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J1eKTpOrexHuKe XOpOIlJO H3AeCTHO, "ITO
06a ..14>cI>eKTa cywecTBeHHo nOBblllJaJOT npoBOft,HUJ.He CBOHCTsa 3J1CKTpH\feCKOrO
K
6eJlH, YMeHbWdH 8 HeM YTe\fKY TOKa It 3aTyxaime CHrHa.na.
B .4flle:ltUiOObiX HepOHblX 80 lOKHGA liOHHhle KaHaJ1b1 pacnpe.n.e.neHbJ HCp3BJlO
MepHo. TaK HaTpHeBble KaH3J1bl H OCliOBHOM paCnOJlOiKellbl D nepe>'B3Tax PaHBbE:', 3
B MC>KnCpeXBaTltoii o6J1acnt IiX 3Ha'IHTenbHO MeHbwe (Shrager, 1987; Meiri e/ al..
1986). B 06Jl3CTHX, npl.IMbIKatoUJ,HX K nepexH3TaM PaHBLe, HMelOTCSI pa3nli'IHble
THlIbl Ka:H1CBbIX KaIl3JlOB, 06J13..13IOWlfe 6b1CTpofi Hnll Me.D.,.1eHHml KHHeTHKOH. KdK Y
aM<p u6HH , TaK Ii Y MJleKonHT310111HX HX nJlOTHOCTb YMeHbWaCTCij no Mepc Y.!laJ1eHIiR
OT nepexBBToH PaHHbc (Grissmer, 1986; RIper, Schwarz. 1989). np.f B036y>K.il C -
ItHH MlleJ1HIIOBOro JlepBHOro BonOKHa npoRe..1eHHe nJ1. oCYUleCTBJlReTCH 1I0CpCllCT-
BOM JlOK3J1hHhIX TOKOR npH6J1H.nne.fJbHO TaK >KC, KaK }f Y 6C3\fHe.nIlItOBOrO HepBHoro
8UJ10KH3, (pHC. 4.4. a). OT.fJUI.{He MuenUHOBoro B0J10KH3 COCTOHT B TO\4. "ITO ero
HCTO-IHIIKH 8KTHBHpYlOIIlero J10KaJlbHOrO TUK8 paCnOJIO>t<CHbl B ncpeXD3Tax PaHBbe
(TO eCTb DIm HCOllHOpOllll0 pacnpe!leJ1enbi B,!1.0Jlb MeM(5paHbI aKCOHa). J1oKa.lbtlble
TOKH, Bbl
OJJllllHe "3 B036y>KJteHHOrO nepeXRHTa, OKa3b18aJOT 3KcnOHCIIUU3.nbHO
YMeHbWatoUJ,eCCR 803D,dkTBHC BllOJ1b HcpBlioro DO",10Klla 113 npOTR>KemHl HeCKonb-
KUX Me>KnepeXB3THblX YtlaCTKOB. COOTBeTCTneHHO. OHH BHOCRT BK..'1BJ1. I:J Acno.nApH-
3aUHtO HeCKOJ1hKH>. nepeXRaTOB PaHBbe (pHC. 4.... 6).
TaKHM 06p330M. npOBelleHUe B M.feJ1HHOBOM HepBHoM AOJlOKHe npmfcxolHT
nocpeAcTBoM nOCJ1eJI.ORaTe",1bHoH aKTHB311lHf ncpc>"BBTOB PallBLe, B XO!l.e KOTOp011
J10....aJ1bHbIC TOKH nOBbiWalOT ux Me"d6pallHblH nOTeHUUaJ1 .!I.O noporoBoro ypOBHS1.
3TOT npouecc -laCTO Ha3blBalOT ca 7hmamOpHbUI. npooeiJeHUe.M (OT J1aTltHCKoro
saltare, «npbJrt:lTb»), nOCKOJlbKY nOTeHuuaJ1 .a.elicTBIUl «nepecKaKHBaeT» OT nepe-
XBaTa K nepexB3T)'. n pH HeH3MeHHOM .!UmMel pc 3KCOHa, \IHCJlWUlJaWfS1 YBeJlH\fH-
BaeT CKUpOC1 b npOBCllemfS1 no HeMY npH6.rHf
HTeJ1hHO B 20 pa3. t\"HeJ1HHOBOe uepB-
Hoe BOJlOKHO npCJlCraBJlHCT co6oi'1 CJlO>KllblH 6J1OJ10ntlleCKIiH Ka6t'JJh C Hep3lmo-
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)IJ1H nCCnellOB3HJ.lst npOBe,JJ.eHHS1 tfepBHbIX HMnVJILCOB II I

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npHMcHeHbI B TaKHX o6JtaCTStx, KdK: ( I ) P acn p ocT p aH e lL (lle n n
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CJlellOBhlX norCHUH310B (Blight. 1985)' ( 3 ) MO
 e.nHpOBat H
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3J1eKTpH'feCKHX CBOHCTB MHe
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CummIns, 19d5); (5) npOBe.n.eHHe B ..1CMHeJlHHI
3H poBaHliO 00 ,
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(\Vaxman, nl, I 8)... (6) npOBeJJ.eHlfe B 6e3MHerU fJIOBO
. M HepBHOM fiO
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Ka (Barach, \\'ikswo, I gS7).


