061LUie nPuhujiIlbl IlOctpoehusi hepbhoo · chcteMbl
OcHoQHoii cTpyKTypHoii rr <f>yHKurroHaJibHoii e.n:rrHrrueii HepBHoii cuc1'eMhI
SIBJISieTCH neii.pon nellfon . B Ka)KnOM HeiipoHe pa3nIHlaIOT :reno If eepBHble oTpOCTKif - .n:eH)J:pHTbI c il:x peuenropHbIMH OKOHtiaHifSIMH If HeiipHTbl' (aKCOHbl) c HX 9cpeKTOpHbIMif OKOHtiaHHHMif (pHc• . 240).
Tena HepBHbIX KJieTOK B uee'rpallbHOM OT){eJie Hep.BHOH CIICTeMbI (rOllOBHOH H cnrrHHOH M03r) 06pa3y10T cepoe' .,uoscoeoe eew,ecmeo.- substantia grisea, ·a Ha nepmpepHrr - eaJ-te1tuu - ganglion (cnrrHaJibHhre 11 aBTOHOMHble - ganglium
spinale et 1mtonomicu m). , ·
HepBHble BOJIOKHa B ueHTpaJibHblX opraHaX HepBHOH CHCTeMbICOCTaBJIHIOt ornosy 6e1toeo Moscoeoeo eew,ec111.8a _: substantia alba - ti BhinoJIHHJOT ¢YHK· WHO npOBO;l{HIIKOB. B nepmpepn11ec1mM OT)teJie nepBHOH CHCTeMbl OHII BXO)tH'f B COCTaB Hep808 -nervi - Ii npOBO)J:HT HepBHble IfMilYJibCbl OT UeHTpa K nepH
<IJepHH (.zi:arrrare.nbHble, rr.TiH 34>¢epeHTHble, BOJIOKHa) HJIH, Hao6opoT, oT neprr f}:lepttn K ueHTpy ( 11yBCTBHTeJibHbfe HJIH aqxpepeHTHbfe BMOKHa).
Bee HeiipoHbl 3aK.'IJOtieHhI B oco6hrii ocToB - Heiipornrr10, o6pa3oBaHHYIO rJIIfaJibHblMH KJieTKaMH, Bb!IlOJIHHIOUlifMif aamHTHyIO, a B ueHTpaJibHblX opra Hax HepBHoii c11c1eMhl 1po<f>nqecKyro H onopHyro ¢yHKun0. B Heil :me npo-
407.
XO)JJIT H KPOBeHOCHhie cocy.n;h , OTl[eJieHHh e 'or Heii:poHOB Me3orJirreii:, SiBJIHJOIIJ.eH· c.si: npoH3Bu.D,HbIM Meao.n;epMbI. , 1 •
Peuer.nopHhre HepBHhie 0Ko,HI1aHHS1 >JyBcrBrrTeJJbllhIX HepBHhIX BOJIOKOH
(peueriTOpbI) BocnpHHHMaJOT BHellIHife If BHyTpeHHife pa3.n;pa:lKeHifSI If riepeaJOT llepBHbie IfMilYJlhCbI no .n;eH.n;pHTaM (peuenropHbIM, IfJIIf aqxpepeHTHbIM, HepBHbIM OTpOcTKaM) B TeJIO HeiipoHa. . • - . .
