(Luận án tiến sĩ) vai trò của ức chế trong vỏ não đối với sự hình thành tính nhạy cảm theo hướng của các nơron vùng vỏ não cảm giác soma

BO mi

UQC M& SHUUG liOC QliMlil.

LmSliJ^

ZKUQtJU VAl aOC i-QlJU ilQP li/, i.Ui
MMSJA SMih HOC
M*

SKAK

S

4

»

N»

TI4IIU

AN

K.vai^4ri
VAI 'i'itO CWi 01; Uiii- *x»ivU.U VU i.^U i^Ul VOl
aU liHUl i^iW-H s u m iiiiiil C^k 2IAL0 iuWlku

Con bGS# i'ho t l d n s i 3)8 Cdc^;
iitfu

•^ I Mir Vf-rN

I

^u^'ah
^ng
^^

iiugD an p b o t;i^D s i kiiaa b p c

HA 1.01 -

ige?


- 2 -

Hp C

L y c
Treng

no

© 1U

Riaii
T(JNG

II

iiUAN

TAI

LlS;U

9-57

I - Cd s3 hinh t h a i cua cac qua t r l n h -&c che d ndroa

9-20

! • Te bao tiTunf^ g i a n Uc chc t r o n g vo nao
2 . GAM - chat t r i m g ^an TSC che t r o n g vo nao
5» Cau t r u e modun va cau t r u e t j p qu-szi
I I - Dan l i § u ve cd che t a c d\^t7 Uc che I a n nhau cua
ndrun vo nao*
I I I - Ciuan nifra ve t r u 3 n g t h y cam va t i n h c h a t
d e t e c t o r cua ndron vc nao*

21-26
:6-32

! • TrUdne; t h y cam cue ndron
2 , Tinh c h a t d e t e c t o r cua ndron vo nao
5 . Gc quan phan t l c h xuc g i e c v i b r i s s
17-

Dan l i g u ve ve;l t r o 5c che d o i vci

thanl-i tirjh c h a t cua ndron vo n o c .
Phan
PHt/OlvG

I-

b\? ulnii

32-57

III

PHAP

J«GHl£f^

Cl/U

Chuan "bj. d^ng v i t t h x n^igna

5b-51
38-39

I I - Kxch t h i c h hTi5n£j taji
I I I - Vi d i | n eye duu£. f?e gJii ho^it tLalx dign cua
ndron va dung cho v i di^n d i .
IV- Ghi d i § n t h e cua ndron va aru' l y cac k e t qua
nghien cUu.
!ã
2.

3.
4.

i)^g
Dtfng
Tinh
Tinh

histogram
vectd-diagrainjii
hf so nhĐy cam t h e o hudng
t h 3 i g i a n tiem t a n g

V- ChUdng trinlri t h y e ng;h i§iQ

30-51


« 3 Phan
K U T

(4UA

lY

Trang

NGfilfiN

Chx&jng

CtfU

I

52-111
3^-70

©AC T i m DAP Ul^G CUA I^OROi.^ VraG VO ISAO CAtl
GlAC-SOriA s o CAP ( S j ) C fxBO K ) I VOI KICH THICH
fcliia.'j HOC, KICii TKIGH Od HOC.
I - i>l§n t h e dap u^ng t r o n g vung c h i e u cua cd quan x u c
giac v i b r i s s .

5^-33

I I - Phan Ung c u a n d r o n vung c h i e u t r o n g vo nao cam g i a c soma Ichi k i c h t h i c h c a c s ^ i v i b r i a e myt CfiCh r i e n g
bift.

5'*—63

I I I - Phan tfng c u a n d r o n k h i k i c h t h i c h c a c sk h a c n h a u t r o n g t n r 3 u g t h y cam*

6^70

Chijfdng

71-'&4

II

bU BIETi iiOI Tlim NHAY CAM THH) EUDfiG CUA NOHON
DUOI TAG DVnO CUA K I C H T H I C K i>OI,Ai^£i HUONG CUA
Y£U TO TiiOI GIAN f>01 VOI SU BIfcdfi DOI TR&^.
I - By b i e n d o i t i n h n h ^ y cam t h e o hUdng c u a n d r o n difdi
t a c dyng cua k i c h t h i c h d o i .

72-76

1 . Sy t h a y d o i t i n h nh§y cam t h e o hUdng 5 c a c n d r o n
ng*uyen p h a t khong nlrify cam t h e o hiidng*
2 . Sy t h a y d o i t i n h nhgiy cam t h e o hUdng d c a c n d r o n
n g u y e n p h a t nh§y cam t h e o hUdng.
I I - I n h h^:fdng cua y e u t o t h 8 i g i a n d o i v d i s y b i e n d o i
t i n h n h ^ y cam t h e o hxrdng cua n d r o n .

77-60

1 . t?y x u a t h i ^ n g i a i d o ^ t'iam d a p v ^ g cua n d r o n
2 . Dien b i e n t h 5 i g i a n t r o n g s y b i e n d o i t l n i i n h ^ y
cam c u a n d r o n .
I I I - C y ^ t h a y d o i t h 3 i g i a n t i e m t a n g c u a p h a n x/ng k h i
x u a t h i ^ n t i n h nh§y cam t h e o hUdng d nhung n d r o n
von k h o n g nh§y cam t h e o hUdng.
ChUdng

III

.

YAl TRO CUA YEU TO KHOWG GI.J^ TBDm

tJU BIEN DOI

Tliai NHAY Qtili ThiiX; HUOriG

81-84


- 4 -

I - i n h hu3ng cua hvTdng k i c h t h i c h d i e u k h i e n d o i v 5 i
dap ting hu:ng*

85-87

I I - by b i e n d o i t i n h nh§iy cam t h e o hudng cua cac ndron
dtr^i t a c dyng cua k i c h t h i c h d i e u k h i e n l e n cac s g i
v i b r i s s co v i t r i khac nhau t r o n g trtf3ng t h y cam.

87-*00

! • Sy b i e n d o i t i n h nh^y cam cua ndron k h i cac s ^ i v i b r i s s
c h i n k i c h t h i c h d i e u khien phan bo d h a i ben cua
s^i
v i b r i s s chj-u k i c h t h i c h hutig.
2« Sy t h a y d o i t i n h nh§y cam theo hUdng cua ndron k h i t h a y
d o i khoang each g i 5 a k i c h t h i c h d i e u k h i e n va k i c h t h i c h
h&ig*

3» Sy t h a y d o i t i n h nh§y cam t h e o htfdng cua ndron k h i k i c h
t h i c h d i e u khien t a c dgng vao s ^ i v i b r i s s nlm khac hang
v d i fi^i chj.u k i c h t h i c h hxhag.
Chtfdng IV

101-111

TAG DU14G CUA PICROTOXIK i)OI VOI THUGITG THU CAii VA SU
II£I^ ^01 TI^iti KHAY CM: TilEO HUCMG CUA INCHON
I - T I C dyng md rgng trtfdng t h y cam cua p i c r o t o x i n d o i v 3 i
cac ndron vung c h i e u thugc cd quan nhjn cam tii cac s ^ i
vibriss.

101-10

I I - Tac dyng cua p i c r o t o x i n d o i v d i sy b i e n d o i t i n h nh^y 103-10
cam t h e o htidng cua n d r o n .
1 . ^ o i v d i cac ndron nguyen phat nhay cam t h e o htidng
2 . Doi v 5 i cac ndj^on nguyen phat khong nh§y cam t h e o hiidng
I I I - By b i e n d o i t h 3 i g i a n tiem t a n g cua phan Ung dxtSi
anh htfdng cua p i c r o t o x i n .

