedconducti onal onganervef ibre.Impulsesare
conveyedbyeachnervef i
breseparatel y,withoutpassi ngfromonef ibre
toanotherandactonl yonthosecel l
si nwhi chthef i
brestermi nate( with
theexcepti onofunmyel inatedfibres–Cgroup) .Forex.Stimulationofone
ofth erootswi llcausecontracti onnotofthewhol emuscl e( asitwoul d
ifexcitationwaspassedf romonenervef ibretoanother) .
4.Thel aw off uncti onalunspecifici
tyofnervef i
bres.Thenervous
fibref unctioni sunspeci f
icandi tisonl ybasedonthef unctionsofthe
worki ngorganandnervouscentrethi sf ibrei sconnectedwi th.This
featurei saveryi mportantf orneurotranspl antation.
5.Thel aw ofrel ativeindefatigili
tyofanervef ibre.Anervei s
practi cal
lyindef ati
gabl einanai rmedium. Tocausethef atigueofanerve,
itisnecessarytoputi ti ntothemedi um wi thoutoxygenori ntothe
medi umofni trogen.

Thecl assi ficationofneurones

Receptorneurones( afferentorsensory)arebi polarnervecel ls
performi ngthef unctionofrecepti onandconducti onofsti mulif rom
peripheralreceptorstothecentralnervoussystem.
Contact,or i nterconnecting,or i nternuncial neurones ( or
interneurons) formoneofthemostnumerousgroupsofneuronesi nthe
centralnervous system.They provi de communi cati ons between the
receptorandef f erentnervecel ls,andaccordi ngtothecharacterofthe
effectproducedbythem, theyarecl assifiedasexcitatoryandi nhi
bitory.
Efferentneurones sendi mpul sestotheperi pheralorgansand
tissues, whicharecarri edal ongtheirl ongaxons. Thosethatf ormmotor
nerve f ibres l eading to the skel etalmuscl es are known as motor
neurones. Thei rbodi eslieintheanteri orhornsofthegreymatterofthe
spinalcord.
Thepri ncipalandspeci fi
cmani festationofthecentralnervous
system( CNS)acti vi
tyistheperf ormanceoftheref lexacts, orref lexes.
Aref l exi saregul arreacti onoftheorgani sm i nresponseto
stimul ationofreceptors, asarule,ef fectedthroughtheCNS. Ex.Drawi ng
theh andasi deaf tertouchi ngthehotobj ect.
Theref lexarci sastructuralbackgroundofref l
ex. Theref lexarci s
thepathwayf orthenervei mpulsesthattravelf rom areceptortoan
effector( worki ngorgan)duri ngtheperf ormanceofanyref lex.

Thereflexarciscomposedof:1).Receptors(recepti
vefi
eld)
,which
acceptthestimul ati
on;
2).Af
ferent( sensory)nervefi
bres,i.
e.theprocessesofreceptor
neuronescarryingwavesofexcitati
ontothecentralnervoussystem;
 4

3).Nervecentre( neuronesandsynapses, analysi

ngani nformation
andconveyingimpulsestotheef fectorneurones);
4).Efferent(motor)nervef ibrestransmittingimpulsesf romthe
centralnervoussystemtotheperi phery;
and
5).Effectororworkingorgan( muscleorgland) ,
responsibl
ef orthe
reflexreali
sation.