HeiipHT (a1reon, IIJU! epeH 1H_hlii HepBHhIH orpocroK) 6bIB1 eT TOJlhKO o.n;mf. 0.H nepe.n:aeT OTBeTHhie IfMilYJlbCbI If3 TeJla HeiipoHa qepe3 CifHanCbI IfJIIf
Ha .n;pyrne HeiipoHbI, HJIH Ha pa6ot1He KJleTKif (M"'1llit1Hble HJIH :lKeJJe3HCThie •
o
ne
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Ha3.SrnaeTC51
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npe.n:crnBJ15leT
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u
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ex
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51
Ha
neprr
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pacnoJ1araerc51
B
IieHTpaJJhHhIX
porax
cnrrHHoro
M03-
ra
IfJIIf
B
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Si.n;pax
CTBOJIOBOH
tlaCTif
fOJIOBHoro
M03fa
If
CBOifM
aK-
.eonoM coen:ttH51ercst c pa0ot1RM opraHoM..Me:lK.ZJ:Y. HHMH Haxo.n;HTCSI rperuii, TaK Ha3hIBaeMhlH BCfJU180'lHblU Heupon; . OH. o6ecneqrrBaeT nepe.n;aqy IfMilYJibCa c qyB
CTBifTeJibHoro
3BeHa
pe$JleKTOpHOH
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Ha
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06wqHo
B
v eT: r OMa.n;Hoe ftJ:IfCJIO HeiipOHOB (o.n;HH HeiipoH MO:lKeT KOH- raKTHpOBaTb CBOHM. .n;eH.n;pHTOM c 6o:nee qeM 4 ThIC. .n;pyrnx HeftpOHOB, a CBOHM Heil:pHTOM C 60J1ee lleM 27 TbIC. HeHpOHOB. . CT H TaKOM IlOCT OeH -
.n:y 803
y:lK.n;eHrre,
B03HifKlliee
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B
o.n;HOM
peuenropHoM
paiioHe,
nepe.n:aeTCH
6eCtlHCJieHHOMy
MHO:lKecTBy
KJleTOK
pa3JIHllHbIX
IfCilOJIHifTeJibHbIX
opraHOB, tlTO H 06ecnet1HBaeT KOOp.n;HHaUifIO IfX WYHKIIIfH.
PA3BUTUE HEPBHOfl CUCTEMbl
' l
Bo3HHKHOBeHHe
HepBHoii
CHCTeMbI
06ycJ10BJ1eH()
•
peaKTlfBHQCThIO opra-;
HH3Ma,
T.
e.
ero
cnoco6HOCTblO
BocnpHHifMaTb
pa3.n;pa)KeHifSI
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HepBHaSI
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o.n;HoBpeMeirno
COtleTaTbC51
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Mhllliell
HOH
¢YHKUHeii,
a
COOTBeTCTBYJOIIJ.a51
1cTpyKrypa-
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co6oii
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a3
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B03HH
pa3BHBaTbCSI y.:lKe caMocT051TeJJhHo, Ho B recHeiiUierr B3alfMOCBSl3H. TaKHMif op raHaMrr y npHMHTHBHhIX Kiflliet1HonoJ1ocTHhI1C (rrr.n:pa) cJJy:lKa't TPH 'rpynnw dre- urraJJH3Hj?oBaHHhIX KJJeToK 9nwremrn1n,Ho Mhlllle1r!1bl'e, ·t1yBeTBHTMbHO-Heps- ·
HbI If HepBHbie (pHc. 241).
. 3nHTeJIHaJlhHO-Mbllliet1Hhie KJleTKif BocnpHHifM3IOT pa3.n;pa:lKeHHe' H nepe
.n;aIOT ero. Henocpenc;xseHHO Ha MblWellHYIO qaCTb. 0TBeTHIHIpeaKUllH B@pmi<at
eTC51 B .ZJ:Bif:lKeHHif. .
llyBCTBHT_eJihHO-HepBHaSI KJleTKa IfMeeT .n;Ba oTpOCTKa, 113 KOTOpbIX O.ll:IfH .