108-11

Phan IV

MJK
fipH
K£T

qui
112-129
I - Ve d | c t i n h dap ung cua cac ndron n h ^ thong t i n
112-116
tu' cac s ^ i v i b r i s s n&n t r o n g vung cam giac-eoma
sd cap ( ^ j ) *
I I - Ve s y b i e n d o i t i n h nhay cam t h e o hu5ng cua n d r o n vung
vo nao cam giac-soma dudi anli huSng cua k i c h t h i c h doi» 116-12
I I I - Ve anh ^vTdng cua yeu t o t h 8 i g i a n va khong g i a n d o i
v 3 i s y b i e n d o i t i u h nh^y cam t h e o hi/dng cua n d r o n .
120-129
! • by t i c h h^g ve t h 3 i g i a n
2 . Tich h;;p ve Idiong g i a n .
Ph^ V
K f T
LJJ I ^
130-131
Phan VI
~*~ Ll|nJ Thm
KHIO
132-148


« 3«
aiMi • ^

C^n^ v ^ «tf p^i&t t r i d a chuof: cuo 000 khog sinh
ly
bpc» trong nhtiiaf; nacs (^Jn day ainb l y bpc than iclnh da dgt
diHyc ahiitxg ffcanh tyu t o l 5 n , do phat b i | n m^t ©5 quy

tic
td cbi/c ha^t d^Qg cua nad b^ - ccf quan diSu khl^n
phtfc
tpp va h9an bao nh'u cua g l ^ i t y cbi3a«
ItbJmg; vin de cc? ban va co clnb of:.uyeo cic nhif
qua
t r l n h tfcb bffp cac lu^ne btfiJng cdD» vol t r S cua qua t r l n b
^c chl tronf bop^ <7<^Q£' pbon xp cS d l l u k l ^ , qu^ t r i o b hiob
th&nh V9 duy t r l t r i oh^,v*v«** tvt ISu iis Xitiu hut 3^
quan
tdm d§c b i g t cua I.P»Pavlov (47) vo ttbilu nh© cinb l y
bpc
oc'i tlluG c^ Mdn z^ cZ^r nfaif nbleu nlfi$c khgcô Cac aha khaa
hpc ViĐt nao cung c5 hiiijpnt; done f^op nboc d^rib trong
linb
vyc nay ( 1 , 2» 45, 6I)« Tuy v|iy, ban c b i c cti»a qua t r l n b tic
ebl« eui^ nbi/ cc^ chl y u l t billn^ phat ^ r i d n VB uac
dime
ttn:^Dg bd cua qua t r J n b nay v5l qua t r l n h ht?ng pbln oht? t b l
nao, t h l troo^' su^t o^t tbo^i gian d a l vln c ^ chiia
dif^^

Kit qua ngblln e^u cua nbiSu t^> t b i t a c g i a
tr^n
pbatD v l coSn t b i g l $ i trong v a i chyc naa g l a day da
chfing
mlnb v a i t r o bSt siic quan tr;mf; cua qua t r l o b i$c chl
dli
v5i toan bBô

trtr^c bic l a sy blnb tbaob cac quo t r l n b then kinb ccf
ban
dam bao cbo nao b<^ t;byc bi^o cbuc nang tbdri^- t i n «
dieu
kbl4n cua slnb ( l a , 1 9 , 39, 31^ 32, 51» ^^, 137, 155)#
Tbeo quan dilm cua o b i l u t;oc g l i f t b l m chl co
voi
t r o quyec djnb d3i vtfi sy blnb chanh va thay ddi cac trii&ng
thy cao chuyen boo cua cac oc?ron vmii; vo nao t h j g i a c ( 6 6 ,
^ ) » cvtii^ ahit viMig vo nao com ginc-sooa (9) vo do do da l a a
nlly sinb sy cln t b i e c phai »es xet l a i kh^i nigm kinb d i e o
vS trur&ng tby ca©t coi trtr&tig thy cam cua neuron l a tS ch5c


« 6 «
ctfoe a b l e , dtfcfc blab tbanb obo' sy guy t y co cbpn I p c ohtmg
loSng xung di^Dg btf5ag tS© d l tli cac td cb5c than klnh t h i p
ban (X6» I t ) *
Dfinir tho^i nhilu t o e gia cung t l o t b ^ fi»Si l i ^ o quao
ob^t c b i glua 5c cfc^ v5i qua t r l n b pbgn cich thdcg
trio
CM giac & ©5c neuron, d§c b i 0 t l a ay blnb tbanb t i n h c h i t
d e t e c t o r ^ cac otfron ims^ vo nao t b i g i a c ( 6 6 , 67, IJ^^t^iSS}
va vo oao cao gioo-^ona (9)«
Hgoai r e COQ CO y k i l n cbo rln{r ay b i l e dSl **mls deo*
trottg fahmp ©li quan b | tt»ng tac gil/a c5c acfron d l t p o ra
ay l i n h bo^t troog vo nao cung l a nh& qua t r l o b ^
cbl
( 1 6 , l&U
So vol t r S quan trpag nbtf v|iy nSn bl^n nay 5c

cbl
da trey tbanb vSo d l tbu but taanb ay cbu y cua n h i l u
nha
kbOQ bj>c» ^^§t troog ahtfng phtfctog pbap co hifu quo dSi v ^
vl#c t i e bl§a vai t r o cua 5c cbl l a pbi^mg pbap v l dlfn d l
(mlcroalectropboroa o ^ l i c a t i o n ) , d(^ CDC c b i t d^i
icbaoe
( o o t a g w i i t t a ) cua geiw»a-oa5ino-*utyric-axtt (UAES) vao
tl
bao tb§Q k i n b . £ | c qua ogaien c5u cbo t h i y ay pbong
toa
Se c b l Be Xao mit t£nb c h i t d e t e c t o r cua nifron (V, &3)
•
Bgi;PQ^c l a l , cung bSng pfaiJcmg pbap vl di#n dl 6tfe g l u t a ^ e t
vao cac ncfrtm aha cacb d i l o ^ l kboaog 200 diet 3CX> mm t b i
thay s u i t bi0n cac natron d e t e c t o r m^i, b o | c thay ddi tCob
nbay cof& da co t r u ^ dSy cua cac acrron d e t e c t o r (XO)*!ihllQ
tac gia cbo r i n g g l u t i ^ a t da boat hoa
cac nc^roa 5o
cbl
trung glaD va k i t qua l a ^ arixat bl0», bo§c tba:7 d d l
tlnb
nbay cas tbeo bti5ng cim cac t;l bao tbap*
ã I t l l p tyc tlâ b l l u val t r o 5c c h l trong vo oao d^l
v5l sy hltxb thnnb tfnh nbay c^a theo hvtm^ cua coc
nc?ron
vung cac3 g l a c - a o o a t cbune cbl d'a sii dyng is^t phtfcfng
pbap
DgblSo c5u kbac, dS l a phifo^ phop kich t b i c b d d i . Fbucme

|*8P nay cbo phep nghien c5u val t r o cua qua t r l n h 5c c b l
xulu bi§n dtf5l aab hating cua kicb t h l c b h«i5og tSa t y nbiSn
troD^ vi#c blnb thanb eac tinh c b i t cua oxyroo vung
cam
giac^aoffla*
Md blnb tbufin l y l cbo v i f c nc^bign c5u noy l a b# t b l o g
ttbfty cao xuc giac rSu eao ( v l b r l s e ) *
Val t r o cua 5o cbl trong vo nao dcac t l a b c b i t cuo cc^on vung c i s giac-aoma se co tbd
dt/;^
l a s sang t o tbdng qua cbt>ng nbifm vy cy t b I sau dSy s
1« % b l € n c5u d|ic t i o b dii? 5tm cm cac nc^on vut^ vo
oao Cora giac-«osia Bcr d p (Sj^) d l i v5i kich t h l c b t b i c b daog
voo cd quau nb^n cam v l b r l a a «
3m fbSn t f c b sy blnb tbanb t i n h nb^y casi tbeo
cna Dc^ron diiP^l aob btn^ig cua klcb t h l c b hi^^g tSs do
t h l c b d l l u k b l l o gfiy ra*

btidng
kich

% s i s b l l u anb bii^ag cua ylu t l t h ^ l gian d l l v^l ay
b i l n 66l t l a b obpy eioD t b e o hiring cua ac^on»
4* IPhSm t l c b d$c t i n h kbdug g i a n cua ay b i l n doi t l n b
nbay c ^ tbeo biiwg cua DC^CHI*
5* K ^ l e o c&u anb bifdisg oua p i c r o t o x i n ( c b I t d^l khang
cua GABA) d l l vSi sy blnb tbanb t i n h nbay cm tbeo btf^ng va
trt/^nc thy coea cua OCJ^OC vung vo nao cam glac-soiBo. qgam do
t h I y dtryc boo c h i t cua 5c cbl va val t r o cua no d e l v5i ay

blnb tbanb cac tfnh c b l t nay cua nc^on*
^*btmg k i t quo thu diiyc trong c6nt" t r l n b nghiSn
nay cbo t h I y rang t

c5u

- t5c c b l troog vo nao do kich t h l c b hb^mxe t § a t a o r e
^B lam v o l t bi§n bo^c Im thay d d l t l n b nhay eaa tbeo h\i3fne


• 8 »

cua fibtfn^ ncrron trong vun^ c b l l u cua cvlbrls«« S^TOog d l l u klfn klcb t h l c b dtfn d;^ d$c t l n b
hogo kbdng c o , ho|ic oo, nbi/n^; o' ni5c C^ khori^; cao#

nay

- QtQ? l u l t 1*ay ddl t l n b nbay ca^a cheo hu$tis cua no?ron
pby tbu§c vao cac y l u t S t h J l gian va khdcc gian cua
coc
klcb t h i c h *
«* v i oSt l y chuylt nbDng k i t qua tbu diiyc coo
gop
phan vao sy EBO^ r^n^ nbtfng kbal nl#9 hi#D nay v l c0 c b l
5c
ebl bdn trong vo nao, v l val t r o cua 5c chl d6i v&l sy blnb
tbanb cac dge tfnb cn5G nan^ i cac ncffron*

fflAH IZ

Bifn nay ^a co n b i l u t a l l l # u d l c$fi d i n sy t d ch5e*
ch5c n&tiQ cua vo nao, din t l n b c b l t oua nc^on va v a l
tro
cua 5c c b l trong vo nao* Dgc b i ^ t cac ddn l l f u v l
blnb
tbnl bpc da g ^ pban dang k l vao v i | c g l a l qii^lt cac
vln
di trSn.