Thesi mpl estref lexarccanbef ormedbytwoneuronsonl y,a

receptorandanef fectorwi thasynapsebetweenthem. Sucharef lexarc
iscaledbi
l neuronalormonosynapti c.
The ref lex arcs of the most ref l exes are mul ti
neuronalor
polysynaptic.
Therearemotor( efferent)andi nternunci alneuronsi nCNS.
Forthenormalref lexactaf eed-back( transmi ttionofthei nformati on
fromthereceptorsofef fectororgantonervecentre)i snecessary. Itis
necessaryf orcontrolandcorrecti onofef f ector' swork.
Functionalsystem( byAnokhi n).
Functionalsystemwasof f
eredbyMoscowacademi cianAnokhinasa
generalprincipleoftheorgani smf unctionsregul ation.
Functionalsystem i sadynami corgani zati on,whi chtemporaryand
selecli
velyuni tessomenervouscentresandperi pheralorgansonthe
basis ofnervous and humoralmechani sms f or reachi ng the usef ul
,
adaptableresul t( homeostasi s) .
Mainstagesofaf uncti onalsystemorgani zation:
I. Af ferent( sensory)synthesi s.Accepti onofthei nformati onf rom
theenvironmentandi nternalmedi umandi tspri maryanal ysistakes
place. Al lthei nformati oncanbedi vi dedi ntocondi ti oni
ng( general)
andstarti ng( mainchange, causeth ef uncti onalsystemf ormati on).
II. Acceptanceofthedeci si
onofacti ng.
III
. Creati ontheprogramofacti ng.
IV. Acceptorofthef utureresul ts.Usi ngtheprevi ousexperi ence, the
organism canexpectthef utureresul tf orcreati ontheopti mal
program.
V. Ef ferentexci tation.Thecommandstransf ertotheworki ngorgans.
VI. F eed-backf romthereceptorsofworki ngorgans.
 5

LECTURE3a
Structural ly-functionaldescriptionofsynapses.F eaturesofconducti ngof
exci
tation through synapse.Synapti c transmi tters.Postsynaptici nhibiti
on and
presynaptic i nhibiti
on,synapti c mediators. Di vergence and convergence are
structural l
y-f unctional pre-condi tions of coordi nati
on. I rradiation and
generalisationofexci tation.Refl
exesare: antagonistic,
alli
ed,
sinergi
c,chain.Pri
ncipl
e
ofgeneraleventualway. Princi
pleofdominant( О.
О.Ukhtomsky),i
tsproperty, stages.

SYNAPSES
Synapsei sastructural -functionalcontactfortransmi ttionof
impul ses.
Synapses l ocated on the body ( soma)of a neurone are cal led
axosomati candthoseonthedendri tesaxodendri ti
c.
A synapse i n the centralnervous system,l ike peripheralones
consistsofthenerveendi ngproper,synapticcl eft,andpostsynapti c
membrane.Nerveendi ngsinthecentralnervoussystem varyi nshape,
general lyl ooki
ng like knobs.Each synapti c knob i s covered wi th a
membrane-socal ledpresynapti cmembrane–andi nsi
dei tthereare
vesiclescontai ni
ng,inawi delyheldview,asti mul atingorani nhi
bitory
medi ator.
Synapsesaredi sti shedasexci
ngui tatoryori nhibitoryaccordingto
theef fectproduced.
 6

Synapti cTransmission
Generalcharacteri sticsofchemi calsynapses:
1.Anacti onpotentiali nthepresynapti ccel lscausesdepolarization
ofthepresynapti ctermi nal.
2.Asaresul tofthedepol arization,Ca2+ entersthepresynapti c
2+
termi nal.Ca entry causes rel ease of neurotransmi tter into the
synapti ccleft.
3. Neurotransmitterdi ffusesacrossthesynapti cclef
tandcombi nes
wi threceptorsonthepostsynapti cmembrane,causi ngachangei ni ts
permeabi litytoionsandi tsmembranepotenti al.
4.Exci tatory neurotransmi tters depol arize the postsynapti c
membrane;
inhibitoryneurotransmi ttershyperpol ari
zethepostsynapti cmembrane.