-
o6pameH- so BHE'4nmo10 cpe,u.y (penenropnLif 1), a JJ;pyroft, 6oJ1ee ;zvrnHiihlfi, co-
e.n;rrHeH c . o6oco6JreHHhIMH Mhllliet1HhIMif KJJeTKaMH. OTBeTHaH peaKUHH Ha BOC· npHH51TOe pa3.n;pa:lKeHrre 3aKJIIOtlae'tC51 B MhllliellH:OM BH:lKeHHH:
Hepsiible KJieTKII paaMema,10TCS1 HOA 3HHTMMeM Fl CBOllMH OTPOCTKaMrr co- e.ll:IfHSllOtcsr c .3IlifTeJIHaJlbHbIMif H MblllietlHbIMif KJleTKaMH. 3a 'CtleT coe.n:rrlfeHifll Me:lK.ZJ:Y co6oii HepBHhie KJJeTKif 06pa3y10T 8u<fJcpyslible cnJ1.ellleHUJI,. ,
B npouece 3BOJIIOUHH cTpoeHHe HepBHoli crrcTeMhl yCJib:lKHHeTCH B cB513H
c ycJJO:lKHeHrreM ¢YHKUHif. Be.n:yrn.rrMrr ¢akropaMH npH 3TOM SIBJI5110TC51, c o.n;Hoii cropoH, .n:mpqiepeHUifaUHSI H KOHUeHTpaUH51 t1yBCTBHTeJ1hHl':\IX KJleTOK, ¢opMH.; pyiorn.rrx cneumprrt1ecKHe opraHbI t1yBcTB (peuenrophl), a c .n:pyroit cTopoHbl - pa3BifTHe MhlllietlHhIX "3JleMeHTOB. .
1 • DpHMHTHBHaH KOHl{enmpaq_ua HepBHhIX 3JleMeHTOB Ha6.nio.n;lJeTCH y:lKe y
HeKOTOpbJX Kiflliet1HOilOJIOCTHblX (Me,n;y3) B BH.ZJ:e HepBHOeQ KIJJlbtfa IlO KpaIO 30H· THKa. 3.n;ech :lKe y HeKorophIX If3 HHX pacn0J1ara10TC5l H cnetfua.llusupoeannble
408 I
1 J 2
.7
8
I I
Puc. 241. lf>ffJioreffes uepsuoii CffCTeMbl
A - cxeMa .zmcjJcj>ysuoii ue suoii cHcTeMbl r '
- smffTeJJff a.llbHOMb!IIIe'I HaSI K.11erKa; 2 - qyscr BffTeJJbHall uepsff asi KnerKa; 3 -ee peuenropnb!ii ff 3' -a<j>cj>eKrop'Hblii: OTpOCTKff; 4 - 9KTO,llepMa;
5 -M . .
7, - MeseHXffMa; 8 - 9HTO,llepMa; B-B - Kou ueurpauff ll uepsuoii cucreMbl (uaceKoMoe); 1 - ·· Ha,llr.llOTO'lffb!H ff DO,llr.llOTO'lffb!H rau·rJJffff; 2 ,.._
ff; - rpy,Auo raurJJffii; 4 - peueniopbl.
peqennwpbt:
rna3KH,
xeMopeu,enTopb
I
H
opraHbl
paBHosecmt
HaH6oJiee
pe3Ko
·
KOHU,eHTpauHSI
Bbipa)KeHa
B
opraHH3MaX,
nocTpOelIHbI.X
no
npHHlj.Hny
,OJ.J;HOOC·
HOCTif If .n:sycTopoHHeH CHMMeTpHH, cnoco6HbIX K HanpaBJieHHOMY nocTyiiaTeJih- HOMy ){Blf)Ke!'IHIO. y Hlf X B03HHKaIOT npo,n:OJibHble HepBHble CTBOJibl BIU{e nap HbIX HJIH HenapHbIX 6p10utmtx yMoeatnbtX t{enoieeK (npH HaJIHtllfH cerMeHTaUHif TeJia).ilpH 3TOM Ha roJIQBHOM KOHUe TeJia pa3BHBaeTC5! «fOJIOBHOH M03D c Ha.ZJ:· fJlOTOtlHbIM If no,n:rJIOTOi:IHbIM raHrJIIf51MH. HaiJ2Aomoq,HblU 2aH2Aut1 xapal<repH-