3/^ CQ go mm m^i mj t^np gvA 'smn m qm
lA pbucrog dl§n blnb t h a i - td ob5c cus qua t r l n b 5c
chl i nc^rdn, nhaag vin d l difcfc cbu y l a cac c l bao
tining
gian 5c c b l tr^Mig vo nao, asrlt sacuMh-^Blno-butyric - c b I t
trung gian boa bpc cua irinap 5c c h l trong vo noo, cac c l u
t r u e sddun vo c l u czuc typ quan (anaeffible) cos n c ^ n *
3^* 2 l bSo trun^: fzimn 5c chg tram- vj a»o
clu t r u e v l bao vo nao da duyc nghlSn cuu tb
ISu ,
nbtfng y ngbia ch5c nan£^ cua tSuf^ l o a i te l ^ o vo v a l
tro
cua Chung trong «y hoji; d^nc t l c b hyp cua vo noo cbo
din
nay vSn coo dgng dtf^W! t i l p tpc tl® biSu*
Jtbl de cgp d4n vIn d l 5c cbl trwifi vo nao, cac
nha
e ^ l d n c5u cbu y d i n chib nang cuo t e bao blnb aoo» KhiSu
y k i l n cao r l n ^ dly l a nbufn^; ntfr<3n trune glen 5c ch5 da

dtfffc chuyen boa va l a ca c b I t cua quo t r l n h 5c cbl ( 1 7 , 1 9 ,
20» 2 1 , 4 9 , 3X, 34t 115» 122, 150)*
Do tfnh c b I t da dgng cua t l bao blnb sao va do k l c b
tbiioc cua than t l bao gu;? nbo, nen vigc ph&n l o p l
tryc
t l l p cac t l bao 5c c b l trong vo nao ca v l mgt cb5c n^ng va
blnb t h a i d i u rSc ph5c t j p ( 1 6 , 1 7 , iO, 36, 3 7 ) .


a lo

3

Mgt sS cac nha kboo hoc do c^ gang tits bl4u o i l 11 Sn b |
glUa clu t r u e cuo t l boo v5l que t r l o b 5c c h l bang
each
ngbiSn c5u cac c b i ulGu gian t l l p (-:^3» 1^1» 1^7)^ TnfSc
hit
i^tfol t a chu y dio c l u t r u e cua xlnop» f^ao l^Jt?^ Gray (1^1 )
v5l phtfo^g pbap c3 dj.nb va nhu^a to cb5c than klnh kbl nghlSn
e5u nhd k l r h bien vl di$n t u da pbgt hign vo ud ta hai
loal
sclnap s
- Lopl I l a l o ^ l CO khe :Elnap rync* aoni; xlnar day 113n
tyCf vung boa* d<5nt: 6 ca uarg trtiac vo raan^^ sau dIu rtfng, co
pban 5ng v5l a c e c y l c h o l l n .
- I*oal I I CO kba 3clnap b^p hc^n, vvn^^ boat dgnf^ hep
khong CO phan 5n^ v5l a c e t y l c b o l l n .
3ov do nSo 1^65 tcblznov (157) chi r o tMm l o p i I
coc t u i yinop hlnb t r o n , con l o a l I-^ co cac t u l srloap

hhv dyc»

va
co
blnb

Oac xlnop l o a l I va l o a l il phfm bo o^t each co
cbpn
Ipc tr§n tb§n vo cr6n coc d e n d r l t cuo nafrdn* l a c lv<35 ittidersen
Va cync sy da quon SQG thay ylnap thuyc l o ^ l I t i p trung chu
y$v tr@n dendrlG cuo not^n thop thuvc byppocoisp* I'ront: kbl do
xlnop l o a i i i pban b5 ci:u ylu 0 cr6n than cua t l boo nay,
6
t l e u nao xlnap l o a l i-£ t^p trung; tv^ti thSn ciia te bso pin:*klnj€
(73)• '^beo cac t a c e l a t r ^ n , srlnop l o a l I l a yinop b.rng ph*n,
con xluap l o a l XI l a :
hSitQ 1966 Lccles (Vu) cune thing n b i t v5l nb$c d^nb t r e o
Theo dtif;, g l a choy't; ylnop 5c cbl phSn b l t r e n than t l bgo iS
boan toon l o g i c , l^am gluo vuu; co xluap hittx^ pbia
(trSn
dendrlc) va nc/l tpo yunc:"(tf dopa dSu axon), c^c xlnsp 5c che
do c h l l n v} t r l tbugn l y l n h i t trong vl^c d l l u h^o qua t r l n b
pbat xun^^; cue n o ^ n .
Iron£; so cac c l bso soo, uu boo ry thuvn^- tgo m

xlnop


a 11 «

l o p l I I trSn tb3n t l uoo ubap ( l a . , 123, 130)« fixoo cua
tl
bao r p too nhonh rpnc tbeo bif5ng ckm ngjjnt:, ugo uhanb
doo
r l l quanb thfin t l bao thop» ^c d l l c noy qUan sot t b i y t r e n
t l bo^ Tp cuo »o nao, cleu noo vS hyppocorop* i>llu do
cho
phep cac nlia nghloti c5u gla tbuy^t rang s t l boo irp co trong
t i t CO coc vung vo nao ©51 va l o t l bao trun^: eloa 5c
che
(13, 19, 6 1 , 9i^. l a , 132, 14G).
Can c5 voo c l u t r u e vs d^c d i e s phan nbanh, o^m
1975
^ n t a g o t h ^ l ('>9» 13^) cbla coc cl bao r p tbanb bal l o ^ l »
- Te b^o rp nho, co cac axon cgin, ph§n b l chu
tron^ l5p i l cua vo nao»

ylu

- T I bso r p l 5 n , too nbanh rpn^, ph8n b l tronf- 1 ^
IIX-V cua vo noo,
hh%n d|nb nay cun{; phu hyp v5l y k i l n cuo ^orlQ-i'adilo
( 1 - i ) . hi^ixiSc c5u clu t r u e cua t l b^o r p tron^^ v ^ g v ^ d^Jng
vo nao nt:i/&l» t a c f:;ia tb*y nhun^: cl bSo any ph&n nbenh trong
g l d l ban nbtfnr. phlan tban^ d5n^::, vudnj^ goc v5l chiou t3nl bSl
trtmc crtttig taE» liiaunt: phllo noy xu;§n t 5 l5p i l xuSnc
l5p
V, CO chl§u dot lUuO-^QOU fids va day kboong iS)0 acD* iiscon
cuo cl boo r p c l i p xuc v5l cac t l boo tbop ahm ti*oof:
phlln«

6 noo khf, naci l^jf73 Glons(ll>) cuut^ quoa s o t t h I y coc bl boo
l o p i i V5 I05I Vi, tu'cync t y nhif cl boo r p l5n vo r p nho cua
Sentogouhal» Cbun*:^ P^^-^n nhonb cbu ylu vudnt^ coc v5l
tbuy
t r t ^ c trunj; ti^o*
oc»rdn
b(/5nc:
tblcft
tpoof

Goc dSn l l g u n£:hien c5u v l anh hiiD'ng 5c chl gitJa
cac
vung cao gloc i oeo cung phu bjp v^-l coc quan s o t
ve
pnun nbonli cua t l boo r p , 'Dual onb hi^'ng cua
kich
boo bpc, blgu qua 5c cbl the hlgn r o c' coc nc^rdn nhm
bmt)s: vuctttj £0c v 5 l ranh chtf tb^p ( 7 , ^)m


sit hyp phiKfnc pbap cbyp onb t y d^n^ bing phong xg ,
pbi/otig pbap Golgl Va klnh b i l e vl dlyn t i i , esc toe gla
fib|t
Ban (147, 1^6) da cho t b i y xlnap gll/a t l boo r p va t l
boo
tbap CO ban c h i t ^i^i^A (eaffinio-axlno-butyric--8xlt)» DSy
la
bSng chSn^; nuo bd sun^, cho gia thuylc t l boo r ? lo t l
boo
truo^: gloo 5c chl trong vo noo»