Neuromuscul arj uncti on

Iti sthesynapsebetweenaxonsofmotoneuronsandaskel etal
muscl e.
Theneurotransmi tterrel easedf romth epresynapti ctermi nalisAchand
thereceptorsonthepostsynapti cmembraneareni cotinic.
1.Synthesi sandstorageofAchi nthepresynapti ctermi nal.
Cholineacetyl transferasecatal yzesthef ormati onofAchf rom acetyl
coenzymeA( CoA)andchol i
nei nthepresynapti ctermi nal.
Achi sstoredi nsynapticvesi cleswi thATPandproteogl ycanf orl ater
release.
2. Depolarizationofthepresynapti ctermi nalandCa2+uptake.
Actionpotenti alsareconducteddownthemotoneurontoi tspresynapti c
2+
termi nalwheredepol arizationopensCa channel s.
2+ 2+
WhentheCa permeabi li
tyi ncreases,Ca rushesi ntothepresynapti c
termi naldowni tselectrochemi calgradient.
2+
3. Ca uptakecausesrel easeofAchi ntothesynapti ccleft.
Thesynapti cvesi clesfusewi ththepl asmamembraneandemptythei r
contentsi ntothecl eftbyexocytosi s.
4. DiffusionofAchtothepostsynapti cmembrane( muscl eandpl ate)
andbi ndingtospeci ficreceptors.
TheAchreceptori salsoaNa+andK+i onchannel .
Bindi
ngofAchtothei r subuni tscausesconf ormati onalchangesthat
opensthecentralcoreofthechannelandi ncreasei tsconductancetoNa+
andK+. Thesearel i
gand-gatedchannel s.
5. Endpl atepotential( EPP)inthepostsynapti cmembrane.
Becausethechannel sopenedbyAchconductancebothNa+ andK+ i ons
 7

increases,thepostsynapti
cmembranepotenti ali
sdepolari
zed.
Thecontentsofonesynapti cvesicle(onequantum)producesamini
ature
endpl atepotential(
MEPP–thesmal l
estpossibl
eEPP).
MEPPssummatetoproduceaf ullEPP.TheEPPisnotanactionpotenti
al,
butadepol arizati
onofthespeciali
zedmuscleendplate.

6.Depolarizationofadjacentmusclemembranetothreshol d.
Once the end pl ate region is depolari
zed,l ocal current causes
depolarizationandactionpotentialsintheadjacentmuscl e.Contracti
on
foll
ows.
7.DegradationofAch
TheEPPi stransi entbecauseAchisdegradedtoacetylCoAandchol i
neby
acetylcholinesterase(AchE)onthemuscl eendplate.
Onehal fofthechol i
neistakenbackintothepresynapti cendingbyNa+–
choli
necotransport.

Synaptictransmi ssioninneuronalsynapse
Inputtosynapses
Thepostsynapti ccellintegratesexci tatoryandi nhi
bitoryinputs.
Whenthesum ofal lthei nputbri ngsi tsmembranepotenti alto
threshol d,itfiresanacti onpotential( onthetri ggerzone-axonhi l
l).
1.Exci tatorypostsynapti cpotenti als( EPSP)
EPSPsarei nputthatdepol arizethepostsynapti ccell,bringi
ngi t
closertothreshol dandcl osertof i
ringanacti onpotenti al
.
EPSPsarecausedbyopeni ngofchannel sthatarepermeabl etoNa+and
K+,si
mi l
artotheAchchannel s.
Exci tatory neurotransmi tters i ncl ude Ach, norepi nephri
ne,
epinephri ne,dopamine,glutamate, serotoni n.
2.I nhibitorypostsynapti cpotenti als( I
PSP) .
IPSPsarethei nputthathyperpol arizethepostsynapti ccell,moving
itawayf romthreshol dandf atherf romf iringanacti onpotenti al
.
- -
IPSPsarecausedbyopeni ngCl orK channel s.Themembranei s
hyperpol arized.
Inhibitoryneurotransmi ttersareγami nobutyricaci d( GABA)and
glycine.

I
nhi
biti
onistheoppositeofexci
tati
oni ni
tsphysico-chemicalnature.
I
ti sanactiveprocess.Therearespeci alinhi
bitoryneurones,produce
speci
ali
nhibi
torymediators(GABAandgl
ycine).
I
nhi
biti
onisasuppressi onofacti vityasaresul tofexci tation.
 8

Othersmaintainedth
atinhi
biti
oninthecentralnervoussystemresulted
fromaconf li
ctbetweenseveralexcitati
ons.Thereareseveraltypesof
inhi
biti
oni
nthecentralnervoussystemofvariednatureandlocali
zati
on.