3yeTCSI CBOHMH CBSl351Mlf c TaKHMif opraHaMH llJBCTB, KaK 3peHHe, OOOHSIHH.e,
8 pe3yJibTaTe oil CTaHOBHTCH Ha,n;cerMeHTc:tpHbI M opraHoM, o6ecneYBB.aIOUUfM 6oJiee 6blcrpoe npose,n:eHHe B036Y*.ZJ:eHHH no sceMy TeJiy. fl"oiJ2Aonwi/,Hbli}, 211!l2· AUt1 cBH3aH c KHI1Iet1Ho:H Tpy6Koii. HaKol'.leu, y BbICIIIHX HaceKOMbIX aa cll'eT scex
-
p10I1IHbIX y3JloB <IJopMHpyeTCH 2pyaHot1 zaH2Aua, a Ha,n:rnoToqHbIH raHrnHii:, BbIIlOJIHSISI CJIO)!{Hyio ¢YHKUHIO, npHo6perneT If COOTBeTCTBYIOW.ee CTpoeHHe, BilJIOTb l1P IlOSIBJieHHSI B HeM accOtf uamusilblX lfeHmpoe. .HaJIIfqlie oocne,n:HHX cnoco6cTByeT ocyw.ecTBJieHHIQ CJIO)KHbIX HHC.TlfHKTOB, KOTOpbie 51BJI5IIOTCH pe- ' syJibTaTOM HcropHqeCKH CJIO)KlfBUierocH B3aHMo,n:eiicTBHSI opraHH3Ma ,c BHeUIHeA
Cpe,n:oii. C .IlOSIBJleHHeM Ha onpe.n:eJieHHOM 3Tane paBHTHSI 3aMKH'yroA COCJ.ztHC·
409
PHc.
242.
PasBHTHe
cnHnnoro
Mosra: .
I - uepsHall nJiacTHHKa; II - He!JBHblii >Ke.11060K i
/// - HepsHall Tpy6Ka; 1 - uenTpaJihHblii cnHH HoMosrosoii KaHa.11; 2 - DJI8CTHHKa noKpb!WKH; 3 -6oKOB8ll n.11acTHHKa; 4 - n.11acTHHKa OCH0-
B8HHll; 5 - nJJaCTHHKa )J.Ha; 6 - raHr.llH03H8SI
,nJiaCTHHK a; 7 - CDHHHOii M03r.
Toe cncreMbl H3 Me3eHXHMbl pa3BnsaeTCH cocy.n:ncTaH (cnMnarnqecKaH) HepsHaH
y xop.n:oBblX HepBHaH CllCTeMa npOHCXO)(IfT He TOJibKO If3 9KTO.n:epMbl, He nocpe.n:CTBeHHO IlO'ABeprarow.eii cH 803.zi:eiicTBifHM BHenrneii cpe.n:bl, HO, B03MO:lKHO, Ha 9HTO){epMbl (no ;n;anHhiM R . Remak, A. Tinel, C. H. 1"'\;ttBeeBoii, 1953) H H3
Me3o,n;epMbl, T. e. TaK :lKe, KaK n. y 6ecno3BOH6IIHhlX. B oTJinIIne oT 6ecno3so HOIIHblX 9KTo,n;epMaJll HaH HepBHaH CifCTeMa paaBHBaeTCH CTporo JIOKaJIH30BaH
HO If3 nepBHOH cnHHHOH nJiacnrnKn. 9rn JIOKaJm3aIUrn 06ycJ10BJ1eHa UCO- 6eHHOCTHMif CTpoeHHH npe,n;KOB xop.n:oBblX, IfMelOllJ,IfX Ha TeJie (3a IfCKJllOIIeHneM He60JlbllIOfO ero yqacTKaj xopoIIIO Bblpa:lKeHHbIH 3aIUHTHblH KO:lKHbIH noKpOB.