JNgoal t l ho^ r p z ^ , t l bao blnb bul cung^ d(;i^ col
la
t l bao trung gloc 5c chl tron^j, vo nao, TbSn cua Coc t l
bao
noy nlo trong I c ^ H , x l l • /ixon cuo t l boo toa nhanh tbanb
hlnb nhi/ c h i l e den cbuc ( 4 9 , 1 ^ , 150) vo too re coc
xlnap
l o o l I I t l l p xuc v&i dopn dIu ax«i cua t l baO tbap ( 1 M , 1 4 5 ,
146) •
Irong nliung nam gin dSy, cac toe g l a i*hgt Ban ( 145 ,
146, 146) c5n thay trong vo nao co bal lo©! t l boo blnb saO
khac cung t b i ^ c I09I t l bao trung gian 5c cbl« £0 l a cac t l
boo blnb âgng nbĐa va cl boo co bal bo d e n d r l t . v5l
pbtrcm^
pbap chvp anb b*ng phong x©, coc coc gla cuasf, thIy boa c h i t
trw5g gian cua coc xlnap nay l o GiiBA#
Ttf cac din l l § u t r e n , co t h i k i t lu0n rln^s trong
vo
nao CO nhllu c l bao trur-r; g i a n , Ve nguygn t:1c, coc t l bao
nay co kha nang gfiy 5c c h l coc t l boo thop vo coc ve
boo
khoc, Tuy tbu§c voo v}. t r l zQa cunc cuo coc phSn nhonb
cua
cbung trSr; t l boo tbap so 05c d j 5c chl co khoc nbau,
thI
blgn bBng sy suy glsm nhleu bay I t ctf&nf; dy lucng xung d^ng
htftyng tSo vo do do d l l u boo qua t r l n b tpo xunp cua t l
bao
t;bop«
2 . G/>r.4- c b i t trum- glnn 5c chl t r o n r vo n.r|Q .

Kbllu y k i l n cho ring c h i t trung gian 5c chl ct^
ban
t r o n t vo aoo l a gaomo-afflla^-'butyrlc-oxlt ( > , 5 , 5 8 , 7 1 , 82 ,
83, 93, 116).


» 13 «
Tbeo din l i $ u cua B0t s i t a c glo (55* 5^) ohnc 60 G^}^i
trotig noo ttPtmg d e l c a o , Ixboang 1,1-^j ool5/g» I^ung
phmmg
pbap pbon*.; x j donh ciu ( I r - GABA) Iversen va c^nc sif (112 ,
113) cbo b l e t , khoang 70 >v c h i t nay t|ip trun^^ a t?n
cung
tban k l n h , 11 >» trSn s y l khdUt: co myelin, 11 v^ t r i n dendrlu
va 6 ^ ti'Sn cl boo ijlia* 3y t i p trung OAB^ nhlSu t r o n g
t#n
cung t b i n klnb cb5nci t o no pbal glif vol t r o quan trpng t r o n g
din truyin xlnap t o l t^ay.
TrOQg nbufng t b l nghi^ra klcb t h l c b t r y c t l l p len
bi
m$t vo noo, doner th<^l v5l v l f c g h i difn t h I 5c cbl J* ncardn ,
tac gla da xac djnb duyc non^ d^ cua GABA t;anc Ifin g ^ 3 I a n
so v5l l5c cbtia kich thlcb» IJIU kb5 ion Ca**^ tronf; cd
than
kinb ohl khi klcb t h l c b vo nao nSnt d^J QiiBA khong tling ISn «
Dllu noy cb5ng t o Gn"*^ lo ylu cl kbdng cbl t h l l u duyc
dl
g l a l phone CABA, tiXcPng t y nhu t i n h c h i t cua coc
fooa
chat

trung glon khgc da dtipyc cdc^ nb§n (112, 1 1 3 ) . ^o cvng 1 P B^t
bing chSfif^ ou^ d l kbtrong
xlnap 5c chl cua vo.nno,
I^htfoG t b l nfchifiD dtPo coc chic vao t l boo bang
phtfcfng
pbap vl dl^n di (mlcz-oelectrophoaras a p p l i c a t i o n ) cua Knlevlc
(116, 117, IIB) va Alexandrov ( 3 , 4 , 5) da thu dtfyc k i t qua
CO y n«:bio don^ k l , c b l r o anb hi:f(5^g cua Gi^A l^n bo<^t d^og
t a o xunt cuo t l boo t h i n klnb vung cac g i o c . Blnb
tbiiWig
nc?r6n phot xun^j tiKme d l l 6n dj^no, v5l xiln s i kboang 5 xung
trong BK?t g l S y . Dwl anb hiWng cua dong dl§fl d l GAB^,
co
earns d$ 20 h^, ^ 70 >^ t^ng s3 oc?rdn ghi duyc, t i n sS
xung
glaffl xuong con 0,6 xung trong ns^c g l a y , 6 ogt o l no^on kbnc,
xung d^ng libdng xulu blgo, Cgc no^rcn noy do b | 5c cbl
bo5n
t o o n , TrontT dieu klgn trOn, v5i difn eye gbl dl§n bSn trtmg
t l boo, coc coc gia da ghi d; ye dl§n t b l tSnc pbSn c;,^
( h y p e r p o l a r l z a t i o n ) * I^bti vgy UA£i do gSy tru ph^n eye
aong
no»rdn a vo noo cut^e t y nhi^ dl$n t b l ijang pb3n eye
troni::
xlnap glu-a t l boo tfo cbl iienshaw va t l baa vpn dyng
tuy
sln^!;*

s 14 9

Myt trong nburng pbusmg pbap xac d^nh co blfu quo
val
t r o trung gian boa bpc cuo GAB/* lo dung cac c h i t d l l
kbang
( a n t a g o n i s t ) cua GAB^ t a c dgng len nc?ron ( 7 9 , 6 ^ - ^ , 9 4 , 105,
106, 107, 135, 136)* Gac cdng t r l n b ngbldn cms cus C u r t i s vo
c^ng sy (04-66) da cho thay 00*51 tac d^fnp, cuo alcalol^^-blcu culln ( a n t a g o n i s t cua G A B ^ ) , bl^u qua 5c onl cua OABA h% s i t
hoan toan* kbdog ahatii^ t b l , nhilu kbl xung ahn cua neuron con
t ^ g bG^c blnb thi^4^« l a c gla cbo s&c^, trong d l l u klfn
ty
nbiSn v5l m^iz m5c dy nao d o , ncfrdn da cf vr&Cf: t h a i b}. 5c c b l ,
v l t b l cac d^mg cua b l c u c u l l n kbdng nbting pbong toa bl^u qua
cua GiB^ so con g l a l pbong nctrdn kbol'*5c cbl truPcfn^ l y c ^ Sou
do S l l l l t o (135, 136) vo HlckS (107) kbl ag:hl§n c5u sy b l l n
d l l trtf^n^^ thy coo cua ncfrdn vung cao g i a c cunc nb§n
tbIy
hlfu qua cua GAB^ b} mit boon toon kbl cung m(^t luc cbo CABA
va b l c u c u l l n t a c dyng I t o acni'dntf
Kit qua t b l nt"bi§ffi cua Stcwat va Cf^ng sy (149) cbo t h | y
ngoal blcuculln c ^ co myt lOal o l c a l o l t khac l o p l c r o t o x l a
cung lo c h i t d l l khon^-: cua GA3A» i^bl dtfo voo t l bao,
cfalt
nay cung l a s giom bi^u qua 5c cbl cua G^BA tdcmg ty nbii
b l c u c u l l n , Cac cdng t r l n b ngbign c5u gin day n h i t cua Alexandr
va c^ng sy ( 9 , IG, 11) cung cnung oinh t a c dpn^: cus p i c r o t o x i n
ttfcfng t y nha b l c u c u l l n trong quo t r l n b b l l n d l l trii&ne t h y cam
Va t l n b c h i t d e t e c t o r cua ncTcc vung cac g i a c & tseo,

Kbl x^^> x l c cac c b I t l a s glao blfu qua 5c c b l cua
GijBi
Sexkov (55) dya trSn col l l # u thyc ngblfm cua nhilu uoc
gla
da duPa »» y k i l n v l vol t r o cua p e n l c l l l n , Tbeo dn^, p e n l c l l l n
cung l a c h i t d l l khanu cua GABA, CO ^ac d]fn£: Im glam 5c
cb
cua GAB A trong vo nao» i;o do p e n l c l l l n da d*/yc dung d l
too
nfaimg Old blnb dpog klnb thyc ngbl^o,
GJ Qhi toe dyne d l l kbang cua blcuiitlin vo plcrotcwrln
d l i v ^ CAi^ khac nbou« Blcuculln co clu tao tiKfug t y nbu phen
ttf GAB/J va cung aha GABA, no k i t hyp v5l **phan nh§c bllt** cua
vung thy c a s trSn ©ong neuron* v l vyy ma sy k i t hyp noy c?ang