Postsynaptici nhibition.There are i nhi

bitory neurons wi th the
inhi
bitorymedi ator.Theaxonsoftheseneuronesf orm nerveendi ngsin
thebodi esanddendritesoftheexci tatorycel ls.Thenervei mpul ses,
generatedbysti mulationofi nhibi
toryneuronsdonotdi fferf rom the
action potentials of ordi nary neurons,but the medi ator does not
depol ari
ze the postsynapti c membrane, but on the contrary,
hyperpol arizei
t.Thi
shyperpol ari
zationi scalledinhibi
torypostsynapti c
potenti al.Thetypeofi nhi bitiondescri bedherei scalledpostsynapti c
becausei tisduetohyperpol arizati
onofthepostsynapti cmembrane.
Presynapticinhi
bition. Asi tsnamei mpl i
es,thistypeofi nhibi
tionis
locatedi nthepresynapti cel ements. Thistypetakespl aceintheaxo-
axonalcontacts.
Thesearethepresynapti ctermi nals,formi ngspeci fi
ci nhibi
tory
synapses. Themediatorssecretedi nthesei nhi
bitorysynapsesdepol ari
ze
themembraneofthetermi nalsforal ongti meandbri ngthemi nastate
similartocathodaldepressi on,whichresul tsi naparti alorcompl ete
blockofconducti onofimpul sestothenerveendi ngs.

NERVE CENTRES
Anerve centre i s a group ofneurones acti ng togetheri n the
performanceofadef ini
teref lexori ntheregul ationofaparti cular
function.
Forexampl e,thecentref orthekneej erkref lexisl ocatedinthe
secondtof ourthlumbarsegments, andthecentreofthepl antarreflex
inthef i
rstandsecondsacralsegmentsofthespi nalcord.
Theproperti esofnervecentres:
Thel aw ofone-wayconducti on.I
ntheCNSexci tati
oncanonl ybe
conveyedi nonedirecti
on,fromreceptorneuronetoef fectorneuronevi a
internuntialneuronebecauseimpul sespassacrossthechemi calsynapses
inonedirectiononly.
Delayed conduction.Excitation is conducted consi derably more
slowlyi n nerve centres than i n nerve fibres.The peri od between
stimulationandresponseisaref lextimeorthel atentperi odofref l
ex.
Theperi odtakenbyintracentraltransmi ssionofexcitati onfroman
 9

af ferenttoanef ferentneuronei sknownasthetrueorcentralref l

ex
time.Sl ow transmi ssionofexci tationinthecentralnervoussystem i s
consequentonthepecul i
ari ti
esofi mpul seconducti onacrosssynapses.
Theperi odf romthearri valofani mpul seatthenerveendi ngtothe
appearanceoftheexci tingpostsynapti cpotenti ali sapproxi matel y0,5
mi lli
second.Thatperi odi scal ledthesynapti cdel ay.Another1 ,
5to2, 0ms
passbetweentheappearanceofthepostsynapti cpotenti alandi tsri si
ng
tocri ticalval ue.Thus, conducti onacrossasynapserequi resaltogether2
or3mi lli
seconds.
Summati onofexci tati onisacharacteri sticofnervecentres, f i
rst
descri bedbySechenovi n1 863.Itsessencei sthatacombi nationoftwoor
moresti mul ationsofperi pheralreceptorsoraf ferentnervesarousesa
ref lex,wh ereaseachtakenseparatel yisnotsuf fi
cienttoel icitthe
reacti on.Twotypesofsummati onareknown: consecutive( temporal )and
spati al.
Consecuti vesummati oncanbei nducedexperi mental lybyappl yinga
seri esofrhythmi csubthreshol dsti mulitoanaf ferentnerveortothe
recepti vef i
el dofaref lex.Impul sesshoul dcometoanervecentreone
af teranotheratshorti nterval( l
essthandurati onofEPSP( 1
2-1 5ms) )
alongthesameaf f erentnervef ibres.
The spati alsummati onoccurswhentwo ormore subthreshol d
sti muliareacti ngsi mul taneousl yondi fferentreceptorsbel ongingto
thesamerecepti vef ield.Ei therofthesesti mul iactingal onewi llnot
inducethescratchi ngref lex,buti ncombi nationtheydo.
Spatialandconsecuti vesummati ontakespl acewhensubthreshol d
sti muliareappl i
edati nterval sshorter1 5mi ll
isecondontwoaf ferent
fibresofthesamerecepti vef i
eld( ofoneref lex).Thepostsynapti c
potenti als( EPSP)areaddedandthesummaryreachescri ti
call evelof
depol arizati
on.