fa HeJ)BHOH llJlaCillHKll CHaqaJJa u6paayeTCH HepBHblH :lKeJio6, a 3aTeM HepB- HaH Tpy6Ka c ueHTpaJibHbIM IlllHHOM03fOBblM KaHaJIOM (pttc. 242). HeiipoHbl,
Pfl3BHBa10w.neCH B HepBHOH Tpy6Ke, CBOifMif OTpOCTKaMn -,n;eH)J:pnTaMn If HeH- prnaMn sc1ynaro1 B CBH3b co BCeMIJ 6e3 HCKJllOIIeHHH 9JieMeHTaMH opraHif3Ma. HepBHaH cncTeMa xop.n:oshlx (no3soHOIIHhlx) B ueJIOM cpaBHHTeJihHO c
6ecno:rnoHoIIHblMH xapaKrepn3yeTCH: ·t) BblCOKoii ,n;ncpqiepeHunautteii opraHoB
qy scrn (peuenropos); 2) nom1pHoH aposo.n:nMoCThlO B ueaH HeiipoHoB 6'1Iaro
,n;apH IfX CHHailTHIIecKHM CB513HM; 3) MifeJIHHif3aI.J,HeH HepBHblX BOJIOKOH, IlOBbl
marow.eii nx npos.o,n;HMOCTh; 4) MOllJ.HblM pa3BHTHeM Ha.zi:cerMeHTHoro opraHa s sn.n:e cemtf.W'llOeo 06pasoeaHU5l (formaho reticull!ris), a y MJieKonnrnrow.nx,
KpoMe Toro, B sg,n;e Kopb1 noJiyIIIapnii roJioBHoro M03ra; 5) rny6oKHM pa3Mew.e
HHeM HepBHblX KJieTOK B ueHTpaJibHOH HepBHOH CifCTeMe (y 6ecno3BOHOIIHblX
KJieTKH B raHfJIHHX Jie:lKaT Ha nepncpepnn, a oTpocTKH nx 6oJiee rny6oKo, qTO o6ycJIOBJIHBaeTCH oco6eHHOCT51MH cpopMnpoBaHHH HX HepBHOH CllCTeMbl); 6) pa3-
)J:eJieHHeM HepBHOH CifCTeMbl Ha u eHT aJibHblH OT,ll;eJI fOJIOBHOH Ii: Ciill HHOH M03f H nepn pnqeCKHH, B COCTaB KOToporo BXO)J:HT CnHHHOM03fOBble, qepeaHO M03f0Bble If sereTaTHBHble HepBbl; 7) HaJIHIIIfeM CnHHHOM03fOBOro KaHaJia c ero
pacwHpeHHeM. B ueHTpaJibHOM OT)J:eJie HepBHOH CifCTeMbl; 8) .u:opcaJihHbIM pac- , IlOJIO:lKeHHeM M03ra no OTHOIIIeHHJO K KifIIIeIIHHKY If xop.n:e. ' •
' .
U.EHTPAJl bH bl D OTJJ.EJI H EPB HO A CH CTEMbl
lI,eHTpaJibHblH
OT)J:eJI
HepBHOH
·cttcTeMbl
TOaorpacpnIIeCKH
If
cpyHKUifOHaJibHO
pa3)tem1eTCS1
Ha
CIII4HHOM
M03f,
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B
H03BOllOIIllOM
1rnHaJie,
H
FfrJIOBHoii
Mo3r,·
noMew,aiow,ttlics.i
B
noJiocTH
Hepena.
KaK
cnHHHoii,
TaK
·n
roJIOBHoii
Mo3r
nocTpoeH
H3
ceporo
n
6eJioro
M03roBoro
sew,ecTBa.