^ 15 «
t l n b c h i t "canb t r o n h " . Con p i c r o t o x i n c b i k i t hyp v5l •*pb5n
d l l u boa", nan t a c Cms kbdng tbeo ccr cbl canb tranb (149) •
Soog du tbeo ctf cbl nao, cac c b I t d l l kbonc cung locn
glao
bl^u qua 5c cbl cua GAi>A d l l v5l oOvQn nbif r ^ a u , Dllu
do
kbing djnb thStn gia tbt^ylt cbo r i n g GABA 1Q c h i t t r u n g gian
5c chl trong vo nao,
Tron^i, Coc t|in cung thin klnb i vo nao GABA di/yc tyo r a
bIng con dLro^.g ^ c o c b o x l n boa a x l t g l u t a s l c v&l sy tbaQ g l s
cua aen g l u t a o e t decacboxylaza vo coenayo Bg (113)•
I^ng
tbtfl CABA cung duyc *o^ tbu IQX v5i o i l y e It/n (Ko » d^S

(K» m 2,2 , 10^5 M) (112, 1 2 4 ) .
Bhm^ k i t qua ngblSn c5u v^ slnb l y ncfron, v l t l
bao
hpc, v l di^c ly hpc va bIng klnii bien vl dlgn ttf da Isisang
d].nb GABA CO du vioM chuan myt boa c h i t trun4' glon ifc
cbl
trong vo nao (55)» GABA diryc too r a tron^: cac xlnap l o a l I I
(xlnap 5c c h l ) gllio t l boo sao vo c l boo t h a p i gSy tfu pbSn
eye san^: t l bao thop va do do 5c chl kba nSng boat d^ng t y o
xung cuo cl bao nay, Nbtf vgy ca ebl 5c cbl cuo te bao triuxg
gian d l l v5l t l bao tbap l o nhh chat t r u n c glon hoc bpc GABA,
bay nol each khac, 5e chl co ban c b I t G^BA.
vl v§y x5 dpnc G^B^ va d|lc b i $ t lo cac a n t a g o n i s t
no trooB v l | c nghlSn ctfu quo t r l n b , 5 c cbl lo boan toon
ly-

cua
byp

3» c l u L-ruc mddun vo c l u t r u e tan ouSn ( c n s s a b l e ^

Sy b l l u b l l t v l ntsuySn t i c boat dyng cua tiicg nc^on du
c5 pbODt': pbu Vo do dour, din dev vIn kbdng cho pbep nbgn th5c
dtfyc dly du cac ntuyOn l y bojjt dgnc cbune* cua ca b^ noo,Cblnh
v l v^y ma vlgc ncblon c5u Qg;uyen t a c td cb5o coc agnr. ncfrdn
duyc c o l nbi7 cs^Jt bL?5c quan trpng t r o n r vlgc kbass pbo
nbifnr
b l m§t cua b^ thing t h i n k l n h .

ic^
Quan nl|c9 vl sy tBn t a i troue vo noo cac t l boo trung
glon 5c cbl CO onb hiftJiag r a t l5n din kbol nlyni bl#n dpi v l
sy t<5 cb5c - ebue n^ng vo noo o 5 i , Tbeo kbal nlgo noy
tbl
cac nc^dn tgp hyp tbonh coc clu tr&i oddun vo cau t r u e tpp
quan*
ã ) f^^*M^ ô coc no^dn tĐp hyp v5l nhgu tbanb cac cgt
hlnb t r y theo chiSu thing d5n^ t p l I s coc raddun, fcfidua glo
nbimg t l bao t h i n klnb cung thyc hlgn myt cb5c n^ng
tbdng
t i n nio do ( 1 6 , 1 9 , ^ 1 , 2 7 , aB, 4 3 , 54, 1 3 I , 1 6 1 ) .
Rgu'S^l dSu t l € n d l cgp d i n c l u t r u e mddun trone vo nao
l a i^ountcastle (^3) va xi»«ell (131)* Kbl k l c b t h l c b l$fi da
0 k h i , cac t a c gia n b ^ t b l y t r § n vune vo nao cas g i a c
co
cac cyt ncfrdn bufng p h i c , c^nb do co cac cyt iKd^on 5c c b l •
l^bun^i luc do cac |»ac gla trSn cbua g l a l t b l c h dtryc
cb5c
a&cs cuo coc cyt ncrron nay* ^ u nh^ co sy phot bifn r a coc
ncfrdn trung glon 5c chc trr^ig vo nao, quon nl^is v l c l u t r u e
cyt din din dtfyc song t o *
»b5n^" din l l ^ u ngbien ctfu vh dlgn sinh l y da c b i r o ,
kbl dlgn eye d l quo cac l^*p cuo vo nao vo ludn ludn
vudng
goc v5l b l Q§t vo nao, tbt/acg ghi doyc coc wjron g l l n g nhau
v l o§t cb5c nang* Cac c^t ncfron co chSc nang c l i n g nbau gbl
duyc c? cac vung cm g i a c ( 4 3 ) , vung th% giac ( 1 0 8 ) ,
vung
tblnh g l l e ( 1 4 , 15) vo vung vgc d^n^^:, ( 7 4 , 7 5 ) . ^^bu v0y c l u

true c^t l a d^c dllts cbung cua vo noo#
Tbuylt clu t r u e cyt ngoy nay duyc g p l l a t b u y l t tsddun,
Tbeo ^ o u n t c a s t l e (43) dan v} ccr ban cua oddun lo v l oddun
v5l di7^^ klnb gin 30 tsco, Cac ncfron trong ©yt vl reddun co
pban 5nt cbpn Ipc v5i myt thorn s i n h I t dj-nb cua k l c b
thlcb
btf&g rSi3 va lo dc^. vJ. stf cing n b I t cua quo t r l n b xu? l y
tbdng t i n * Khoang vol trSm v l sddun t^p b ^ tbonb mddun lt?n,
c l c l u C90 ph5c tgp bc/n, l^ddun l5n co dpng p h l l o , d a y kbooog


TT;

i V-W^#5J

« 17

30 - 2G0 Dcm, doi ^^00 - 1000 ocm, xuy§oc5 l5p l i t 5 l l ^ V
cuo vo noo* Trong oddun l5n d l l n ro coc quo t r l n b xu" l y
tbdn^r t i n pb5c top licfn v5i nhleu thoo s i khnc nhau cuo k l c b
thlcb* i>o l o dirn v|. xu ly vo t r u ^ l n dpc t h e r e t i n , t l l p xuc
v5i hong l o y t dtfo^i^:; %?ao vn ay^ f^c dwcni ro ('^3)*
t>btf v^y, vl n^^uyen t i c cau t a ^ , eac mddun l5n co cs?
s^ blnb t h a i tacmr; del r o , nbtfn cac v l ludtiun - dc?n v l cb5c
nanf' co* ban - c o kich cbuf5c 3G oco khdnf phu hyp v5l
sy
phan nbanh cua ogc cyi t h i n klnfi d l til t h a l ^ i u s t 5 l vo noo,
Tbeo t a l ll&u dQ dupyc c nj: nhgn (5G), ®0t cyi than
kinb
thalamus - vo noo tSa nhonb tronr vung c b l l u 6 vo noo trong

