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

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1.01
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-
2 -
H p
C L
y
c
no
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1
U
Riaii
II
T(JNG
iiUAN TAI LlS;U
I-
Cd
s3
hinh
thai
cua cac
qua
trlnh -&c
che
d ndroa
!• Te bao
tiTunf^
gian Uc chc trong
vo
nao
2.
GAM - chat
trimg ^an
TSC
che trong vo nao
5»
Cau
true modun va cau true tjp qu-szi
II-

Dan
li§u
ve cd che
tac
d\^t7
Uc
che
Ian
nhau cua
ndrun
vo
nao*
III-
Ciuan nifra
ve
tru3ng thy
cam va
tinh
chat
detector cua ndron
vc
nao*
!•
TrUdne;
thy cam cue ndron
2,
Tinh chat detector cua ndron vo nao
5.
Gc
quan phan

tlch
xuc
giec vibriss
17-
Dan ligu
ve
ve;l
tro 5c che
doi vci
b\?
ulnii
thanl-i tirjh
chat cua ndron vo
noc.
Treng
9-57
9-20
21-26
:6-32
32-57
Phan III
PHt/OlvG
PHAP
J«GHl£f^
Cl/U 5b-51
I-
Chuan
"bj.
d^ng
vit thx

n^igna
II-
Kxch thich hTi5n£j taji
III-
Vi
di|n eye
duu£.
f?e gJii ho^it tLalx dign
cua
ndron va dung
cho
vi
di^n
di.
IV- Ghi
di§n
the cua ndron va
aru'
ly
cac
ket
qua
nghien
cUu.
!• i)^g histogram
2.
Dtfng vectd-diagrainjii
3.
Tinh hf
so

nh§y
cam theo hudng
4.
Tinh th3i
gian
tiem
tang
V-
ChUdng trinlri thye ng;h i§iQ
38-39
30-51
«
3 -
Phan
lY
Trang
KUT
(4UA NGfilfiN CtfU 52-111
Chx&jng
I
3^-70
©AC
Tim
DAP
Ul^G
CUA
I^OROi.^
VraG
VO
ISAO

CAtl
GlAC-SOriA so
CAP
(Sj)
C
fxBO K)I VOI KICH
THICH
fcliia.'j
HOC,
KICii TKIGH Od
HOC.
I-
i>l§n
the dap
u^ng
trong vung chieu cua cd quan xuc
5^-33
giac vibriss.
II-
Phan
Ung
cua ndron vung chieu trong vo nao cam giac-
5'*—63
soma
Ichi kich thich
cac
s^i
vibriae myt CfiCh rieng
bift.
III-

Phan tfng cua ndron khi
kich
thich cac
s<Ji
vibriss
6^70
khac nhau trong
tnr3ug
thy
cam*
Chijfdng
II
71-'&4
bU BIETi iiOI Tlim NHAY
CAM
THH) EUDfiG
CUA
NOHON
DUOI
TAG
DVnO
CUA
KICH
THICK
i>OI,Ai^£i
HUONG CUA
Y£U
TO
TiiOI
GIAN

f>01
VOI SU
BIfcdfi
DOI
TR&^.
I-
By bien
doi
tinh nh^y
cam theo
hUdng
cua ndron
difdi
72-76
tac
dyng
cua
kich
thich doi.
1.
Sy thay
doi
tinh nh§y
cam theo
hUdng 5
cac ndron
ng*uyen
phat khong
nlrify
cam theo

hiidng*
2.
Sy
thay doi
tinh nhgiy
cam theo
hUdng
d cac ndron
nguyen
phat
nh§y
cam theo
hUdng.
II-
Inh
h^:fdng
cua yeu to
th8i
gian doi vdi sy bien doi
77-60
tinh nh^y
cam theo
hxrdng
cua ndron.
1.
t?y xuat hi^n
giai
do^
t'iam
dap

v^g
cua ndron
2.
Dien
bien
th5i
gian trong sy bien doi
tlnii nh^y
cam cua ndron.
III-
Cy^thay
doi
th3i
gian tiem tang cua phan
x/ng
khi 81-84
xuat
hi^n tinh nh§y
cam theo
hUdng
d
nhung
ndron
von khong
nh§y
cam theo
hUdng.
ChUdng III .
YAl
TRO CUA YEU TO

KHOWG
GI.J^
TBDm tJU
BIEN DOI
Tliai NHAY Qtili
ThiiX;
HUOriG
-
4
-
I-
inh hu3ng
cua
hvTdng kich
thich dieu khien doi
v5i 85-87
dap ting
hu:ng*
II-
by
bien doi tinh
nh§iy
cam theo hudng cua cac ndron
87-*00
dtr^i
tac
dyng
cua kich thich dieu khien
len
cac

sgi
vibriss co
vi
tri khac nhau trong
trtf3ng
thy cam.
!•
Sy bien doi tinh
nh^y
cam cua ndron khi cac
s^i
vibriss
chin
kich thich dieu khien phan bo d hai ben cua
s^i
vibriss
chj-u
kich thich
hutig.
2« Sy
thay doi tinh
nh§y
cam theo
hUdng
cua ndron khi thay
doi khoang
each gi5a
kich thich dieu khien va kich thich
h&ig*
3»

Sy thay doi tinh
nh§y
cam theo
htfdng
cua ndron khi
kich
thich dieu khien tac
dgng
vao
s^i
vibriss
nlm
khac hang
vdi fi^i
chj.u
kich thich
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-
TIC
dyng
md rgng trtfdng
thy cam cua picrotoxin doi
v3i
101-10
cac ndron vung chieu
thugc
cd quan
nhjn
cam
tii
cac
s^i
vibriss.
II-
Tac dyng cua picrotoxin doi vdi sy bien doi tinh
nh^y
103-10
cam theo
htidng
cua ndron.
1.
^oi
vdi cac ndron nguyen phat
nhay

cam theo
htidng
2.
Doi v5i cac
ndj^on
nguyen phat khong
nh§y
cam theo
hiidng
III-
By bien doi
th3i
gian tiem tang cua phan
Ung dxtSi
108-11
anh
htfdng
cua picrotoxin.
Phan IV
MJK
fipH
K£T
qui
112-129
I- Ve
d|c
tinh
dap
ung
cua cac ndron

nh^
thong
tin
112-116
tu'
cac
s^i
vibriss
n&n
trong vung cam giac-eoma
sd cap
(^j)*
II-
Ve sy bien doi tinh nhay cam theo
hu5ng
cua ndron vung
vo nao cam giac-soma
dudi anli huSng
cua kich thich
doi»
116-12
III-
Ve anh
^vTdng
cua yeu to
th8i
gian va khong gian doi
v3i
sy bien doi
tiuh nh^y