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CJlO.)KHbIe npo6.1eMbl. CBH3aHHbie C O!l.HOApe\feUllort 3.fJeKTpHllecKoH . .
MUOfllX KJ1CTOK B 06LeMHOM n:1 aKTHBHOCTblO
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TOKa n 'u _ H Re.lleT ceuR IIp3KTIi'reCKH KaK HCTO'IItHK nOCTORHHoro
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'jHT "i o B o ll H oe BOJlOKHO. C l IHT3eM,
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, llOcTaro'UlO fieJlHI< OTHOC
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malOU!cro HepBHoe 80J10KHO ) Ha 4 4 ( )
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CJ1e:).\! C \! . Irv.\. IIHoro Y"IacTKa BOJlOKHa B oKpYlf\afOul Y IO
'0/ JllCJJbHulM conpoTJm rrCHHeM P T ..
BE:1CToyeT pacn _.
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n pe..'1CJ1CHHfO 3apfl.ila, npellCTaBJlCHHO'dY H3 pHC. 4.4 (a )
OCKOJ1LKY nOTellUH - .
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npOCTpaHc
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IIOC p;JcnpelJ,CJleHHt' nOTeH
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1ReTCSI
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vpaHLj RBJlHeT c a ( 1 ) 't' u oTeHUH3J18 Ha BHeWlieH JlOBepXHOCTH MCM-
tt:.. n TpexcpH3HOH ( ? ) 60Jl ee n -
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,
Y>K...'1eHHR R0J10KHa H ( 3 ) J -'
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J ,I.fc
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( ncnOllellUHallLIlU C Y Re J
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M pa!UianbHoro paCCTOstfU1H OT B0J10KH3
OTeHL.UI3JI 'J,OCnlraCTCR Y 15
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m KpOBOJlbT. .





142
4 npPOAA 6OnOTEHLlIltAn08
 3 d>YHKLlOHAllbHA CTPYKTYPA nEPV1cpEPV14ECKOVl HEPBHOIi! CVTCTEMbi 143
(a)
Cyr.t\WH nOTeIlUIiaJ10B, C03)).aBat:MblX B IumHOJ-t TO'IKe BceUf HCTO"lIlHKaMH BHYTpH HepR-
Itoro CTBOJJa. 06b1ljllOH 4>OpMOH :1I\CTpdKJIeT01.IHoro n.n. HepBHoro CTBo.na, Bhl3b1BaeMO-
ro 3J1eKTpH'leCKOH CTHMYJ1ffUHeH, ftBJJAeTCR Tpexcpa3Hhlif OTBeT aMnJ1Hl)'.lI.0H B HeCI\OJ1L-
1<0 MHKpOBD.nbT, I<OTOpblH YMeUbwaerCR npH YJJ.dJ1HHH OT HepBa, npli'-leM 3TO YMeHb-
weH.fe 60 'Iee Bhlpa/KeHO AlIA BhlCOKO"laCTOTHOH cocraBnffJOUl.eH n.n.
O..lHHM H3 cnOCO()OR YRenWleHHR penfCTpHpYCMblX CHf'HdJ10B npH BHeKneTO"l-
HOM OTReJ],CHHH HBJIReTCR H3MeHeHHe napaMeTpOB OKpY>KaIOWeH BOJ]OKHO Cpe.n.LI
(HBrPY3KH). JInSi 3Toro MO>KHO nOBblCHTb y'teJJbHOe COnpOTlfBJ1emiC HJJH YMHbWHTb
06"beM OKpY>f<at01l1ero npOBOllH.,K paCTBopa.
npCn.weCTBYlOll1ee H3JJO>KCHHC 06"bHCHReT MeXaH1f3\fbl 3J1eKTpOrene3a 3Hr,
OTBO.!utMOH OT nOBepXHOCTIi PYK, 1I0r, HJlU JUlua. AHanOrHljHhlM 06pa30M ItHTepnpe-
THPy'OTCR \1HOrHe 6H03.'1eKTpH'leCKUe SlBJ1CHHR, KaK HeflporeHHble (Rbl3B8HHble no-
TCHUli3J1b1, perHcTpHpyeMble OT npOBomlUUfX nYTeH n CmlHHOM M03re H ceHcopHblX
ueHTpOB B r0J10BHOM 03re), TaK 1'1 MI40reHHble (31\.\r CKeneTHblX MblWU, 3Kr
cep;u.{a H 3ff r.na.n.KHX MbIWU).