pbam v l CO diio^T klnli 30U-iiOO ocs vo t l l p ::&; xlnop v ^ gan
5000 ncfron* iJbimg v l ©ddun v5l dMn^ klnb gen 30 oco c b l co
t b l boo i^hm 110-260 acfron (^3)* 5y fflSu tbuln tiny chl co cbl
g l a l chlcti Khi phat hlfn r a coc t l boo 5c cbl trong jsddun ,
Cac t l boo 5c ctil difyc bav^s P^ldsng
biltfng t§t3 bo^Jc ba'l cgic xun^ dyn^ tbeo coc sg'l qu^t nfji/yc.Khl
coc cl bga nay htrng phan chWH^, se ifc che coc ce boo
xung
quanb vo too ra coc vunf *^ c h l quonh c^c mtcoc bt/ne pban •
Do k i t qua cuo qua t r l n b 5c c h l , sy hung pbin duyc kbu
tru
trong phafts v l cuo dc?n vJ ch5c nan^>
^^bt^ vvy vl ffiddun l o dcm vJ. cti5c non^^ diPyc hlnb
thonii
diTol Goc dyne cuo coc t i n hi^u bir5n£. vam^ Hlnb doni; vn k i c h
cbtf5c cua no phy thuyc r a t nbiau voo co' s^ blnb thol bsn dau^
libim^:; cd^^ t r l n h ngblcn c5u ve blnb t h a i hpc cuo i-arlnf o d i l l a (122) gop phan c l o l tnlch vol t r o cuo 5c chl
trong
vo noD del v5l t<^ ch5c - ch5c nan^: cuo coc ©cjdun. Joe
gla
da figrblen c5u ty o i sy pbjn nbonb cuo t l boo r p trong
cac
l5p cuo vunc vpn dyng. Axon cuo l o ^ i t e bao noy too
nbonb
t^ong phao v l nhif^: p h l l o tban^, d5nf: v5l ciileudny khoong
200 f3ca) dal 1000 - SOuO occ;, vudnfv; .:; oc v5l b l (u§i vo nao ,
xuyen t u l5p Ul din l5p V vS tl^l^^WssMM^'^'o
co coc
t l boo thop nIo t r o n c phleri. ^''bu wy^by^^hl^^,^^* t;o^ r p - t l

sif* *-K1« AZ ^^^ — •**^'>c4f eye no^oti c^:dv*^<9>-?blln^Ai^ae d5rig.


a IC Klcb t b i / ^ vo hlnb 6ont, cuo cnc cyt noy do d^c diem ptxbv^ nbonb
vo chieu doi cuo oxon co boo r p qvye\^ d^nh,
Tbeo Sentagothal (50) trong vo 000 coc raddun co
blnb
done vn kich thtr5c kbnc nhau l o uuy thuyc voo clu tpo cuo t l
boo rp* T I boo r p nho a l5p Xi tao cbonb cac cyt co
diwng
klnh 50 -• 100 oco, phu byp v ' l klcb thir5c plmo nh^^nh cuo axon.
T I boo rp l5n o» l5p I I I tao thirinh coc oddun co dgng pblln,doy
kboong 23-3O oco, dal £:ln 5G0 oco vo sau t 5 l5p U d i n 1 ^ V,
Tuy cbo din nay vSn cbua co oyt quon d l l a thing;
nbIt
vl blnb doat:, klcb thu?t'e cua oddun, vl ctf chl coc dyng 5c c b l
l | n nbau trong oddun, vo g l i ^ coc oddun, nbi/nr coc cbo ngbldn
c5u dIu edng nb^n r i n g ylu ul khdnf: t b l t h l f u dixyc tron£ clu
true mddun lo coc uti boo trun?^ fclon 5c c h l , Cac t l b^o
noy
dong val t r o quoo urpni; tron^^ vl0c v6 cb5c ciou hoo boat dyng
cua oedUQ v5i coc o5c dy pii5c typ khsc n h a u , ( 1 6 , 18, 4 9 , 5G ,
5 2 , 5^).
b) gap Q^h C^aagp^lt)
B^n^^ phuc?n^ phop s5 dyne nbllu dlfn eye (c;-5 vl
dlpn
e y e ) , ghi hOQv tlnb d l | n cuo oyt nhon ncfron, k i t hgp
v5i
ph\f0tk£ pbgp xoc dj.nb t l n b c h i t va ffi5c dy ti^tii: uac GI^O cbung,
oyt s i toe glo (<-tj «. >2, 34) da cbo tboy m^i anb htKm^. cua

coc t i n lil^u htf&c^ t § o , troofy cac t^ chSc uhln klnh, oa tr*/5c
b l r lo tronr, vo bon c5u djil noo, hlnb thooh c l c t&p quin nc?ron
v5l hlnb done' vo klcb thi/'c khac nhou,
Tbap Kogon vo c^tc s^ ( ^ , 3Gt 34) t h l c#p quin
noron
l o oyt ohm coc no?ron cure i trong tr^nf: t h o l biinf: phan, d u ^
bao quoch bSn^ vonb doi coc u l boj bj. 5c c h l , Vg c^v. uruc
vonb d.ol QDC V • r.o3 b} t'c c h c , Vo clu t r u e coc typ quan ncfron
CO nbl/ng d$c ctiea co* bat^ sau s
- tidl c$p quia boo gom coc ncfron cun^; I05I 0'
trgng
t b s l bimg pblo kinoc nhou vn nao t r e n cun^r myc a5c td ch5c cuo
cun*? nbnn x a . th^ia chl lo cung cron ffiyt l5p t l boo, ^ihi;
vgy


» 19 a
(J oSl l5p i:l bjo vo nao cung oyt luc co chc hlnb tbanb nbleu
t^p quISn no?ron* 2>o do t^p qu5n co cSu t r u e khccpr c:lon
theo
c h i l u RcnnCf kboc v5l ofidun co c5u t r u e tbeo c h l l u dpc
suit
be day cuo vo ooo,
- Coc t§p quin CO klcb tbtr5c khac nhau, pby tauyc
vao
bon chlu va tbdn^ so cuo t i n hlgu h;?^5nf^ t a o , voo n i c
thane
t l l n boo cuo dyn^r v^t* C dpni: vgc bye th^p (bo s o t , li/5nf: tb§)
kich tbif5c coc tgp qoSn r a t l5n (40U - 500 ocm), coo c? d^ng
vgt CO VU, dt^^^nr klnh cuo t^p QuBn c h i khoonc. 15^ - *i'X meo.

- 1§p quSn c h i blnb thinnh kbl co klcb t h l c b h i ^ g too ,
vl VQy no khI09I
klcb thich bu5n^ tfio t^o nen coc t^p qu5n co hlnh done
va
kich tbu5c ttfcfng &£;• ho toe dyn^ cuo mdi uri^&n£: 18n c c
thl
sdnr ludn ludn blSn dyn^ sen clu t r u e kbdn; {;lan vo chc?l gian
cuo t§v quon cun?? ludn ludn tbgy d e l , i^ol coch kboc, coc t§p
quSn di/yc hlnh t:honh cheo ngu^/Sn tac xoc su*t» thln^r k g ,
ngbia l a sy thom gla cuo coc nc/ron voo c l u t r u e c^p quin d l l n
ro theo nf^^uy^n t a c xac sUwt, con higu quo boot dynr cuo
t^p
qu»in I s kit; quo tblot: k§ hogt dyof: cuo cac nc?rdn clu
thonh
t | p quIn d o ,
- Vnnh doi coc ue ooo bj. 5c che boo quonh cyp quBn cung
ludn ludn rhoy ddl du^5l onh htfonr cuo kich t h l c b hif&vj^ cao •
Gblnh CQC ac^dn bj. 5c chl nay quy dj.oh cau true khdne
glon
cua t^p quIn,
- till quon b | ttfcfnt,: toe gltfo coc titntm tronK- cuni
oyt
t§p qu5n boo giS cun£; 1? quon hf dSng pha* Hf s l tif^Vi^
quan
difcfn^ v5l s i s t r j tv^/^t doi kho coo, lig-tfjc I p i hf so
ttfcfng
quan gl5n ndton n^o crong trung tuo bt/ag phan vo'i n o ^ n
nao
ngoal vanb doi tic cue boo gic^ cune; fflonf! dIu 3n vo do do

oil
quan h0 ciKmr toe tron£: tru^f- hyp noy lo 0/14^*^ P^'^^*
v | ci/ c b l hlnh thonh coc c^p quan ncTrdn cho din noy v8n
con CO nbuj]^ y k i l n kboc nhou, Tn,y nbien cusif; co myt 00 quon


« 20 a

d l l o Chung nhtf snu s
- Coc ncfrdn tboo e l o clu uhonb ojpt tjlp quIn dIu phol co
nhi/ng yeu uc hlnb th.nl chimg baodau, cunt" num tr^n nht?n£: phSn
nbgnh cu^n s^i tnan klnb ba5ng tao x u l t phac tu ayt so thy cao
chl n^^opi v i boy tJlr c^yt nhom cl boo naty i tklnb
t b i p bfSti cheo cung phon xg,
- ^c? nh hlnh uuoi trSn c h i l o t l e n d l can t h l e c ban ^ l u ,
eon vol t r o cd cb5c ndn coc t^p quIn I p l tnuvc vc coc
dong
t i n blgu btf^nt; t3đ, Coc k i t qun ne:biĐn eSt'^ cua nliieu toe glo
da cbSnr ralnb r i n g cf dlBu klyn n^n, a l l quon hg tutfn^toe
gli/o coc nc?ron r i t y l u vo kbdng mong ulnh dj.nh ha&n^ r o r f c ,
- Hlnb ^vin\^ vn klcb thi/Sc cua tgp quSn con ph-^
cbuyc
voo quo t r l n h 5c c b l xoy ro cJ vo noo, '^o t r l n h noy chu
ylu
do toe d^ng cu? coc t* bao 5c ch*^ t r u n r cinn tgo non (3i:)*
!Dheo quon dicD cuo coc cac gla t r i n day t b l odl
tpp
quSn ncfron dt/y'c coi nhif opt do"n vJ. ch52 nano at/ dSng nb^tj
,