cam theo
hi/dng
cua ndron. 120-129
!• by
tich
h^g
ve
th3i
gian
2.
Tich
h;;p
ve
Idiong
gian.
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
trida
chuof:
cuo 000 khog
sinh
ly
bpc»
trong
nhtiiaf;
nacs
(^Jn
day
ainb ly
bpc than
iclnh da dgt
diHyc ahiitxg ffcanh tyu
to
l5n,
do
phat
bi|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 gl^i ty cbi3a«
ItbJmg;
vin de cc?
ban va co
clnb of:.uyeo cic nhif
qua
trlnh
tfcb bffp cac lu^ne btfiJng cdD»
vol
trS
cua qua
trlnb
^c chl tronf
bop^
<7<^Q£'
pbon xp cS dllu
kl^,
qu^
triob hiob
th&nh V9 duy trl
tri
oh^,v*v«**
tvt ISu iis Xitiu
hut
3^

quan
tdm d§c bigt
cua
I.P»Pavlov
(47) vo ttbilu
nh© cinb ly bpc
oc'i tlluG
c^
Mdn
z^
cZ^r nfaif nbleu nlfi$c khgc«
Cac
aha khaa
hpc Vi§t
nao
cung c5
hiiijpnt;
done
f^op
nboc
d^rib
trong
linb
vyc
nay
(1,
2»
45,
6I)« Tuy
v|iy,

ban
cbic cti»a
qua
trlnb tic
ebl«
eui^
nbi/
cc^
chl
yult
billn^
phat ^ridn
VB
uac dime
ttn:^Dg bd
cua qua
trJnb
nay
v5l
qua trlnh
ht?ng pbln oht? tbl
nao,
thl
troo^'
su^t
o^t
tbo^i gian dal vln
c^
chiia
dif^^

Kit
qua
ngblln
e^u
cua
nbiSu
t^>
tbi
tac
gia tr^n
pbatD vl coSn tbi gl$i trong vai chyc naa gla
day da
chfing
mlnb
vai tro
bSt
siic
quan
tr;mf;
cua qua
trlob i$c
chl
dli
v5i
toan
b<i
qua trlnb
td cia5c - cbi/c
Dang:
ou©

nao
b0,
B«
trtr^c bic
la
sy blnb tbaob
cac quo trlnb
then kinb ccf
ban
dam
bao cbo nao
b<^
t;byc
bi^o cbuc nang
tbdri^-
tin
«
dieu
kbl4n
cua
slnb
(la,
19, 39,
31^
32,
51»
^^,
137,
155)#
Tbeo

quan
dilm
cua
obilu
t;oc
glif tbl m
chl co
voi
tro
quyec djnb d3i vtfi
sy blnb
chanh
va thay
ddi
cac
trii&ng
thy
cao chuyen
boo cua cac
oc?ron
vmii;
vo
nao thj giac
(
66,
^)»
cvtii^
ahit viMig vo
nao com
ginc-sooa

(9) vo do do da
laa
nlly
sinb sy
cln tbiec phai »es xet lai kh^i nigm kinb dieo
vS trur&ng tby ca©t
coi
trtr&tig
thy cam cua
neuron
la
tS ch5c
«
6
«
ctfoe
able,
dtfcfc
blab tbanb
obo'
sy
guy
ty
co
cbpn Ipc ohtmg
loSng xung di^Dg btf5ag tS© dl tli
cac
td cb5c
than
klnh thip

ban
(X6»
It)*
Dfinir tho^i nhilu toe
gia
cung tlo
tb^
fi»Si
li^o quao
ob^t
cbi
glua
5c
cfc^
v5i
qua trlnb
pbgn cich thdcg trio
CM
giac
&
©5c
neuron,
d§c bi0t
la ay blnb tbanb tinh
chit
detector
^
cac
otfron
ims^

vo nao
tbi giac
(66,
67,
IJ^^t^iSS}
va vo
oao cao gioo-^ona (9)«
Hgoai
re
COQ
CO
y
kiln cbo
rln{r ay bile dSl **mls
deo*
trottg fahmp ©li
quan b|
tt»ng
tac
gil/a
c5c
acfron
dl
tpo ra
ay linh bo^t troog
vo nao
cung la nh&
qua trlob
^ cbl
(16,

l&U
So vol
trS quan trpag nbtf
v|iy
nSn bl^n
nay 5c cbl
da
trey
tbanb
vSo
dl tbu but
taanb ay
cbu y cua
nhilu nha
kbOQ bj>c» ^^§t troog ahtfng phtfctog
pbap co
hifu
quo
dSi
v^
vl#c tie bl§a
vai tro cua 5c cbl la
pbi^mg
pbap vl
dlfn
dl
(mlcroalectropboroa
o^lication),
d(^
CDC

cbit
d^i
icbaoe
(ootagwiitta)
cua
geiw»a-oa5ino-*utyric-axtt
(UAES)
vao
tl
bao tb§Q
kinb.
£|c
qua
ogaien
c5u cbo
thiy ay pbong toa
Se
cbl
Be Xao mit t£nb chit
detector cua
nifron
(V,
&3)
•
Bgi;PQ^c
lal,
cung
bSng pfaiJcmg
pbap vl
di#n

dl
6tfe gluta^et
vao cac
ncfrtm aha
cacb
dilo ^l kboaog 200 diet 3CX> mm
tbi
thay
suit bi0n
cac
natron
detector
m^i,
bo|c
thay
ddi tCob
nbay
cof&
da co
tru^
dSy cua cac
acrron detector (XO)*!ihllQ
tac gia cbo
ring gluti^at da boat hoa
cac
nc^roa
5o cbl
trung
glaD
va

kit
qua
la^
arixat
bl0»,
bo§c tba:7 ddl tlnb
nbay
cas tbeo bti5ng cim
cac
t;l
bao
tbap*
•I tllp tyc
tl©
bllu val
tro 5c chl trong vo
oao
d^l
v5l
sy
hltxb
thnnb
tfnh
nbay
c^a
theo
hvtm^
cua
coc nc?ron
vung

cac3 glac-aooat cbune
cbl
d'a
sii
dyng
is^t phtfcfng
pbap
DgblSo c5u
kbac,
dS
la
phifo^
phop
kich tbicb
ddi.
Fbucme
|*8P
nay cbo
phep
nghien
c5u
val tro cua qua trlnh 5c cbl
xulu bi§n dtf5l aab hating
cua
kicb thlcb h«i5og tSa
ty
nbiSn
troD^
vi#c
blnb

thanb
eac
tinh cbit
cua
oxyroo
vung cam
giac^aoffla*
Md blnb
tbufin lyl
cbo
vifc nc^bign c5u noy
la
b# tblog
ttbfty
cao
xuc giac
rSu eao
(vlbrlse)*
Val tro cua
5o
cbl trong vo nao
d<!l
v^l
sy blnb tbanb
cac
tlab cbit cuo cc^on
vung
cis giac-aoma se
co
tbd