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n.n., perHcrpHpyeMLIH B tleKOToport TO'iJ<C OKono B0J10KHa OTHOCHleJlbHO 6eCI<OHe"l-
HO Y.!laJ1eHflOH TO"lKJi (ee MOllenHpyeT pCcPepeHTHbiA 3JteKTpOA, norpy>t<eHHhlH B Ka-
MCpy C BOnOI\IIOM). YBenli1.IHTcR. TOT >Ke pe3YJlbTaT J].ocTHraeTcR npH np8KTH1.JeCI<H
nonHOM YMeHbllJeHHH 06'beMa pacTDopa. OKPY>K aI0 1l1erO HcpBHoe BOnOI\HO. B Ka>K-
AOM 1f3 Tnx cnYl.JaeR o()w.ec BHeK..'1eT01.IHoe COllpoTHDnemie TOKY, reHepMpyeMO"dY
6HOJ1eKTpH"IeCKliM HCTO"lHHKOM 6y.aeT 6onbwe, H Torlla no 3aKoHY OMH pcrHcrplt-
pye\tblH nOTeHUHaJ1 yneJ1WlBTcSI.
B TOM c.i1)"-1ae, Korlla HCTO"lHHKOM CHruana flBJtReTCH HCpDtlLIH CTBon C He-
CKOJlLKHMJ.f TbiCRl.Ja'dH OAHOBpCMellllo B030Y>f<Jl.eHHblX HCpBHblX BonOKOH, 3KCTpa-
KneTO'IHoe nOJle B oJ.Hopol1nofl OeCI<OHe1.JHOH cpe..1c 6YJler TaKHM if<e, KaK H Y 0.111'1-
H01.JHOrO BOJ]OKHa (pJ.fc. 4.5). I3HeKJleT01.JHblH 3J1eKTpll"leCKHH nOTeHllHa..1 RBJlHCTCR
S'" + S,
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4.3 «!>YHKLU10HAnbHAfi CTPYKTYPA nEPJ.1<IJEPH4ECKOH
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(6)
s.
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c1>YIII<UUOlldJ1bHOH 3.rleMeHT0\1 HepBHoH CIfCTeMLJ HB.1S1eTCSI pecf17.eKmopHaq iJya
(paic. 1.6. a), BK.rlf01i8JOUl.aR B ce6A c..1e;tylOUl.ue KOMnOlleHTbl: (I) OpcUH 'lYtJl:ma, co-
CTORlIlJfH 113 peuenTopOH onpeJJ.eJleHHOH Mo;xanbHOCTH. pearJfpYIOWHX Ha TaKMe CTHMY-
JlbI OKpy>+<afOU1eif cpeJ1bl, KaK .n.aB..1CHHC, TemepaT)'pa, npJ.lKOCHOBeHHe II .llpynte; (2)
n}'\um ceflcopHbLX HepoHbix 00 ?OKOH (ItHOr..W aHaTOMHlleCKH 06'be!I.HHeHHblX B ceu-
COpHhlii lIepB), nepe,J::IJOIllHX HHcpopMaUHtO, 3aKollHpoaaHHYJO l.JacroToH c.neJJ.OB8HHR
Hepomx li\fnyJ1bCOB (noTeHllH3.'10B .n.eHCTBJlR. 1111), OT ncpHcI>cJ.Ht'ieCKHX ceHcopHhlX
pt:llenropoA I< J1PyntM HCpBlfblM K.1eTKa\1, pacnOnO>KeHHblM B UHC (rOJJOBHOH H cnHH-
HOH \103r); (3) IlHC, KOTopaSJ IIrpacT pOJlb u.eHTpanhHoro HHTerpnpYlOllLero OT.n.en3.
OLtfllHBatOl1lero nocryn81OUIylO uucpopMauulO U IIPH Hco6xo1.HMOCTH Bblpa6aThiBafome-
ro «.n.nuraTeJJLlloc» pemeHHe B BHlle noc.ne.n.on3rCJ1bHOCTH DB. B MOToHeHpoHax: (4)
ilellt!anze.1lJHOe HepOH.Oe nOIlOKHO (aI<COH MOTolleHpoua), CBH.iblBalOw.ee UHC C ne-
PH(j>CpH'leCKOH (cKeneTHOI-I) 'dbIWu.eii; (,5) lJCnOAHUmcAbHblu OpcQH (B JlaHHOM npHMe-
: BOJIoKHa CKe.neTHOH MblWULI. KOTopble co....pawalOTcSI B OTBeT lIa lIepBlibie IIMnYJ1b-
, JlOC1)'lIafOU1Hc no )).BHraTenbHblM aKCOJ-la).
:J npocreHWHM npHepoM pecpJleKTOpHoH .n.yrH HB.'1ReTCH KOlleHHbLu pecjJAeKc.
, TOT he rJ "'K C B
t' 't'o c bl3bJDaCTCH JICrKHM y'tapOM no nOD.I<OJlCHHOMY CYXO>KHJ1HK) HU)k.C
ko.fJCHa n p
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MHHI..I 1Ia3
, blBaeMblC _4fbLwe'iHbiAt/l oepemeHQAt.U. 3TO B036Y>K!1eHUe BbJ3b1BaeT
MHOrO'iHCJI
8 eHHhle uepBHble HMnYJlbCbl, 1\0TOpble no CCHCUpHbIM (acpcf>epeHTHhIM)
OOKHaM nocTV n 
P 'J alOT B CnlfHHOH M03r H JI.a_'1ee nepe.llafOTCR Ita 'dOTUHCHfJOHb1. B
e3YJIbTare .