Transf ormationoftherhythmofexci tation.

Nervecentresarecapabl eoftransforming( changi
ng)therhythmof
theingoi ngimpulses.
Therearetwotypesofi tsphenomena:decreasingtypeandincreasi ng
type.
Decreasi ngtypeofatransf ormationisconnectedwi ththel ow
labi
lityofsynapses( 1
20-150imp/sec).
Increasingtypeofarhythm' stransformati onhasseveralreasons: 1
.
Longexci tingpostsynapticpotential
,whichtriggersasecond,thi rd,
etc.,
actionpotenti alaf terthetermi nati
onofthef i
rst;2.I
fthenegati ve
after-potenti ali s strong enough to reach the cri ti
call evelof
depolarizationofthemembrane, itmaygiverisetoanotherimpul seandso
 1
0

on;3.Ci
rcul
ati
onofnervei
mpul
sesi
ncl
osedneuronalchai
nsoftheref
lex
centre.

Af teraction.
Ref l
exactsdonotendsi mul taneousl ywi thcessati onofthesti mul i
causingthem, butafteracertai n,someti mescomparati velylongi nterval .
The phenomenon,cal led af ter-acti on,has the same reasons as the
transf ormati onoftherhythmofi ncreasingtype.
Posttetani cpotenti ation.
This phenomenon decl ares i tself as i ncreasi ng of the reacti on
strength af ter previ ous rhythmi c subthreshol d sti mul ation.The
mechani smoftheposttetani cpotenti ationthef ollowing:increaseofthe
mediator quanti ty and i ncrease of the postsynapti c membrane
excitability.
F atigueofnervecentres.
Fatiguei satemporaryreducti onoftheworki ngcapaci tyofacel l,
organororgani sm asawhol eresul tingf rom prol ongedexerti onand
passingaf terarest.
Themyoneuralj uncti onbecomesf atiguedmuchearl ierthanthe
muscl ef i
bres,andi nconsequence the muscl ei sprotectedagai nst
exhausti on due to prol onged exerti on by the bl ocking of i mpul se
transmi ssi onf romnerve; andi ntheorgani smaswhol ethenervecenters
tireevenearl ierthanthemyoneuralj uncti ons.
Thef atigueofnervecentersmaybecausedbyasharpf al linthe
reserves of synthesi zed medi ator i n the nerve endi ngs,di minished
sensitivityofthepostsynapti cmembranetothemedi ator,orareducti on
ofitsenergyreserve. Itisnecessarytonote, thatthesynapsesarevery
sensitivetol ackofoxygen.
Fatiguei nmuscl eisbroughtaboutbytwopri ncipalcauses:
1.
One i s the accumul ation of metabol ites i n the muscl e duri ng
contracti on( i
nparti cular, ofl acti cacidthati sf ormedduri nggl ycogen
breakdown) ,
whi chproducesadepressi veef fectontheworki ngcapaci ty
ofthemuscl ef i
bres.Partoftheseproducts, andpotassi umi ons, diffuse
from thef i
bresi ntotheperi cel l
ularspaceandexert ani nhibitory
infl
uenceonthecapaci tyofthemembranetogenerateacti onpotenti al
s.
2.Thesecondcauseofprogressi vef atiguei sgradualexhausti onof
theenergyreserveofthemuscl e.