dt/;^
las sang to
tbdng
qua
cbt>ng nbifm
vy
cy tbI
sau dSy s
1« %bl€n c5u
d|ic
tiob dii? 5tm cm
cac
nc^on
vut^
vo
oao Cora
giac-«osia Bcr
dp
(Sj^) dli v5i
kich thlcb tbicb
daog
voo cd quau nb^n cam vlbrlaa«
3m fbSn tfcb
sy blnb tbanb tinh
nb^y casi
tbeo
btidng
cna Dc^ron diiP^l aob btn^ig
cua
klcb

thlcb
hi^^g tSs
do
kich
thlcb dllu
kbllo gfiy
ra*
% sis bllu anb bii^ag
cua
ylu
tl
th^l
gian
dll
v^l
ay
biln 66l
tlab
obpy eioD
tbeo
hiring
cua
ac^on»
4*
IPhSm tlcb d$c tinh kbdug
gian cua
ay biln
doi
tlnb
nbay

c^
tbeo
biiwg
cua
DC^CHI*
5*
K^leo
c&u
anb
bifdisg oua
picrotoxin
(cbIt
d^l
khang
cua
GABA) dll vSi
sy blnb tbanb tinh nbay cm tbeo
btf^ng
va
trt/^nc
thy
coea
cua
OCJ^OC
vung vo nao cam
glac-soiBo.
qgam
do
thIy dtryc
boo

chit
cua 5c cbl
va
val tro cua no
del v5i ay
blnb tbanb cac
tfnh cblt
nay cua
nc^on*
^*btmg kit
quo
thu diiyc
trong
c6nt"
trlnb
nghiSn
c5u
nay cbo
thIy
rang t
- t5c
cbl
troog
vo nao do
kich
thlcb
hb^mxe t§a tao
re
^B
lam

volt bi§n bo^c Im
thay
ddl
tlnb
nhay eaa
tbeo
h\i3fne
•
8
»
cua fibtfn^ ncrron
trong
vun^
cbllu
cua
c<f
quan
nb^
cam
vlbrls««
S^TOog
dllu
klfn
klcb thlcb
dtfn
d;^
d$c
tlnb nay
hogo kbdng
co,

ho|ic
oo,
nbi/n^;
o'
ni5c
C^
khori^;
cao#
-
QtQ? lult
1*ay
ddl
tlnb
nbay ca^a cheo hu$tis
cua
no?ron
pby tbu§c
vao cac ylu
tS thJl
gian va
khdcc
gian
cua coc
klcb
thich*
«*
vi oSt ly chuylt nbDng kit
qua tbu
diiyc
coo

gop
phan vao sy
EBO^
r^n^
nbtfng kbal nl#9 hi#D
nay vl
c0
cbl 5c
ebl bdn
trong vo nao, vl val tro cua 5c chl d6i
v&l
sy blnb
tbanb cac
dge tfnb cn5G
nan^
i
cac
ncffron*
fflAH
IZ
Bifn nay ^a
co
nbilu tal
ll#u dl
c$fi din
sy
td
ch5e*
ch5c
n&tiQ

cua vo nao,
din
tlnb
cblt oua nc^on
va val tro
cua 5c cbl trong vo
nao*
Dgc bi^t
cac
ddn llfu
vl blnb
tbnl bpc
da
g^
pban
dang
kl
vao
vi|c glal qii^lt
cac
vln
di
trSn.
3/^
CQ
go
mm
m^i
mj
t^np

gvA
'smn
m
qm
lA pbucrog dl§n
blnb thai
- td
ob5c cus qua trlnb 5c
chl
i
nc^rdn,
nhaag vin
dl
difcfc cbu
y la cac
cl
bao
tining
gian 5c cbl
tr^Mig
vo nao,
asrlt sacuMh-^Blno-butyric - cbIt
trung gian boa bpc cua
irinap
5c chl trong vo
noo,
cac
clu
true sddun
vo clu

czuc typ
quan
(anaeffible)
cos
nc^n*
3^*
2l bSo
trun^:
fzimn
5c
chg
tram-
vj a»o
clu
true
vl
bao
vo nao da
duyc nghlSn cuu tb ISu
,
nbtfng
y
ngbia ch5c
nan£^
cua
tSuf^
loai te
l^o
vo val tro
cua Chung trong

«y
hoji;
d^nc
tlcb hyp cua vo noo cbo
din
nay vSn coo
dgng
dtf^W!
tilp tpc
tl®
biSu*
Jtbl de cgp d4n vIn
dl 5c cbl
trwifi
vo nao, cac nha
e^ldn
c5u cbu y
din chib nang cuo
te bao blnb
aoo» KhiSu
y kiln
cao
rln^
dly
la
nbufn^;
ntfr<3n trune glen
5c
ch5 da
dtfffc

chuyen boa va la
ca cbIt
cua quo trlnh 5c cbl (17,19,
20»
21, 49,
3X,
34t 115»
122,
150)*
Do
tfnh
cbIt
da dgng cua tl
bao
blnb
sao
va do klcb
tbiioc
cua than tl
bao gu;?
nbo,
nen vigc ph&n lopl
tryc
tllp cac tl bao 5c cbl trong vo nao ca vl
mgt
cb5c
n^ng
va
blnb thai
diu rSc

ph5c
tjp
(16, 17,
iO, 36,
37).
a
lo
3
Mgt
sS cac nha
kboo hoc
do
c^
gang
tits bl4u oil 11
Sn b|
glUa
clu
true
cuo tl boo
v5l
que trlob 5c chl bang
each
ngbiSn
c5u cac cbi
ulGu
gian tllp
(-:^3»
1^1»
1^7)^

TnfSc hit
i^tfol
ta chu
y
dio
clu
true
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 klrh
bien vl
di$n
tu da
pbgt hign
vo
ud

ta hai
loal
sclnap
s
- Lopl
I la
lo^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
acecylcholln.
-
I*oal
II

CO
kba 3clnap
b^p
hc^n,
vvn^^
boat
dgnf^
hep va
khong
CO
phan
5n^
v5l
acetylcbolln.
3ov
do
nSo
1^65
tcblznov (157) chi
ro
tMm lopi
I co
coc
tui yinop hlnb
tron,
con loal
I-^
co cac
tul srloap
blnb

hhv dyc»
Oac xlnop loal I
va
loal il phfm
bo
o^t
each
co cbpn
Ipc tr§n tb§n
vo
cr6n
coc
dendrlt
cuo
nafrdn*
lac lv<35 ittidersen
Va cync sy
da quon
SQG
thay
ylnap thuyc lo^l I tip
trung chu
y$v tr@n dendrlG
cuo
not^n thop thuvc
byppocoisp*
I'ront:
kbl do
xlnop loai
ii pban b5

ci:u
ylu
0 cr6n
than cua tl boo
nay,
6
tleu
nao
xlnap loal
i-£
t^p
trung;
tv^ti thSn ciia
te
bso pin:*klnj€
(73)•
'^beo
cac tac
ela
tr^n,
srlnop loal I
la
yinop
b.rng
ph*n,
con
xluap loal XI
la
:<inop
5c