-
Cepoe Moarosoe seecTBo coeroH'f ns nepsin,ix KJie'foK M. nx , o'Ppoe'PKOB,
,
M03ra BOKpyr ueHTpaJihHblX M03f0Bb1X IlOJIOCTeii: l.(eHTpaJibHOro CHHHHOM03- roooro KaHaJia n *MYllOIIKOB roJioBHoro M03ra. B cnnHHOM . Mo3re cepoe se w.ecTBo MaKpOCKOilHIIeCKH BocnpHHHMaeTCH KaK CilJIOlllHa51 Macca, B ·TO BpeMH
KaK B fOJIOBHOM M03re OHO npe.n:crnBJieHO 60JibllIHM IIHCJIOM 51.zi:ep, COCTOHW,IfX H3 CKOnJieHHH H Cl.(enJieHHH HepBHblX KJieTOK, BblaOJIHHIOIUHX O)J:HOpo,n;HyIO
tPYHKUH. 11o.n:o6Hble cpyHKI.J,HOHaJibHble H,n;pa fHCTOJIOfHIIeCKH o6Hapy)KHBaIOT CH If B c;nnHHOM M03re• .UpyraH oco6eHHOCTb CTpoemtH fOJIOBHoro M03ra MJieKO·
410
. IlHTaIOIII.IfX Bbipa:lKaeTCH B TOM, ,IJ;TO B noJiyrnapHSIX 60JibllIOfO ,M03fa, a TaK:lKe B M032Ke11Ke cepoe Mo3rOBoe BeI.IJ.ecTBo 3a.1eraeT If Ha nepmt)eprrrr B BHJJ.e Kophl 6oJibllIOro M03ra If KOphl M03me1ma. Kope roJIOBHoro M03ra <fJYHKUifOHaJibHO 110.n;1rnHeHhl Bee rny6meJie:lKaw.rre oT.n;eJihIceporo Mo3rosoro BeIII.ecrna, KoTOphre B ueJIOM <flopMlipyJOT TIOJJ.KOpKOBOe cepoe Mo:rroBoe BeI.IJ.eCTBO, IfJ!If TIOJJ.KOpKOBbi e SIJJ.pa. . .
KpoMe KOpbInoJiyrnaprrii If· 110.n;KopKOBhIX H,n;ep, KaK B cnrrHHOM Mo3re, TaK If B fOJIOBHOM IfMeeTCSI cemttamoe o6pasoeanue, KOTOpoe XOTSI H . IfMeeT npHMif• nrnuoe crpoe1m:e, 110 BhlnOJIHHe 1 Be'CbMa sa21rnyro cpyHRII)1RJ. Cena 10e o6pa3o-
11,l:(y'I' B ueHlpaJlbl OH HepBHOH cncreMe B pa3HblX Ha11paBJieHllSIX. - npo,n;OJibHO If nonepe1rno, 11'ro no3BOJIHeT eii 6hITh cBH3aHHoii co BceMrr oT.n;eJiaMrr ro,n:oBHoro rr cnHHHoro M03ra. TaKHM 06pa30M, peTHKYJIHpHaH ¢opMaurrH npe.n;cTaBJIHeT co6oii HecneuH<IJHIIeCKYIO aKTifBifPYIOIII.YIO CifCTeMY M03ra, BKJIJOIIaSI Kopy noJiy rnaprr'ii (y MJieKOTIIfTaIOW.IfX), KyJJ.a HanpaBJISIJOTCSI BOCXOaRU{ue nymu peTHKY1:15lp
HOH <flopMaurrH. Or Kophl 3aBHCHT B ro :lKe speMH rr caMa peTHKYJIHpHaH <flopMa-
UifH 6Jiaro.n;apH HaJirr11rr10 HUCXOORU{UX nymea, IIJJ.YI.IJ.HX If3 KopbI B npoMe:lKyrn11- HbIH rr cpe.n;Hrrii o3r. •
y Hif3llIIf X l103BOHOII H&IX, He IfMeIOW.If X KOpbI, peTHKYJIHpHaH cpopMaUifH
HBJIHeTCH rna'BHblM n.eHTpoM Koop,n;HHauHH peaKuHii rr 110Be,n;eHHS1. Bo36y:lK.n;e Hrre B peTHKYJIHpHOH ¢opMaUHH npoTeKaeT .n;rrcpcpy3HO He3aBHCifMO OT IfCTOII HifKa H c11eumpHKif pa3.n;pa:lKeHHH. B cnrrHHOM M03re pernKymipHaH ¢opMaUH51 pacnoJiaraeTCH c 6oKoB OT .n;opcaJibHhIX croJI6oB ceporo M03rosoro · BeIII.ecrna If EOKpyr neHTpaJibHOro CTIIfHHOM03fOBOro KaHaJia_. CHJibHee OHO Bblpa:lKeHo
B rpyJJ.HO:ii If oco6eHHo B rneiiHoii 11acrn cnHHHoro M03ra. B npo.n;oJ1roBaTcM,