Sechenovmadethetheoryaboutanactiverest.Sechenov(1
903)f i
rst
showedthatrecoveryoftheworki ngcapaci
tyoff atiguedmusclesinthe
humanarmaf terprol
ongedexerti
oninlifti
ngaload,markedlyquickened
 1
1

i
fworkwasperf
ormedwi
ththeotherarmduri
ngtheperi
odofrest.

Kindsofinhibi
tioninCNS
Recurrent(Renshaw)inhibi
tion.Renshawcel lsarei nhibi
torycellsin
theventralhornofthespi nalcord.Theyrecei vei nputf rom col
lateral
axonsofmotoneuronsandwhensti mulated,negativelyfeedback(inhi
bit)
onthemotoneurons.Thi smechani sm li
mitsthemuscl ehypertoni
cityon
thel evelofthespi nalcord.Disease i nfantil
ecerebralparal ysi
si s
causedbydisordersofthiskind inhibi
tion.

Reciprocalinnervati
on. ( Ex.Innervationofantagoni stmuscl es.
Flexionandextensi onref lexes,inspirationandexpi rati
on.)Af ferent
nervef ibresformarborizationsinthespi nalcord,someofwhi chconvey
excitationtothemotorneuronssuppl yingthef lexormuscles,otherto
the neurons,f orming i
nhibitory synapses wi th the motor neuron,
suppl yi
ngtheextensormuscl es.

Lateralinhi
biti
on.Themainideaofthiskindofinhi
biti
onisininhibi
tion
ofthel ateralchannel
s,conductingthei
mpul sationbyinhi
bitoryneurons
throughrecurrentcol l
aterals.Thi
skindofi nhi
biti
onisbasi cf orthe
“funnelofSherri
ngtone”formingandiswidelyspreadinsensorysystems.

Maintypesofunconditionedreflexes:
1.Antagonist(reci
procal)reflexes(flexi
onandextensi
on;
inspi
rationand
expiration;i
nspirati
onandswal l
owi ng)
.Theycannotbecarri edout
simultaneously.They are based on the antagoni st (reci
procal)
inhi
biti
on.
2.Synergicreflexes–ref l
exes,realizi
ngthesamemotoract.F orex.:
Movementofarmsandl egsatwal ki
ng.
 1
2

Alli
ed–ref l
exes,whichreinforceandi ntensi
fyeachother.F orex.
:
Sali
vationasresul tofsi multaneoussti mulationofmechani cand
chemicalreceptorsi smorethanth eirsum asresul tofseparate
stimulati
on.
3.Chai
nandrhythmi crefl
exes.Compl exref l
exactsi nwhichoneref lex
excitesanotherandisreplacedbyi tarecalledchainreflex.Theyoften
occurinadef i
nitesequenceofthesamesi mpleref l
exes,whi chare
repeatedrhythmi cal
ly.
Theseref lexesincludewal ki
ng,scratchingand
otheractions.

COORDI NATIONOFREF LEXES

Theinteracti onofneurons, nervouscentresandnervousprocessesi n
thecentralnervoussystem thatensuresi tsharmoni ousacti vityi s
calledcoordi nati on.
Therearetwopri nciplemorphol ogicalbackgroundsofcoordi nation–
convergenceanddi vergence.
Convergence.I mpul sesreachi ngthecentralnervoussystem al ong
variousaf ferentf ibresmayconvergeuponthesamei nternunci aland
efferent neurons.Thi sf act underl ies the pri nci
ple ofconvergence
establ i
shedbySherri ngton.F rom 2to6thousandofsynapsesmaybe
formedononeneurone. ButPurki nj e’
scel lsmayhaveabout200thousand
synapses. MotoneuronesandPurki njecel lsmaybetakenasexampl esof
convergencecentres.
Convergenceexpl ainsthespati alsummati onofexci tationandthe
phenomenonofoccl usion.
Pri
nci ple of the f inalcommon path.One and the same ref l ex
movementmaybecausedwi thgreatmanydi fferentsti muliacti ngon
variousreceptors. Oneandthesamemotorneuronmaybei nvolvedinmany
reflexarcs. Ef fectorneuronsf ormaf inalcommonpath, asSherri ngton
calledit. Thebackgroundofi ti sconvergenceofdi fferentaf ferentf ibers
onthesamemotoneurone( “funnelof Sherri ngtone”). Asresul t,
amountofef f erentf i
bresi sl ess, thanth eafferent.
Reflexes,whosearcshaveaf i
nalcommonpathmaybedi videdi nto
alli
edandantagoni stic.
Fasi
l i
tati onconsi stsi nsi mul taneoussti mulationoftwogroupsof
afferent f ibres ( each of whi ch causes a weak ref lex muscul ar
contracti on) ,produci nganef f
ectwhosemagni tudei smorethanthe
arithmeti calsum ofthoseref lexestakenseparatel y.I
ti saresul tof
convergenceonsubthreshol dneuronesbyspati alsummati onmechani sm
(summati onofEPSP) .
 1
3