cbl*
hSitQ 1966 Lccles (Vu) cune
thing
nbit v5l nb$c
d^nb
treo.
Theo
dtif;,
gla
choy't;
ylnop
5c cbl phSn
bl tren
than tl
bgo iS
boan
toon
logic,
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
chlln v} trl tbugn lyl nhit
trong
vl^c
dllu
h^o
qua trlnb
pbat
xun^^;
cue
no^n.
Iron£;
so cac
cl bso
soo,
uu boo ry
thuvn^-
tgo m xlnop
a
11
«
lopl II trSn
tb3n tl
uoo ubap
(la.,
123,
130)« fixoo
cua tl

bao rp too
nhonh rpnc
tbeo
bif5ng ckm
ngjjnt:,
ugo uhanb doo
rll quanb thfin
tl
bao thop» ^c dllc
noy
qUan
sot
tbiy
tren
tl
bo^
Tp
cuo
»o
nao,
cleu
noo vS
hyppocorop*
i>llu
do
cho
phep cac
nlia nghloti
c5u gla
tbuy^t

rang s tl boo
irp
co trong
tit CO
coc vung vo nao
©51
va
lo
tl bao
trun^:
eloa
5c che
(13,
19,
61,
9i^.
la,
132,
14G).
Can c5
voo
clu
true vs
d^c
dies
phan
nbanh,
o^m
1975
^ntagoth^l

('>9»
13^) cbla
coc
cl
bao
rp
tbanb
bal lo^l »
-
Te
b^o
rp nho, co cac axon
cgin,
ph§n
bl
chu ylu
tron^
l5p
il cua vo
nao»
-
TI
bso
rp
l5n,
too
nbanh
rpn^,
ph8n bl
tronf-

1^
IIX-V
cua vo
noo,
hh%n
d|nb
nay
cun{;
phu hyp
v5l
y
kiln
cuo
^orlQ-i'adilo
(1-i).
hi^ixiSc
c5u clu
true
cua tl
b^o
rp
tron^^
v^g
v^
d^Jng
vo nao
nt:i/&l»
tac
f:;ia
tb*y

nhun^:
cl bSo any ph&n nbenh
trong
gldl ban
nbtfnr.
phlan
tban^
d5n^::,
vudnj^
goc
v5l chiou t3nl bSl
trtmc crtttig taE»
liiaunt:
phllo
noy
xu;§n t5 l5p il xuSnc l5p
V,
CO chl§u
dot
lUuO-^QOU
fids va day
kboong iS)0
acD*
iiscon
cuo cl boo rp
clip
xuc
v5l
cac tl boo
tbop ahm

ti*oof:
phlln«
6
noo
khf,
naci
l^jf73
Glons(ll>)
cuut^
quoa
sot
thIy
coc
bl
boo
lopi
i
V5 I05I
Vi,
tu'cync
ty
nhif
cl boo rp
l5n
vo rp nho cua
Sentogouhal»
Cbun*:^
P^^-^n
nhonb
cbu ylu

vudnt^
coc v5l
tbuy
trt^c
trunj;
ti^o*
Goc
dSn llgu n£:hien c5u
vl anh
hiiD'ng
5c chl
gitJa
cac
oc»rdn
vung cao
gloc i oeo
cung phu
bjp
v^-l
coc quan sot ve
b(/5nc:
pnun nbonli
cua tl
boo
rp,
'Dual onb
hi^'ng
cua kich
tblcft
boo bpc,

blgu
qua 5c cbl the
hlgn
ro
c'
coc
nc^rdn nhm
tpoof
bmt)s:
vuctttj £0c v5l ranh chtf tb^p
(7,
^)m
sit hyp phiKfnc
pbap
cbyp
onb ty
d^n^
bing phong xg
,
pbi/otig
pbap
Golgl
Va
klnh bile
vl
dlyn
tii,
esc toe
gla fib|t
Ban (147,

1^6)
da cho
tbiy xlnap gll/a
tl boo rp va tl boo
tbap CO
ban
chit ^i^i^A (eaffinio-axlno-butyric 8xlt)» DSy
la
bSng
chSn^;
nuo bd
sun^,
cho gia thuylc tl
boo
r? lo tl boo
truo^:
gloo
5c chl trong vo
noo»
JNgoal
tl
ho^
rp
z^,
tl bao blnb
bul
cung^
d(;i^
col la
tl bao trung

gloc
5c chl
tron^j,
vo nao,
TbSn cua
Coc tl bao
noy
nlo
trong
Ic^
H,
xll
•
/ixon
cuo tl boo toa nhanh tbanb
hlnb
nhi/
chile den cbuc
(49,
1^,
150)
vo
too
re coc xlnap
lool II
tllp xuc
v&i dopn dIu ax«i
cua tl
baO
tbap

(1M,145,
146) •
Irong nliung
nam
gin
dSy,
cac
toe
gla
i*hgt
Ban ( 145
,
146,
146)
c5n
thay trong vo nao co bal
lo©!
tl boo blnb saO
khac cung
tbi^c I09I
tl bao trung gian 5c
cbl«
£0 la cac tl
boo blnb
©gng nb§a
va cl boo co bal bo
dendrlt.
v5l
pbtrcm^
pbap chvp

anb
b*ng
phong
x©,
coc coc gla
cuasf,
thIy
boa
chit
trw5g gian
cua coc xlnap nay lo
GiiBA#
Ttf
cac din
ll§u
tren, co
thi kit lu0n rln^s
trong vo
nao
CO nhllu
cl bao
trur-r;
gian,
Ve
nguygn
t:1c,
coc tl bao
nay co kha
nang gfiy
5c chl coc tl

boo
thop vo coc
ve
boo
khoc,
Tuy tbu§c
voo
v}.
trl
zQa cunc
cuo coc
phSn nhonb
cua
cbung
trSr;
tl boo tbap
so 05c dj
5c chl co
khoc
nbau,
thI
blgn bBng
sy
suy glsm nhleu
bay
It
ctf&nf;
dy lucng xung
d^ng
htftyng tSo

vo do do dllu boo qua trlnb
tpo xunp
cua tl bao
t;bop«
2.
G/>r.4- cbit
trum-
glnn
5c chl
tronr
vo
n.r|Q
.
Kbllu
y
kiln
cho
ring chit
trung
gian
5c chl
ct^
ban
tront
vo
aoo
la
gaomo-afflla^-'butyrlc-oxlt
(>,
5, 58,