^ b i l u t^p quIn nam trSn coc c?5c dd td ch5c kboc nhou cuo cung
pban xp tpo thnnh o^c bloc ch5c nanj^, cun^ thoo r l a boan
thonh o j t nblyo vy tinto djnh noo do ( ^ , 3a, 5^), 0 dSy
ta
they cuori diem oddun vr quon dllmtgp quan g | p nbou, ?roDg ©yt
chu^g oyc nao do co t h l bleu oddun nbif ©yt bloc ch5c nan^v,
I^ con CO nJitfn^: dieu kboc b i y t ve c h i t i l t , coc toe g l a
cue ca lisi cuan diliii ^^u d i den k i t lu§n t h l n t n h l t l o
coc
no^ron kh6a^ bogt dyn£.: oyt coch r iong l e i d(/5l cac dytf
cua
coc cin hlyu bu-^ng tSo ebun^' ci?ye xh cb^c I j i uhonb totie. nhoo
tbeo b l ngonf; vo bl doy cuo vo noo, tco uhoah coc dcfn vJ cbSc
naQti: stf dSnr:* Cblnb co c h l noy cho phep g l o l thlcb du^jic pbSn
nao nbtfng dgc t l n b co? ban trong bjpt d^n$ t l c h hyp cuo by noo,
Thoo glo hlnb uhonb coc d^c t i n h noy khong ^hl t h i l u dtryc vol
t r o cuo coc quo t r l n h t;c c h l tronf vo noo.


u y - •piii; ^ ^ u iii^ CO ciiu TAG LU.W bc G:U: MH miih
CLL ItOI.Ui. VO ULO

throng sinh l y a;/ron, coc thyc n^bi§m c l l i e n quan
v l f c clo b l l u toe dyr^ 5c chl ben trong vo nao 6a vo
diiyc t i l e bonb bScg nbl§u pbtfcrng pbap khac nhou*

sSn
dsioe

V5l pbuc^ng pbap fc:bl dlgn uhl cgooi vfi t o o , bong

loat
cac cdnc crlnb ( 2 5 , 33» ^^2, 5 6 , ^Z^ 6 3 , *?9, 127) dS cfal r o t
dif5l anh btfd^ng cuo b i t c5 oyu l o ^ l klcb chlch buSng t&n noo ,
dnp 5D£* CUO nitron vo cao tbt^cng chlo thonh nhilu pha* i-' da e l
nt?ron, pbo dIu bt*Qg phin (bi/n^?; phin ea d p ) v c l tho^l
glon
ngin (kbooUG IO--3O ooec) tSn s i xunc tang l § n . Sou do l o pha
5c c b l keo d a l (350 - 2P0 o o e c ) , ncn?on QIOQ bc^c n^^iing
phot
xung, 21ep sou gjloi dogu 5c chl & m^t sc nc?roc. co bl^o ttryng
t3n£: phot xung: trt^5c kbl tr:5 1^1 blnb thiidn^ (pbo bilnt;
pbin
sou 5c chl hoy lo bun^ phin tbS d p ) * 0 ©yt ec I t ncfron,
5c
cbl phat t x l e n n^oy sou kbl klcb t h l c b btfSnt ^^^ ^k/ ro ( 5c
cbl 6CP c a p ) , 6ou do ay phot xun^ cuo ncfron oo'i tang ISn*
Sht? v§y tron^^ d^p 5ng
g l o i dojn htrui^ ph'n vc g i o l
soc tbIy ^ coc t l boo tbu^c
klcb t b l c h d§c hi§u xoy ro*
5^^, 53) do k i t l u | n t

coa t * boo tb§n klnh co xen
ki
doan 5c c b l , 3$c dlecs nay
quon
t i t ca coc vun{ vo nio kbl
co
Can c5 voo do Serkov ( 5 1 , 5 2 , 5 3 ,

1 - tic c b l l o thonh pbSn kbdn^^ t b l t h l l u dtfyc t r o n ^ dap
5n^5 cua t l bao than kinb dci vtfi cnc klcb cblch btf5ng t g o ,

a- "Otfp 5nc, cun ncrron ©one d^c t l n b pbo, t i ^ l o t r S n o^t
ncfron ca bal quo trlixh htpn^ phin va 5c c b l dIu x u l t hiyn vffi
il^loh t y tbo'l gian khoc nbou,
tJc c h l x u l t bign oyt trong trtf&ng hyp t r e n gpl l a
cbl btf5ne cao, mQc trong nhung dang 5c cho cuo vo nao,

5c

M^t trong nbung phiK^ag pbap ngblgn e5u co blyu qua quo
t r l n b pbat t r l l n vo keo d e l cua 5c cbl btf^ng tSo l a
phuwie


m 22 ^
pbap klcb thici: d d l , Klcb t h l c b t b 5 nblu gpl l o k l c b t h l c b
d l l u k h l l n i klcb tblch tbu hoi Epl l o kich t h l c b b 5 n o Hal
klcb t b l c h coch cb^v nhtfnt: kboon^ thitfl gian nbot dj.nb gpl
l a tbSl glao gl*^ bol ^^ch chlch* t^gir&l dau t l e n dung pbtfcfng
phop noy lo Eruca vo c^nt; sy (bO) v5l snyc dlcb t l o
bllu
trtf^n^ thy coo cuo ncfrtKi vm^^ cam g i a c (y o a o , l^lch
tblcb
htfng duyc dyt voo truct cOo cua trtfo'nfr chp cotr, klcb t h l c b
diau k b l l n dtfyc dyt t p i coc diC'm khoc nhau tronF crtf^)g thy
Coo* Inb btfotsg 5c c h l do klcb tblch d l l u khiSn gSy ro da l a o
eiim sw' dop tfn^^ cua ncfron d l l v5l klcb chlch Lu^> lya
voo

k i t quo noy t a c gia da pbSn l o p i di/jflc tru^nc thy cao cua
ncfrdn, Sou do n3© 1979 i-askln vo c^n^ sy (119) op dyng pbtfc^g
pbap uren d l n ^ l S n ci/u anli b u t ^ cua 5c chl d e l v5l dap tfng
•^51^^" cua ncfron kbl klcb t h t c b d l l u k b l l n nam 0 coc vJ. t r l
kbac nhau trong trtf^nc thy casi vo tbSi gion gltfa boi h
kich
thich cwu^ khac nhau* C^a phan t l c h coc toe ^us tbIy
anb
htfc^ 5c chl couf; oynb kbl kich cblch dieu k b l l n con?;
nao
gin trung tao trtfStig thy cao va bigu qua 5o c h l l5n nbIt kbl
t h ^ l glon Gliiia b*«l klcb t b l c h khdnj^ qua 20 moac*
Tuy vSy t r a a : cnc t b l ngbl|o trSn cbtfo co odt
olnh t r y c t l l p noo kb^nf: d}.nh 5c chl htftfnfv turn co ban
sau xlnr^p,
i>l g l a l quyli; van d l noy, coc nho nghldn c5u da
9§t trong hai pbtfcfng phop sou t phucfnir pbap g b l dlgn
trong vi boo vo pbtftyr^ pbap dtfa coe c h i t d e l khong cua
trung (3^1on 5c c b l voo trong t l boo bang v l dign d i ,

ch&xg
chit
onus
thl
chit

Kit quo gbl dlgn t h l trone ^e boo da c h i r o , dign t b l
5c c b l sau xlnap *• I n h i b i t o r y Postsynaptic i ' o u e n t l a l ( X P ^ )
dc^c too 3?o t r e n ncfron kbl co kich tblch t r y c t i l p ISn
bl

D;^t vo noo ( 5 5 ) , len cac s y l tbSn klnh vholomus - vd noo(l02)
va do b i t ky oyt kich t h l c b htf5nfi tao noo ( 5 3 , 5^*, 5 5 , 56 ,
6 2 , 6 3 , €>4, 6 5 , 1 0 2 ) .