Occl
usion.
Occl
usionconsi stsessentiallyinsimultaneoussti mul
ationoftwo
groupsofaf ferentf i
bres(eachofwhi chcausesastrongref l
exmuscul
ar
contraction),producinganef fectwhosemagni tudei slessthanthe
ari
thmeti calsum ofthoseref lexestakenseparatel y.Iti
saresultof
convergenceonthreshol dneuronsby“allornothing”l
awmechani sm.

Irradiationofexci tati on.

Divergence( whenf ibersoftheoneneuronbranchandproj ecttothe
different neurons) i s the morphol ogicalbasi sf or the i rradiation.
Impul sesarri vi
ngatthecentralnervoussystem duri ngstrongand
protractedsti mul ationi nduceexci tationnotonl yi ntheneuronesofa
givenref l
excentrebutal sointhoseoftheothercentres.
Thisspreadofexci tati onthroughthecentralnervoussystem i s
known as i rradiation.The maxi mall evelof the i rradiati on called
general izationofexci tati on,whentheexci tati oncoveredal ltheCNS.
Factors,whi chattempt thei rradiation:1 .Strengthofsti mul us
(exampl eonthef rog'sref lex);2. Excitabilityofastructure;3.Bl ocking
thei nhibitorysynapsesandrel ievingpostsynapti cinhibiti
onbyi njection
ofstrychni neoratetanustoxi n. I
f0, 1%sol utionofstrychni nei sinjected
undertheski nofaf rogi ti ssuf ficientaf teraf ew mi nutesmerel yto
touchi tsl egortoknockonthetabl eonwhi chi tissi ttingtoi nducea
most i ntense general exci tation of the central nervous system
accompani edwi thconvul sionsofal ltheskel etalmuscl es;4. Thestageof
adevel opmentofthenervoussystem i nontogenesi s( anew-bornbaby
hasaconsi derablymoreextensi vei rradiationofexci tationthroughthe
CNSthani ntheadul t).

Theprinci pleofthedomi nantwasf ormul atedbyUktomskyasthe

basicworkingpri ncipleofnervecentres.I tisthattheacti vityofthe
nervous system as a si ngle entity is characteri zed in the natural
conditi
onsoftheorgani smbytheexi stenceofdomi nantf oci
ofexcitation,
whichchangetheacti onofal lothernervecentresandsubordi natethem
tothemsel ves.
Ex.
:Pain;f oodandsexi solati
on;def ecationandotherdomi nants.
Motherdomi nant,prof essionaldominants.
Dominant f ociof exci tation are characteri zed,according to
Ukhtomsky, bythef oll
owi ngbasi cfeatures:1).Heightenedexcitabil
ity;
2) .
 1
4

Stabili
tyofexci tati
on;3)
.Capacitytosummateexci tation;4).I
nerti
aof
excitation.
Theri seofadominantexcitationinanycentrei salwaysaccompani ed
withamoreorl essmarkedconj ugateinhi
bitionofothercentres.
Stagesofadomi nant:
1.
Averywi deinfactunlimitedirradiati
on;
2.Concentrationoftheexcitationinthedef i
nitecenterofCNS.