71,
82
,
83,
93,
116).
»
13
«
Tbeo
din li$u
cua
B0t si
tac
glo
(55*
5^)
ohnc 60
G^}^i
trotig
noo
ttPtmg del
cao,
Ixboang
1,1-^j
ool5/g»
I^ung
phmmg
pbap
pbon*.;

xj donh ciu (Ir - GABA)
Iversen va
c^nc
sif (112
,
113) cbo
blet,
khoang
70 >v chit
nay
t|ip
trun^^
a t?n cung
tban klnh, 11
>» trSn syl
khdUt:
co myelin, 11
v^
trin dendrlu
va
6 ^ ti'Sn
cl boo
ijlia*
3y
tip
trung
OAB^
nhlSu
trong
t#n

cung
tbin klnb cb5nci
to no
pbal glif
vol tro quan
trpng
trong
din
truyin
xlnap
tol
t^ay.
TrOQg nbufng tbl nghi^ra
klcb thlcb tryc tllp len bi
m$t
vo noo,
doner th<^l v5l vlfc
ghi
difn thI
5c cbl
J*
ncardn
,
tac gla da xac
djnb duyc
non^
d^
cua
GABA
t;anc

Ifin
g^
3
Ian
so
v5l
l5c
cbtia
kich
thlcb»
IJIU
kb5
ion
Ca**^
tronf;
cd
than
kinb
ohl
khi klcb thlcb vo nao
nSnt
d^J
QiiBA
khong
tling ISn
«
Dllu noy
cb5ng
to
Gn"*^

lo ylu cl
kbdng
cbl
thllu
duyc dl
glal phone
CABA,
tiXcPng
ty
nhu tinh chit
cua coc fooa
chat
trung
glon khgc
da
dtipyc
cdc^
nb§n
(112,
113).
^o
cvng
1P
B^t
bing
chSfif^
ou^
dl
kb<In£ d}.oh
GAB/,

la chit
d^n
truyin
trong
xlnap 5c chl cua
vo.nno,
I^htfoG
tbl
nfchifiD dtPo
coc
chic vao
tl boo bang
phtfcfng
pbap vl
dl^n
di
(mlcz-oelectrophoaras
application) cua
Knlevlc
(116,
117,
IIB)
va
Alexandrov
(3,
4,
5)
da thu
dtfyc kit
qua

CO y
n«:bio
don^
kl,
cbl ro anb
hi:f(5^g
cua
Gi^A
l^n
bo<^t
d^og
tao xunt
cuo tl boo
thin
klnb vung
cac
gioc.
Blnb
tbiiWig
nc?r6n
phot
xun^j tiKme
dll
6n
dj^no,
v5l xiln si
kboang
5 xung
trong
BK?t

glSy.
Dwl
anb
hiWng
cua dong
dl§fl
dl
GAB^,
co
earns d$
20
h^,
^
70
>^
t^ng
s3
oc?rdn ghi
duyc,
tin sS xung
glaffl
xuong con
0,6
xung
trong
ns^c
glay,
6 ogt ol no^on
kbnc,
xung

d^ng
libdng xulu
blgo,
Cgc no^rcn
noy do
b|
5c cbl
bo5n
toon,
TrontT
dieu
klgn
trOn,
v5i difn eye gbl dl§n bSn trtmg
tl boo, coc coc gia da ghi
d;
ye
dl§n
tbl
tSnc pbSn
c;,^
(hyperpolarlzation)*
I^bti
vgy UA£i
do
gSy tru ph^n eye aong
no»rdn a
vo noo
cut^e
ty

nhi^
dl$n
tbl
ijang pb3n eye
troni::
xlnap
glu-a
tl boo tfo cbl
iienshaw
va tl
baa vpn dyng
tuy
sln^!;*
s
14
9
Myt
trong
nburng pbusmg
pbap xac
d^nh
co blfu
quo val
tro trung
gian
boa bpc cuo
GAB/*
lo dung cac
chit
dll kbang

(antagonist) cua
GAB^
tac dgng len
nc?ron
(79,
6^-^,
94, 105,
106,
107,
135,
136)*
Gac cdng
trlnb
ngbldn cms
cus Curtis vo
c^ng
sy
(04-66)
da cho thay
00*51
tac
d^fnp,
cuo
alcalol^^-blcu
-
culln (antagonist
cua
GAB^),
bl^u
qua 5c

onl
cua
OABA
h%
sit
hoan
toan*
kbdog
ahatii^
tbl, nhilu kbl xung
ahn
cua
neuron
con
t^g
bG^c
blnb
thi^4^« lac
gla cbo
s&c^,
trong dllu
klfn
ty
nbiSn
v5l
m^iz
m5c dy
nao do,
ncfrdn
da

cf
vr&Cf:
thai
b}.
5c cbl,
vl
tbl cac
d^mg
cua
blcuculln kbdng nbting
pbong toa
bl^u
qua
cua
GiB^
so
con glal pbong
nctrdn kbol'*5c
cbl
truPcfn^
lyc^
Sou
do
Sllllto
(135,
136)
vo
HlckS
(107) kbl
ag:hl§n

c5u sy
blln
dll
trtf^n^^
thy coo cua ncfrdn vung cao giac cunc
nb§n tbIy
hlfu
qua cua
GAB^
b} mit
boon toon kbl cung
m(^t luc
cbo CABA
va blcuculln tac dyng
Ito acni'dntf
Kit
qua tbl
nt"bi§ffi
cua
Stcwat
va
Cf^ng
sy (149) cbo
th|y
ngoal
blcuculln
c^
co
myt lOal olcalolt
khac lo

plcrotoxla
cung lo
chit
dll
khon^-:
cua
GA3A»
i^bl
dtfo
voo tl bao,
cfalt
nay cung
las giom bi^u
qua 5c cbl cua
G^BA
tdcmg
ty
nbii
blcuculln, Cac
cdng
trlnb
ngbign
c5u
gin
day
nhit
cua Alexandr
va
c^ng
sy (9,

IG,
11) cung
cnung oinh
tac
dpn^:
cus picrotoxin
ttfcfng
ty
nha
blcuculln trong quo trlnb blln dll
trii&ne
thy cam
Va tlnb
chit
detector cua
ncTcc
vung cac
giac &
tseo,
Kbl
x^^>
xlc
cac
cbIt
las
glao blfu
qua 5c cbl cua
GijBi
Sexkov
(55) dya

trSn
col
ll#u
thyc
ngblfm
cua nhilu
uoc
gla
da
duPa »»
y
kiln
vl vol tro cua
penlcllln,
Tbeo
dn^,
penlcllln
cung la
chit
dll
khanu
cua
GABA,
CO
^ac
d]fn£:
Im glam
5c cbl
cua
GAB A

trong vo
nao»
i;o do penlcllln da
d*/yc
dung dl
too
nfaimg Old
blnb
dpog
klnb thyc
ngbl^o,
GJ
Qhi toe dyne
dll kbang cua
blcuiitlin
vo
plcrotcwrln
dli
v^
CAi^
khac
nbou«
Blcuculln co clu tao
tiKfug
ty
nbu phen
ttf
GAB/J
va cung aha
GABA,

no
kit
hyp
v5l **phan nh§c
bllt**
cua
vung thy
cas trSn ©ong
neuron*
vl vyy ma
sy
kit
hyp
noy c?ang
^
15
«
tlnb
chit "canb
tronh".
Con picrotoxin cbi
kit
hyp
v5l •*pb5n
dllu
boa",
nan
tac
Cms
kbdng tbeo

ccr
cbl canb tranb (149) •
Soog
du tbeo
ctf
cbl nao, cac
cbIt
dll
kbonc
cung
locn glao
bl^u
qua 5c cbl cua
GAi>A
dll
v5l oOvQn nbif
r^au,
Dllu do
kbing djnb thStn
gia
tbt^ylt
cbo
ring
GABA
1Q
chit
trung
gian
5c chl trong vo nao,
Tron^i,