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!tau' J lOr CHrHanbl, KOTopble Bbl3b1BafOT ec COKpdweHHe H npHROJ1.RT K pe3KOMY
TH it<eHHfO KOflC"IHOCTJf. 3TO .t CCTb XOpOIlJO 113BeCTllblri KonCIHlblH petPJleKC. OTMe-
. "ITO Ha
O T Lj3nbHbiM CTHMY.'lOM ,lL'1SJ MblWUbI nOCJ1Y>KH.no P aCTR>KeHHe Tor,!la KaK
ro ·
RBHJIOCb ee cOKpaw.eHHe. 3T;:I npOCTaA pe4>J1CKTOpHBH ilyra HMeT "'Horo
(a)
[kH'TJ'C11Uollbi n ...
KUI1CWUX
nop'Ja.I.HhlA
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Cc.Ia.1MU1HhlA
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(r)
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PMC. 4.5 BneK.1eTO'lHOe OTBe.n.eHHC n.n.. BBelUleTO'IHble nOTeHUHanbi 3apenrCTpHpOBaHbI B 06"b
CMHOM npODQJ1I11IKe He;J.aJleKO OT nOlJCpXItOCni R03y>t<.l1eHHOrO CCAdJUll.lUiorO Hepea IDt8'1eTpOM
MM.lLnS. y.'1Y'lweHHR OTHoweHUH CHrtlaJI/WYM 6bU1H ycpeJl.HcHbi 128 OTP..eTOB HepBa Ha .:JneKTp'-
LfeCKuii CTHMyn. COCTaBHm'. Illl perHcTpHpoBBJIH (a) OT OJJ.HoRpeMeHHo B036Y>Kl1CfUlbiX J1Bllra-
TenbHoro Ii ceHcoVHoro CTBO-'10B (5.. + S.), (6) OT H36lipaTeJ1bHO B0.36YWJ.lCHtiOrO ./J.BHraTMbHoro
CT130Jla (5",). Ii (0) H36Hp8TcnbHo BOJ6YH\llf'HHOrO ceHcopHoro CTB0J13 (S.)

144
4 npV1PO.QA 6V10nOTEHl..\Lt1A1l0B
4.4 3flEKTPOHEVIPOrPAMMA (3Hr)
145
cxonHoro C OTpUUaTe'1bHOH 06paTHOH CBS13bIO, D KOTOpOH ynpaBnHCMblH napaMeTp
RRJ1ReTCH .:VJHHO MblU1Ubl (rlfc. 4.6. 6). U !l.aHHOM CJ1YlJ3e UIIC .aeiicTByeT KaK
3leKTpoHHhlH peryJ1snop, MbllllettHoe BepeTeHo KaK .naT'mK LlJIliHbI, HepBIiLte DO-
JlOKH3 KBK npoRo1HHKH CHrHaJlOR neTJlIt o6paTHo-A CB3H. 3 MbJWua K3K 06"beK1'
ynp3RJ1eHHSI.
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nOCKOJIbK)' B CHHanCf npOHCXOllJ1T ..1HrpY3JUJ ..,e.D.HBTOpa OT npeCHII3nTHlJeCKOH
MfM6paHbI K nOCTCHflanTH\feCKOH. 3TOT Me!lJIfHHbJii npouecc aaJlep)f(HBaeT pCJcnpo-
CTpaHeWfe HepBHO-MbJWC\fHOrO CJfrHa.na. Ewe OIlHa 3cuep>KKB 8 peaJ1H3BUHH .llBHra-
TcnbHOii KOM3HJlbJ CBS13ana C .ian33.!1.bJB3UHCM MeXaHHlJeCKOrO COKp3w,eHHH OTHOCH-
TenhHO H3LJ3J1B 3J1e1<TpHlJeCKOH BKTlfB3UHH MbIUIU.. ha 3an.ep>KKa. HaJb1BaeMaH
opeMeHeM 8036Y:JK:aeH.uJI- cOICpaUfeHUJl. S1BJIHeTCS1 CBOHCTBOM CaMO" M bUll LlbJ.
npH nORTopHOH CTHMYJlSUlJUf MeXBHHlleCKHH O'rBeT MbJllIUbi CYMMHpyeTCH. ECJJH
blWUY CTHMYJIHpOBaTb HB BbJCOIUfX \f3CTOTax, TO ee OTBeTbJ CJ1HBafOTCR B eBHoe
JL,lHTeJlbHOe COKpaweHHe. H33bIDaeMoe memaHgcoM (H.IJH memallUlI.eCICU.Af CO-
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(JinTCHTHoro nfpHon.a) nOSlBJJCHHH BbI3B3HHOro 3J1CKTpHllfcKoro OTBcra II MbJUJue. HepB CTHMVJlH-
PYIOT BllBYX paJHblX Y'1;!CTKaX C HJBfCTHbiM paccTmm.leM D Mf' tlHMII. -
CJ.1HAnTH4ECKAJI nEPE.QA4A
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Toro, IiMCIOTCSI H KOHTBKTbI Me>t<.ny HeHpoHaMH Ii \fbllllelIHbJMH B0J10KllaMH. 3TH
Hep8H.O-.AfbiUle r iHble CUHanCbt HaX03HTCfI B H6oJlbWHX cnellHa.1Ji3HpOB3HHhlX 06
JldCTMX MhlllIe4HblX BOJtOKOH, H33bJBaeMLlX KOHqeool; n.aCmlJ.1iKoii.. B KB>K.QOM C}f-
Hance nepCJl3"ia CHrH3JJ3 OCY1..l1eCTB_'lAeTCst ;.nCKTpOXJfMUYCCKUM cnoco6oM. IIpH
!l.enOJISlpH3al1lHl npecHHanTHliecKoro BOJlOf\Ha Bbl)1eJ1S1eTCS1 cneuua.1bllOC BeweCTRU,
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al{emfJAxo:UJH (Ach), KOTOpLIH .D.1-I4>CPYHlJ.HpYCT liepe3 Y3KYfO cHH3nTH"iecKYfO Ulcnb
WHpHHOI OKO.nO 20 HM. 3Ta We_'lb 3anOJIHCHa 06b1"lHOH MC}f{KJlCTOliHO.' }f{HJJ.KOCTbJ(}.