« 23

MSo 1977 iiellweg (102) kbl klcb t b l c b d§y tbSn
klnh
I n t r o o r b l t o l l s (trorv^ do co nbonb di ttf thy can; t h l v i b r i s s )
do quon sot thIy bo 109! phon 5ng cue ncrron VUJH' COO
gioc
aooQ t

L09I 1 s Slyn t b l bung phIn sou xlnop - xung—Sxcltatoxy
Posta^^naptlc p o t e n t i a l - spike (Ei^SP-splke),
L09I 2 % Blgn t h l 5c chl sau xlnop ilx'^'^
ijopl 3 t d o CO dlgn cbl hung pbIn sou xlnop •* xung —
dlyn t h l 5c cbl sou xlnop
(iiPSP ~ spike - IPtJP)
Lopl 3 thtfmig ggp bcfn co*
Trong nbtfng tints 1979-19^^ Pbaooskl ( 6 2 , 6 3 , 6 ^ , 65) g b l
dlyn t h l trong t l boo vung cats gioc khi klcb t b l c h d^c
bl^u
voo coc thy coo bne v i b r i s s cunc QUon sot dtfyc pbon 5ng ttfcfng
t y , % ^ o l 3^0 Coc glo con ghi dtfyc l o g l EPS** ^ IPSP (kbdng; co
xung),
Coc n^hlGO c5u t r e n ncyron vtaif:; tblnh ^ l a c vo vunc
g l a c ( 5 3 , 3 ^ , 56) oun^: deu gbl d i ^ coc l o g l phon 5ng
tren,

tb}.
nol

Kbu v^y trons chonh phIn dlgn t h l tron;: te boo cua pbIn
l5n neuron dIu co dlgn t h l 5c chl sou xlnop*
So sanli tbtfl (^lon t i l o tone cuo ^ S P va 1P£5P 0
ncfron
kbl CO kich tblch htf5nc t S o , nga^i ta nh§n t b I y IFt^
xult
hl§n cb^is bcfn i-PSi' -vhoong 0,fc - 1,2 osec ( 5 3 , 5 5 , 6 5 , IO2
,
1 5 ^ ) . s l l l g u noy thu dtfyc khonc nhling 6 t l boo vung coo gioc
kbl klcb t h i c h ngogl v l , 00 con ^ vuni, t h t gloC kbl
kich
t b l c b doi t h t vo kbdnc tboy ddl kbl k l c b t h l c b t l o ubj (55 )
Dllu do cb5c£ t o r^n^; sy n^tfn^- t r g tron^r d2n t r u y i n dl^n t b l
5c chl sou xlnop xoy r a cggy chlnh tronc vo coo ch5
kbdng
pbal trSn citrSnrr dSn truyen u^:ool vo* libtf v ^ ' l e n tron^ vd noo


a 2 4 ar

pbol CO Q^c nban t l noo do IOD tfc chl quo t r l n h din t r u y i n
btf5ng tfio* Rb§n t l noy cd t h l lo t l boo trun^:; gion do nb^c
t 5 l 5 vrStim IPSP xuau hlgn trong trtfStog byj> nay lo li^i^P bal
xlnop, t r o n r do xlnap 5c chl gltfa t l bao t r u n c iilon vo
tl
bao tbap da co toe dynt ban c h l sy Ion t r u y ! n btfng pbIn

Qgay trong t e boo thap*
IrSn coc vung vd nao cd l | p , boon toon l o p l trtf
opl
dtfSn^ l l § n bf btf5n0 uSo* Si"^ xulc bl^n kbl k l c b t h l c b t r y c
t l l p l e n Boo (59, 6 0 ) , Dl5u do kbIng dj-nb thoo EPSSP
do
cblnb coc t e boo 5c c b l trung glon t r o n c vd noo gay r o *
Co i;al l l g u cho ran^:: iPbP tarong vo noo chu ylu do
5c
cbl qu^t ngtfyc tpo ro(130)* Song uheo Searkov ( 5 5 , 56)t t h i
gltfo 5c c b l btf5n=^ tStn va 5c chl qu§t ngtf^'c co nbimc^
dllo
kbnc nbou. t5c chl htf^m taa yu:1t hlfn ngoy t r o n r l*n
klcb
cblch dau* IFaP cuo ^1 boo vun^^ thlnti g l a c co tiial
gian
t l l o vam: thoy ddl tuy thuyc voo vJ t r l cua klcb t h l c b
btf^nf- t6o» Tb$l gian t l e o ton^: a an nbIt kbl klcb thlcb vao
tholaous ( 0 , ? - 1,3 o s e c ) , dal bcfn kbl klcb t b l c h voo
hoi
nhon sou cya cu noo sinb t u ( I 9 6 - 2,5 osec) va doo
dyng
l5n khi kich t:ilch ban^; §m tbog (7 - 23 msec), IPBP
glao
bin xuinc khi klcb t b l c h btfSng too l o nbimij lopc klcb t b l c b
llSn t y c ,
%tfyc I p l IPiiP ^ t l boo vunc vd noo v$n dyng c b l xulc
blyn kbl klcb t h l c b n^tfyc c b l l u voo bo chop bloe nbtfnci; l o a t
klcb t h l c b llSn t p c , i#oat klcb chlch co tSn s i cof^i^ l5n
,

t b l I?SP cang tan£; vo coni' keo del* Tbi^l glen cleo
tong
cua li'SP tron^ 5c cbl q u | t nguyc doo dgng kbodng 2 , 6 - 6 , 2 ©see
t r u n c blnb 3 , 9 osec (55)*
Tv^ sy khoc b l y t t : d n , t i e gla d l din k i t lugn rang 5c
c b l btf5ng tSo kbdng pby thuyc voo 5c cbl qu§t ngtfyc,
Cac
t l boo trung ^lan thoo g l a voo 5c c b l q u | t ngijpyc kbdn^^ a^a
trong khSu dBn t r u y l n btf5n(r tlio*


« 25 *
lihtf v4y tronc vd olo tSn t p l bol bg t h i n g t l bSo^trung
glon 5c c h l , ddo boo hoi qu5 t r l n h 5c chl kbic nbou l o
ac
cbl htf5npj too vo 5c chl qu^t ngtfyc,
Tac dyuf 5c c I l3n nhou cda t l bSo vd noo con
dtxyc
ngblen c5u quo sy x u l t bL^n ^ ^ vS l^^ ^ t l bio kbl klcb
tblch vSo coc d i l o r S t nbd ben trorig noo ban^: vl dl§n eye .
Pbtftfnc phop noy L^?1 t i t l 5 pb.fofnc pbSp klcb thich
trong

noo, Asanuoa vo c^n^- sy {7!}) kbl dun^^ xunj- dlgn co ctfmig d^
4 OCA Vo keo c^i 0,2 o s e c , klcb t b l c b vno coc dieo 0
Icfp
I I , I I I , iv vung vd noo vyn dyng a oeo do gbl di^yc coc dl^n
t b l oou xlnop cuo coc t l boo quanb diem k l c b t b l c h ,
Tr§n
kboon^: coch 50> oco tbeo htfc'ng nao ngaog vo xucn^:; sSu phio

dtf5l dlgn t b l gbl dtfyc tbuye l o y l n>dt x l n a p , con t r e n khoang
each 100 oco dl^n t a l cbuyc l o g l do xlnop* irong do
HSP
g§p nbleu htfn ti^^'^ Loal I ? ^ oyt xlnop co ch^l {-Ion
tllo
tong O^tj - 1,4 o s e c . Vol pbtrcfng phop klcb t h l c b trong nao ,
Cac t l boo cbju anh htfo^ 15n nhou trong phpo vl rynrc
bc^
so v5l pbiftfng phop t'fcmg quon cheo oa oyc 00 toe glo
kboc
do duns ( 2 9 , 46, 9^., 133), Phon clcb ttfbo^t
t l n b tpo xun^T: cuo hoi ho^c vol t l boo cung o§t l u c , coc t a c
glo cbo thIy oah hn'cna^;,tfcebl gltfo cac t l b*?o xulc
blgn
tronf^ khoong 250-3y3 ©co tbeo bt^ng nIo n^ong, Sy khoc nbau
noy 00 the do phucfng j hop gbl tront* t e boo nbay hcfn so vSx
pbtfo^t^; pbap gbl nijOoi t l bao, Tv^ nhlSn phtfcfng phop kich
t b l c b trong nao cd nh(?ic dleo lo cdng dign klcb t h l c b t a c
dj^ng kbdni^ cbl l§n tb£.n t l bdo, co con co t h l len ca nhtfng
s y l quanb do, ida thai ddl ph2n ndo dnh htfqrng co tbyc
cua
qua t r l n h 5c c h l trong nao,
Pht^jfng pbap dtfa cac c h i t vao t l bao bSng v l dlgn dl l a
pbtf(A3g pbap thuyn l y l bc^n cd* V5l oyt Itfyai;: nbd coc c h i t ,
d i ^ quo v l dlgn eye t r y c t l o p voo t e bdo do cho phep
quon