Coc t|in cung
thin
klnb
i
vo nao
GABA
di/yc tyo
ra
bIng
con
dLro^.g ^cocboxln
boa
axlt glutaslc v&l
sy
tbaQ gls
cua aen
glutaoet decacboxylaza
vo
coenayo Bg
(113)•
I^ng
tbtfl
CABA
cung duyc
*o^
tbu
IQX
v5i
oi
lye It/n (Ko »

d^S
(K» m
2,2
,
10^5 M)
(112, 124).
Bhm^
kit
qua
ngblSn
c5u
v^
slnb ly
ncfron,
vl tl bao
hpc,
vl
di^c
ly hpc va
bIng klnii bien
vl
dlgn ttf
da
Isisang
d].nb
GABA
CO du
vioM
chuan
myt

boa
chit
trun4'
glon
ifc
cbl
trong vo nao
(55)»
GABA
diryc too
ra
tron^:
cac xlnap
loal II
(xlnap 5c chl)
gllio
tl boo sao vo cl boo
thapi gSy
tfu
pbSn
eye
san^:
tl bao thop va do do 5c chl
kba nSng boat
d^ng
tyo
xung cuo cl bao nay,
Nbtf vgy
ca ebl
5c

cbl cuo te
bao triuxg
gian
dll
v5l
tl bao tbap lo
nhh
chat
trunc
glon hoc bpc
GABA,
bay
nol
each khac, 5e chl co ban
cbIt
G^BA.
vl v§y
x5
dpnc
G^B^
va
d|lc
bi$t
lo cac antagonist cua
no
trooB vl|c nghlSn ctfu
quo
trlnb,5c
cbl lo boan toon
byp

ly-
3» clu
L-ruc
mddun
vo clu true
tan ouSn
(cnssable^
Sy
bllu bllt
vl
ntsuySn tic 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
ly
bojjt dgnc
cbune*
cua ca

b^
noo,Cblnh
vl
v^y
ma
vlgc ncblon
c5u
Qg;uyen
tac
td
cb5o coc
agnr.
ncfrdn
duyc col
nbi7 cs^Jt bL?5c
quan
trpng
tronr
vlgc kbass
pbo
nbifnr
bl m§t
cua
b^
thing
thin
klnh.
ic^
Quan
nl|c9

vl sy
tBn tai troue
vo noo cac tl boo trung
glon 5c cbl
CO
onb
hiftJiag
rat
l5n din kbol nlyni bl#n dpi
vl
sy
t<5
cb5c
- ebue
n^ng
vo
noo
o5i,
Tbeo kbal
nlgo
noy tbl
cac
nc^dn tgp
hyp
tbonh
coc clu
tr&i oddun
vo cau
true tpp
quan*

•)
f^^*M^
«
coc
no^dn t§p
hyp
v5l
nhgu tbanb cac
cgt
hlnb try theo chiSu thing
d5n^
tpl Is coc
raddun, fcfidua glo
nbimg
tl bao
thin
klnb cung thyc
hlgn myt
cb5c
n^ng
tbdng
tin
nio do (16, 19,
^1,
27,
aB,
43,
54,
13I,
161).

Rgu'S^l dSu tl€n dl cgp dinclu true mddun trone
vo nao
la
i^ountcastle
(^3)
va
xi»«ell
(131)*
Kbl
klcb thlcb
l$fi
da
0
khi, cac tac gia
nb^
tbly tr§n vune
vo
nao
cas giac co
cac
cyt
ncfrdn
bufng
phic,
c^nb
do co cac
cyt iKd^on
5c cbl
•
l^bun^i

luc do cac
|»ac
gla
trSn cbua
glal
tblch dtryc
cb5c
a&cs
cuo coc
cyt
ncrron nay*
^u
nh^
co sy phot
bifn
ra coc
ncfrdn trung glon 5c chc
trr^ig
vo nao, quon
nl^is
vl clu
true
cyt din din
dtfyc song
to*
»b5n^"
din
ll^u ngbien ctfu vh dlgn
sinh ly da cbi ro ,
kbl

dlgn eye dl
quo cac
l^*p
cuo vo nao vo
ludn ludn
vudng
goc v5l bl
Q§t
vo nao,
tbt/acg ghi doyc
coc
wjron gllng
nhau
vl
o§t
cb5c
nang*
Cac
c^t
ncfron co
chSc nang cling
nbau
gbl
duyc
c?
cac vung
cm
giac (43), vung
th%
giac (108), vung

tblnh glle
(14,
15) vo vung
vgc
d^n^^:,
(74,
75).
^^bu v0y
clu
true
c^t
la
d^c
dllts cbung
cua vo
noo#
Tbuylt
clu
true cyt
ngoy nay duyc
gpl
la tbuylt
tsddun,
Tbeo
^ountcastle
(43) dan
v}
ccr ban cua
oddun
lo vl oddun

v5l
di7^^
klnb
gin 30
tsco,
Cac ncfron trong
©yt
vl reddun co
pban
5nt
cbpn
Ipc v5i myt
thorn
si nhIt dj-nb
cua klcb thlcb
btf&g rSi3
va lo
dc^.
vJ.
stf cing nbIt
cua quo trlnb
xu?
ly
tbdng
tin*
Khoang vol
trSm
vl
sddun
t^p

b^
tbonb
mddun
lt?n,
cl clu
C90 ph5c tgp
bc/n,
l^ddun
l5n co
dpng phllo,day kbooog
TT;
i
V-W^#5J
«
17
30 - 2G0
Dcm,
doi ^^00 - 1000
ocm,
xuy§oc5 l5p li t5l
l^
V
cuo
vo
noo*
Trong oddun
l5n dlln
ro coc quo
trlnb
xu"

ly
tbdn^r tin
pb5c
top
licfn v5i nhleu thoo si khnc
nhau
cuo
klcb
thlcb*
i>o
lo
dirn
v|.
xu
ly vo
tru^ln dpc there
tin,
tllp
xuc
v5i hong loyt
dtfo^i^:;
%?ao vn
ay^
f^c
dwcni
ro
('^3)*
t>btf
v^y,
vl

n^^uyen
tic
cau
ta^,
eac
mddun
l5n
co
cs?
s^
blnb
thai
tacmr;
del
ro,
nbtfn
cac vl
ludtiun - dc?n
vl
cb5c
nanf'
co* ban -co
kich cbuf5c 3G
oco
khdnf
phu hyp
v5l
sy
phan nbanh
cua

ogc cyi thin klnfi
dl
til thal^ius t5l
vo
noo,
Tbeo
tal
ll&u dQ dupyc
c
nj:
nhgn
(5G),
®0t cyi
than kinb
thalamus
-
vo noo
tSa
nhonb tronr vung cbllu
6
vo noo
trong
pbam
vl
CO diio^T klnli 30U-iiOO ocs
vo
tllp
::&;
xlnop
v^

gan
5000 ncfron*
iJbimg
vl
©ddun v5l
dMn^
klnb
gen
30
oco cbl co
tbl
boo
i^hm
110-260
acfron (^3)*
5y
fflSu
tbuln tiny
chl co cbl
glal
chlcti Khi
phat
hlfn
ra coc tl boo 5c cbl
trong
jsddun
,
Cac
tl boo 5c
ctil difyc bav^s