npH ,.!J.OCTH>KeHHH HeJipo\feJI,HaTOpoM nOCTCHflanTtp.ICCKOH MCM6pCHfb1. OH B3mfMO-
.a.cHcTByeT C paClIo.o'IO>KeHHhlM Ha HeH peLlenTopoM, lITO BbJ3blBaeT KpaTKoepeMeH-
HYIO !l.enO.'U1pU3al1ltlO MCM6paHbi H rCHepdUHtu n.IJ., KOTOphiH p3cnpocTpaHHeTcst no
Mblwe4HOMY BOlIOKH}' OT KOHuesof! nJJaCTlllIKJL npouecc 3J1CKTpOXHMH"iCCKOn ne.
pcn.a"iH 8 CJ1HCJllce XBp3KTepH3yeTcH 3a1.ep>KKOH nopSlllKa 0.5-1.0 MceK. nOJlpo6.
HOCTH Me>KHf'HpOHHOH It HeprHlo'MblIllClIIiOH CJltlaIlTH\feCKOH nepeJt.B4H H3JJO>KeHbI B
Klllire Kandel ef al. ( 1991 : rJJaBbJ 9,10 H 11).
4.4 3nEKTPOHEHPOrPAMMA (3Hr)
CKOPOCTb npol:Jen.emm B nepHrpepHlJeCKOM HepBe H3MepHJOT npH nOMOWH ero CTH-
MYJUIUHH B ruyx Y'tBCTKax, paCnOJIO>KfHHbJX Ha H3BeCTHOM paCCTOHHH npyr OT .npy_
ra. Pa3HoCTb Me}f\JlY .!JJIHHHOi, }I KOpOTKOJ1 Jan.cp>KKaMH perHCTpBLf.HH nOTellLJ.HaJl3
J.eHCTB.fH (.naTEHTHblMH nepHOJlaMH) .a.aeT BpCMH npOReJleHHSI no YliaCTKY HepBa
t.1 e >K!J.y CTHMYJ1HPYIOW,HMJf napBMI-I 3JJeKrpOlloB (pIfC. 4.7). 3H3R paccToS1HHe Me>KJJ.Y
HHMU Mbl MO .
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146 4 nPIIIPO.QA 61110nOTEHULo1AnOB
4.4 3J1EKTPOHEPOrpAMMA (3Hr) 147
me oco6ble 3<1>cpepeHTHble aKCOHhI HCrCTan1BHOH HepaHO" CHCTeMbi (HBnpHMcp,
cocy.'J.oJJ.RHraTenhHhle). nOHHTHO, TO npH 3neKTpHlJeCKOM BO.:l6YMW.CHHH Deero
HepRHoro CTRona COCTaSHOH flll HB..'UleTCSI Cyt.iMOH HH.n.HBH.n.yanbHblx nll, reHepH-
pyeMblx pa3HblMH 3KCOHBMH.
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BCHHO OT CCHCOpHblX aKCOHOB npOH3BonbHoro HcpBa (HanpuMep. OT HKpOHO>KHOrO
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n.aeT ..1BHraTeJlbHbIX B0J10KOH. HCCneJlOHdHHSI :1KCTpaKneTOIJHblX n}J. ceHcopHblX
HepOOB HMelOT CYlU,CCTBCHHoe 3H3'iCHHC .LU1.H lUlarHOCTHKH nORpC>K.L1.eHHH nepH<p-
pHlfeCKOH HepBHOH CHCTCbl.
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nOnbJYIOT CKOpOCTb Ii 33..'lep>KKY npOBeJJ.eHHSI nIl. B TO >he BpeMSI, Da>KHOe 3HalfC-
HHe HMelOT H xapaKTepHCTHKH 3KCTpaKJIeTOIIHblX MblWCIIHblX nIl, Bbl3BaHHbJX B
MblUJue THMIi HepRaMH. npH HCCJ1ellOSaHHH Bbl3BaHllbiX MbJUJe'IHbIX nOTeHllHa.rIOB,
HX .l1J1HTeJ1hHOCTb MOiKeT 6bllb BeCbM3 HllcpOpM3THBHb1M napaMeTpOM. D.eJIO 8 TOM.
liTO 3aMe)V1eHHe npOBO.l1HMOCTH 8 HeKOTopblX ABHraTeJ1bHblX HepAax MOiKCT "pH80.
)J..HTb K Jdlld3AbJB3HmO H paCCOrJ1aCOBaHHIO aKTHBaU,HH lJaCTH MblWU, TaK '1TO pe-
3Y.'1bTHPYIOIllHH 3KCTpaKJ1eTOlIHblH nIl MOiKeT CTaTb 60Jlce npO.n.OJ1>KHTenbHblM H
Mlloro4>a3HblM. B non06HhlX CJ1r«aMx aMrIJlHTYlla COCT38Horo 3KCTpaKJ1eTOIIHoro f11l
HepBa MO>KeT 6blTb CyulccTBeHHo CHH>KeHa H3-3a pa36poca 80 BpeMeHH HHD.HBH!l.Y-
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1lJI. B ptU10He 33nJJCTbH HMeeT TPcxCPa3HY rt-.; p H, pacno.'IO>KCHHblMH H3 JBIJHCTbe H OKOJ]O J10KTR.
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se.'1lillHHe Ii cpopMe OTBOQHMLlX COCTaBH bI )"13 n.n CTKax OTBe.i1elHfH, CJIe.nyeT npn.:JHaTb. lITO pa3.1HlJHH B
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BocnpOH3Ben.fHO c pa3peweHHH aBTopOB no KHHre' JAR L .
tromyograplzy, 2nd ed., Philadelphia: Lippincott, '19:'7 . enman, A. E. Ritchie. Clinical Elec-
TKaluf. pacn0J10if<CHHblM \te>KD.Y 6
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nepen. YCT3HOBKO.1 3eKTp:IOBH::eHHfl cnpOTHBneUHSJ .1JIeKTp.flJeCKoro KOHTaKTB,
B >KY Heo XO!UfMO 06e3>KH pHTb (paJn.cJI 5 5)
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c nOMOIlJ,bIO HH3KOW MRUlHX lH'" TOLiHbJe nOTeHllHanbJ penicTpHpYIOT
3Ha'leHHSI KO"" y 't'4>epcHuHalILHblx YCHmneJIcfl, HMelOw,HX BblCOKHe
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PHC 4 8 Ul(UeHm oC/la6.'leIiUIl Cll/1t/Ja3Hoo CUcHaAa KOCC ( p a3.!len 6 5 )
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, CCTeRaR H3BOD.K3 He nOJIHO T 10
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aij HanOliKa OTpaif<aeTCR . 3TOM pHCYHKe ceTe-
pa3JfeCeHHbJX no B CMeH:a:apa3J1Wme Ha'lBJlbHOro H KOHeilHoro ypOBHeft KpHBblX,
MeeK). p nOJlynepHO ceTeaOH CHHYCOHanbHOH BOJlHbI (10
llJlH CHHif<eHHSI HaBon.OK H MOn. .... .
6eJIH coen.HHRIOUlHC 3J1e H'i'HUHpYIOw'HX BJIHSlHHH (rJl3Ra l) BXO.!lHble Ka-
6JJHe n.pyr K n.Pyry H 3KKT:afJlHreJIeM, c.nc!l.yeT pacno.'IO>KHTb KaK MO>KHO
pa3HO nOMeCTJ.fTb B 3KpaH OB ' B HeKOTOpbJX CJIYlJaHX naUHeHT3 ueJ1ecoo6-
CJtrHaJ1 CHJIbHO JiCKB>KeH xa o P HHYIO KO\lHary H.fJH Ka6HHY. FCJ11f perHcTpHpyeMLa'1
J{ THlieCKHM WYMOM (lfanpHMe p
OOMHblX YCHnHTeneH) TO · TenJIOBbiM WYMOM 8h1CO-
!JCpeaHeHue CIlr?HU/lOO' (pa3: ;)HlfeHlfSl OTHOmeHIJSI CJffHaJl-WYM npHMeHRfOT
3KCTp3KneTOlfHble OTBeTbI (cocma{JHbie nomeHL4uaIZbl. oeiicmauJl) ceHcopHblX
HepBOB MOiKHO nerKO H3MepHTb...IJ1H qJAHlHtOrO HJ1H ..IJOKTeBOro lIepBa PYKH. C 3TOH
l.leJIblO Ha 113J1bLlbl Ha!lCaaIOT KOJ1LUCBbre cnfMY_l'Jl1pYIOIllHe 3J1eKTpOD.bI (pHC. 4.8).
PerHcTpauHH 3KcmpaK.1cmOtlHbJX norneHl{fJGilOO OT .!I.BYx r«acTKoB, paCIIOJlOiKell-
HblX en.OJlb lIepBa Ha H3BeCTHOM paccToSiHUH Jlpyr OT Jlpyra, n03B0J1S1CT oueHHTb
CKOpOCTb npOBe!l.eHHH no ceHcopHblM BOJIOKHdM. B CJ1Y'iac J10KTeBOro uepBa (OH
npoeWipyeTcSI rnaAHhlM OOlM30M HB rpcTHH H lJeTBepn.1 naJ1bUbI) 8bBBaHHblH n.n.