P^l<j
b^
coc
luSng
xun^^^
dsng
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 *^
chl
quonh

c^c
mtcoc bt/ne
pban
•
Do
kit
qua cuo qua
trlnb
5c chl,
sy hung pbin
duyc
kbu
tru
trong
phafts
vl cuo
dc?n vJ
ch5c
nan^>
^^bt^
vvy
vl
ffiddun
lo
dcm
vJ.
cti5c
non^^
diPyc
hlnb

thonii
diTol
Goc
dyne
cuo coc
tin hi^u
bir5n£.
vam^
Hlnb
doni;
vn
kich
cbtf5c
cua no
phy thuyc
rat
nbiau voo
co'
s^
blnb
thol bsn
dau^
libim^:;
cd^^
trlnh
ngblcn c5u
ve
blnb
thai
hpc cuo

i-arln-
f odilla (122)
gop
phan
clol tnlch
vol tro
cuo
5c chl
trong
vo
noD del
v5l
t<^
ch5c
-
ch5c
nan^:
cuo coc
©cjdun.
Joe gla
da figrblen
c5u ty oi sy
pbjn nbonb
cuo tl boo rp
trong
cac
l5p
cuo
vunc vpn
dyng.

Axon
cuo
lo^i
te bao
noy
too
nbonb
t^ong
phao
vl
nhif^:
phllo
tban^,
d5nf:
v5l ciileudny khoong
200
f3ca)
dal 1000
- SOuO
occ;,
vudnfv;
.:;;
oc
v5l
bl
(u§i
vo nao ,
xuyen
tu
l5p Ul din l5p

V
vS tl^l^^WssMM^'^'o
co coc
tl
boo
thop
nIo tronc
phleri.
^''bu
wy^by^^hl^^,^^*
t;o^
rp
-
tl
sif*
*-K1«
AZ
^^^
—
•**^'>c4f eye no^oti
c^:dv*^<9>-?blln^Ai^ae
d5rig.
a IC -
Klcb
tbi/^
vo hlnb
6ont,
cuo cnc
cyt
noy do

d^c
diem
ptxbv^
nbonb
vo chieu
doi
cuo
oxon co
boo rp
qvye\^
d^nh,
Tbeo
Sentagothal
(50) trong vo 000 coc raddun co blnb
done vn kich thtr5c
kbnc nhau lo
uuy thuyc voo
clu
tpo
cuo tl
boo
rp*
TI
boo rp 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.
TI
boo rp
l5n o» l5p
III
tao
thirinh coc oddun co dgng pblln,doy
kboong 23-3O
oco, dal
£:ln
5G0
oco vo sau
t5
l5p
U din
1^
V,
Tuy cbo
din nay vSn cbua
co
oyt

quon
dlla
thing;
nbIt
vl blnb
doat:,
klcb
thu?t'e
cua oddun, vl
ctf
chl
coc
dyng 5c cbl
l|n
nbau trong oddun, vo
gli^
coc oddun,
nbi/nr
coc cbo
ngbldn
c5u
dIu edng nb^n ring
ylu
ul
khdnf:
tbl
thlfu dixyc tron£
clu
true mddun lo
coc uti

boo
trun?^
fclon 5c chl, Cac tl
b^o
noy
dong val tro
quoo
urpni;
tron^^
vl0c v6 cb5c ciou hoo boat dyng
cua
oedUQ
v5i coc o5c
dy pii5c typ khsc
nhau,(16,
18,
49,
5G
,
52,
5^).
b)
gap
Q^h
C^aagp^lt)
B^n^^
phuc?n^
phop
s5 dyne nbllu dlfn eye (c;-5
vl

dlpn
eye),
ghi
hOQv
tlnb
dl|n
cuo
oyt
nhon ncfron,
kit hgp v5i
ph\f0tk£ pbgp xoc dj.nb
tlnb
chit
va ffi5c
dy
ti^tii:
uac
GI^O
cbung,
oyt si toe
glo
(<-tj
«.
>2,
34)
da cbo
tboy
m^i
anb
htKm^.

cua
coc
tin lil^u
htf&c^
t§o,
troofy
cac
t^
chSc uhln
klnh,
oa tr*/5c
blr
lo
tronr,
vo
bon c5u djil
noo, hlnb
thooh clc t&p quin nc?ron
v5l hlnb
done'
vo klcb
thi/'c
khac
nhou,
Tbap
Kogon
vo
c^tc
s^
(^,

3Gt 34) thl c#p quin noron
lo
oyt ohm
coc
no?ron cure i
trong
tr^nf:
thol
biinf:
phan,
du^
bao
quoch
bSn^
vonb
doi coc
ul boj
bj.
5c chl,
Vg
c^v.
uruc
vonb
d.ol
QDC
V •
r.o3
b}
t'c
chc, Vo clu

true
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
tbsl bimg pblo kinoc nhou
vn nao tren
cun^r myc a5c
td
ch5c
cuo
cun*?
nbnn
xa.
th^ia chl lo
cung
cron
ffiyt
l5p

tl 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 true khccpr
c:lon
theo
chilu RcnnCf
kboc
v5l ofidun

co
c5u true
tbeo
chllu 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
tin hlgu
h;?^5nf^
tao, voo
nic
thane
tlln
boo cuo
dyn^r
v^t*