MomHO perHcTpHpOB£JTb OT pa3J1HlJHLlX \-teCT BLlOJIb TpaeKTopHH Hepa8 BnJIOTb AO
nOLlMblweliHoH Bnan.HHbl. B CJ1ae cpeAHHHoro HepBa (npoeuHpyeTcSI Ha 60J1LWOH
H YKa3arenbllblH naJ1bUbl) l III MO>KHO perHcTpHpoRaTb OKono HepBa H Bblwe JlOKT".
CnlMynHPYIOUlHe (ocoOeHHO JJ.J1HTeJlbHhle) H\tnYJIbCbI MorYT Bbl3blBaTb HC
TOJlbKO Heo6xoJlHMOe R03t1Y>KJJ.eHHe HepBa, HO H npHBeCTH K COKpallleHHIO Mh/JUUbl.
n.BH>KeHHIO KOHeqHOCHI H n.pYI HM HCiKCJIaTeJ1LIILIM noeX3M. 3TU nOMeXJ1 OTpB>Ka.
IOTCH Ha perHCTpaWfH nn. H Ha3blealOTCSI apmeljJaKmO{ cmUMYJ1Jll{UU. ApTC4>3KT
MO>KHO CHH3.Hb HJ1.1 nOJIIIOCrLIO YCTpaHHTL nYTeM KOMcpopTHoro H pacCJJB6J1e1lHOrO
pacnOJlO>KCIIHSI KOHellHOCTU, a T3K>Ke 6J1aroD.apSi npHMeHeHHtO KOpOTKIiX H HHTeH-
CUBHblX eTJfMynoB (npSlMoyroJlbHble HMnYJ1bCbl OKOJlO 100 B 3MnJ1HTyn.o H H JI)1H-
TenbHOCTblO 100-300 \tKceK). TaKHe HMnYJlbCbI B036Y>Kll3IOT KpynHble, 6b1CTpO-
npOSoD.HUlHe CCHCOpHblC HCpBHblC BOJIOKHa. HO He 3KTHRHPYlOT TOHKHC 60J1eBblC
B0J10KH3 .UHf coce.Jlllfe MblWUbJ. 4T06bJ YMeHhlllHTh apTc4>aKT CTHMY_I'JS1UIfH. npli\4e-
HstlOT H30JlHpYIOw.ee YCTPOHCTAO (H30nHpYKHUHrl TpaHCQ>opMaTOp, D.HOD.HYIO MOCTO-
BYIO u,enb, onnilieCKYIO pa3BSl3K}'. H T. J1,..). TaKoe yeTpoHcTAO H3o.ulpyeT (<< raJ1bBa-
ItHlleCKH pa3Bfl3b18aeT») 6HnOJlflpHbie CTHMYJIHPYIOWHC 3J1CKTpOLlbi OT 3eMJ1H (aT
o6wcI'u IlpUBO.ad). Te\l caMbiM nyTb CTlIM}lJlHpytOlllero TOKa orpaHIfHBalOT }"fBCTKOM

148
4 nPIIIPOAA 6l.10nOTEHV1AnOB
CKOPOCTb npOBEAEHHJt no .QBJ.1rATEnbHOMY HEPBY
HJMepeHU5t CKOpOCTH npOBen.eHHst no )J.BHraTenbHOMY HepBY in viv? MU>KHO BbI:
nOnJlHTb TaK, KaK nOKa3aHO Ha pHC. 4.7. IlanpHMep, Mano6eplloBblH HepB neBOH
IiOfH MO>KHO cTliMynJtpoBaTh n03BD.H KOJleHa (npJ1 O)J.HOM 1-I.{MepeHHu) H n033lUi no-
l1bl>KKH (npH JlPyroM H3MepeHHH). npH 3TOM nil perHCTpHpYIOT nOBepXIIOCTHblMH
HnH JtrOJ1bLlaTblMH J1eKTpo.aaMH. pacnOnO>KeHHblMH Ha CTone.
PEC!»nEKTOPHblE 3KCTPAKnETOllHbiE nOTEHLJ.HAnbl
4.5 3nEKTPOMVlorPAMMA (3Mr)
149
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HOro no.:u<OJ1eHHOrO HepBa (06 YBe.rJHemm aMnJlHryl1.bI CTHM)'J13 MO>KHO CY..1liTb no pdcryllleM)'
apTeq:.ar\1)'). n03;uun1 nOTeHl.tHa.n (II-BoJIlla) stBJJHCTCH HH3KonoporoBhlM OTBeTOM. OH 1l0JiBJIfleT-
01 np:1 M3J1oA CH.'1e CTHMYJ1ltpytOlltero TOKa. KOTopaH He.zJ.OCTaT04H3. IIT06b1 Bbl3BaTb MblweHblH
OTBeT (M-BO.1HY). YBe.'1HeHHe aMnJliTY.llbi M-BOJJHbI cOnpOBO>K.'J,aeTCA CHH>KeHUeM H-BOJIHbI.
BocnpoH3Be.aeHo C pa3pewemcsI aRTopoB no KHHrc: J. A. R. Lenman. A £:. Ritchie, Clinical Elec-
tromyography, 2nd ed., Philadelphia: Lippincott, 1977.
B HeKOTOpbJX CJJY'iaSix npH CTHMYJ1SIUUH nepu<l'epl14eCKOro HepBa H )KCTpaKJleTO'i-
Holi penlcTpaUHu nll B paHoHe MbnllUbI, HHHepBHpoBaHHoH .!laliliLIM HepBoM, MO>K-
HO H86nlOn.aTb nOflBJ1eHHe BToporo nOTeHUHaJld, KOTOpblH B03HHKaeT n03.nHee
06b1tJtlOfO Il1l, cooTBeTcTBytow.ero npHxon.y CHfHaJla no HepBY OT CTHMY'IHPYIOLll.Ii}'
3neKTpo;t.oB. EcnH y3CTOK CTHMYJ151UHH lIepaa npJt6nJi3HTb K MbIUJu.e, TO 38D.ep>KK8
nepBoro nll YMeHbwaCTCSf, a aToporo 1111 ysenH4l-tAaeTCst. TaKaH oeo6eWIOCTb
331.epiK3HHoro nll CBHl1.eTeJlbcTBye-T 0 TO", 'iTO BO.:l6Y>KllCHHe Hepoa CHa'-lana pac-
npOCTpaluteTCfl B npOKCUMa 7bHOM Hanpao..leHUU aT CHIMYJ1HPYIOWHX 3neKTp O .!l.OB
K UHC, a 3aTeM B03Rp3U{aeTCH B ducmaAbHoM Hanpae'leHllU o6paTHo K Mblllille.
Be rJJi1rHHa 3aJl€'p>KKH BToporo OTBeTa HBJ1S1eTCfl ononHe D.OCTaTO'iHOH JJ.,l1H Toro, 'iTO-
Obi B036Y>KJleHHe pacnpOCTpaHHnOCb npOKCHManbHO no ceHcopHblM BOJIOKH8M AO
cnHHHoro M03ra 11 BepHYJ10CL 06paTHO no D.BIUaTenbHblM aKCOH3M. .npyrHMu cno-
8aMH, 3a.n.cp>KaHHb1H rill UMeeT pecpneKTopHYIO npHpo.n.y.
ECJUI cTHMynHpoBaTb 33;t.HHH 60Jlbllie6epUOBblH HCpB lIorn, MO>KHO Bbl3BaTb
n03..lHHH nOTeHI{H8J1 OT TpHLJ.ellCa HKpOHO>KHOf. MblWLlbi (puc. 4.9). 3TOT 'lJIHHHOJla-
TeHTHblH OTBeT 06J1a.n.acT HHJKHM noporo'l U Ha6nlOJlaeTCR npH CTHMYJIax. HHTen-
CHHHOCTb KOTOpblX 3ua'-ntTe.lJbHO HJ1>Ke ypOBHst. Heo6xoJlJ.IMoro !UUl nOSIBJ1emm
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trom . . C pa3peWeHIiR aBTopOB no KHHre: J. A. R. Lenman, A. E. Ritchie, Clinical Elec-
yography, 2nd ed.. Philadelphia: Uppincott, 1977.
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ambnto, DeLuca, 1984; Gerber el al., 1984).

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BOCnpOli3SeJteHO no I<Htre B. S. Llprndn. E. N\assie. R E. Kleiger. Clinical Scalar Electrocardi-
ography. 1972 c pa"peweHIiR H3.lJ,aTfJlbCTBa Yearbook Medical Publishers, Inc., Chicago
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penonflpluawUf. nOCJ1e LJero HacTynaeT OTHOCHTenbflO npo.nOJ1iKUTenbHoe TUzamo
deIlOAflpu3aL{uu. j])1HweeCH 200-300 MceK. q>833 KOHellHOH penOJlHJH1.iaU.HH ooc-
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tal   (',.,t,cal Re'lews in Bioengineering. I. No. I, p. 5, 1971 H no CTaTbe D Durrer t,t aI. "To-
HHJj c hon ofthe Isolted HUH1dn Heart,': Circulatiofl, 1970,41,899-912. npHBe.n.eHo C pa3perne-
P peW€'1II1R H3MTe.nbCTBa The ChemIcal RubberCo. u American Heart Association,lnc.

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C pa3pemeHHH H3.l1aTe.nbCTBa BOCnpOH3Bc.J.cHO no KJU,re A. C. Guyton. Strucrure and Function vJ
the Nervous System, 2nd ed., Philadelphia: \\1'.8. Saunders, 1972.
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tional Federation in Electroencephalography and Clinical Neurophyiology'" EEO Journal, I 58.
10 (Appendix). 371 -375.
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BocnpoH3Hc..1etlO no CTThe H: H. .j8S, G E P fi ld  C Erickson (Eds.), Springfield. III..
Epilepsy and Cerebral / DcaltzatlOn. \. . en Ie , . .
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LIS System. 2nd ed., Phllaof'lphia: W.B. Saunders. 1972.




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