C
dpni:
vgc bye th^p
(bo sot,
li/5nf:
tb§)
kich tbif5c
coc
tgp qoSn
rat
l5n (40U
-
500
ocm), coo
c?
d^ng
vgt CO
VU,
dt^^^nr
klnh
cuo
t^p
QuBn
chi
khoonc.
15^
- *i'X
meo.
- 1§p quSn
chi

blnb
thinnh
kbl co
klcb thlcb
hi^g
too ,
vl
VQy
no
kh<inr tSn tpl vinh
ctfu,
i^i
toe
dyn?.
cuo
s§t I09I
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 cc
thl
sdnr
ludn ludn
blSn
dyn^
sen clu
true
kbdn;
{;lan vo
chc?l
gian
cuo
t§v quon cun??
ludn ludn
tbgy
del,
i^ol
coch
kboc,
coc
t§p
quSn di/yc
hlnh
t:honh
cheo

ngu^/Sn
tac
xoc su*t» thln^r
kg,
ngbia
la sy
thom
gla
cuo
coc
nc/ron
voo clu
true
c^p
quin dlln
ro theo
nf^^uy^n tac
xac
sUwt,
con
higu
quo
boot dynr
cuo
t^p
qu»in Is
kit;
quo
tblot:
k§ hogt

dyof:
cuo cac
nc?rdn
clu
thonh
t|p
quIn
do,
-
Vnnh doi
coc
ue ooo
bj. 5c che boo
quonh
cyp quBn
cung
ludn ludn
rhoy ddl du^5l onh htfonr
cuo
kich
thlcb
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 sl
tif^Vi^
quan
difcfn^
v5l sis trj tv^/^t doi
kho coo,
lig-tfjc Ipi hf

so
ttfcfng
quan
gl5n ndton
n^o
crong
trung
tuo bt/ag phan vo'i no^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/
cbl

hlnh
thonh
coc
c^p
quan
ncTrdn
cho
din
noy
v8n
con
CO
nbuj]^
y
kiln
kboc nhou,
Tn,y nbien
cusif;
co
myt
00
quon
«
20
a
dllo
Chung
nhtf snu
s
- Coc

ncfrdn tboo elo
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
xult phac
tu
ayt
so thy cao
chl
n^^opi
vi boy
tJlr
c^yt
nhom cl

boo
naty i t<l
ch5c
ch?*n
klnb
tbip bfSti
cheo cung
phon
xg,
-
^c? nh
hlnh
uuoi trSn
chi lo
tlen
dl can
thlec
ban
^lu,
eon vol tro cd cb5c
ndn
coc
t^p
quIn Ipl tnuvc
vc coc dong
tin blgu
btf^nt;
t3®,
Coc
kit qun ne:bi§n

eSt'^
cua
nliieu toe
glo
da
cbSnr
ralnb
ring cf dlBu klyn
n^n,
all
quon
hg
tutfn^-
toe
gli/o
coc
nc?ron rit ylu
vo kbdng
mong ulnh dj.nh
ha&n^
ro
rfc,
-
Hlnb
^vin\^
vn klcb
thi/Sc
cua
tgp quSn
con

ph-^
cbuyc
voo quo trlnh 5c cbl xoy ro
cJ
vo noo,
'^o
trlnh noy chu
ylu
do toe
d^ng
cu?
coc
t*
bao 5c
ch*^
trunr cinn tgo non
(3i:)*
!Dheo
quon
dicD
cuo
coc
cac gla trin day tbl
odl tpp
quSn
ncfron
dt/y'c
coi
nhif
opt

do"n
vJ.
ch52 nano
at/
dSng nb^tj ,
^bilu
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 rla
boan
thonh
ojt 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 thl bleu oddun
nbif ©yt
bloc ch5c
nan^v,
I^
con
CO
nJitfn^:
dieu kboc
biyt
ve chi
tilt,
coc
toe
gla
cue ca

lisi cuan diliii ^^u
di den
kit lu§n thlnt nhlt
lo coc
no^ron
kh6a^
bogt
dyn£.:
oyt coch riong lei d(/5l
cac
dytf
cua
coc
cin hlyu bu-^ng tSo
ebun^'
ci?ye xh cb^c Iji uhonb
totie.
nhoo
tbeo bl
ngonf;
vo bl
doy
cuo vo noo,
tco uhoah
coc dcfn
vJ cbSc
naQti:
stf
dSnr:*
Cblnb

co chl noy cho phep
glol
thlcb
du^jic pbSn
nao
nbtfng dgc
tlnb
co? ban
trong
bjpt
d^n$
tlch hyp cuo
by
noo,
Thoo
glo hlnb
uhonb
coc
d^c
tinh
noy khong
^hl thilu
dtryc vol
tro cuo coc quo trlnh
t;c
chl
tronf
vo noo.
uy-
•piii;

^^u
iii^
CO ciiu
TAG
LU.W
bc
G:U:
MH
miih
CLL
ItOI.Ui.
VO
ULO
throng sinh
ly
a;/ron,
coc thyc
n^bi§m
cl lien
quan sSn
vlfc clo
bllu
toe
dyr^
5c chl ben
trong
vo nao
6a
vo
dsioe

diiyc tile bonb bScg nbl§u pbtfcrng
pbap khac nhou*
V5l
pbuc^ng
pbap fc:bl
dlgn uhl cgooi vfi
too,
bong loat
cac
cdnc crlnb
(25,
33»
^^2,
56,
^Z^
63,
*?9,
127)
dS cfal
ro t
dif5l
anh
btfd^ng
cuo
bit
c5
oyu lo^l klcb chlch buSng t&n noo
,
dnp
5D£*

CUO
nitron
vo
cao tbt^cng chlo thonh
nhilu pha*
i-'
da el
nt?ron,
pbo
dIu bt*Qg phin
(bi/n^?;
phin ea
dp)
vcl tho^l
glon
ngin (kbooUG IO 3O ooec) tSn si xunc tang
l§n.
Sou do lo pha
5c cbl keo dal
(350 - 2P0
ooec),
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
dp)*
0 ©yt ec It
ncfron,
5c
cbl phat
txlen
n^oy
sou kbl klcb thlcb

btfSnt ^^^
^k/
ro ( 5c
cbl
6CP
cap),
6ou
do
ay
phot
xun^
cuo
ncfron
oo'i
tang
ISn*
Sht? v§y
tron^^
d^p
5ng coa
t*
boo
tb§n
klnh co xen
ki
gloi dojn
htrui^
ph'n
vc
giol doan

5c cbl,
3$c dlecs nay
quon
soc tbIy ^
coc tl boo
tbu^c tit
ca coc
vun{
vo
nio
kbl co
klcb tblch
d§c hi§u
xoy
ro*
Can c5 voo do
Serkov
(51,
52,
53,
5^^,
53) do
kit lu|nt
1-
tic
cbl lo thonh
pbSn
kbdn^^
tbl thllu dtfyc
tron^

dap
5n^5
cua tl bao than kinb dci
vtfi
cnc klcb
cblch btf5ng
tgo,
a-
"Otfp
5nc,
cun ncrron
©one
d^c
tlnb pbo,
ti^
lo
trSn
o^t
ncfron ca bal quo
trlixh
htpn^
phin
va 5c cbl
dIu
xult
hiyn vffi
il^loh
ty
tbo'l
gian khoc nbou,

tJc
chl xult
bign oyt
trong
trtf&ng
hyp tren gpl
la
5c
cbl
btf5ne
cao,
mQc
trong
nhung
dang 5c
cho
cuo vo nao,
M^t
trong
nbung phiK^ag
pbap
ngblgn e5u
co
blyu
qua quo
trlnb pbat
trlln
vo keo del cua 5c cbl
btf^ng tSo
la

phuwie
m
22
^
pbap klcb
thici:
ddl,
Klcb
thlcb tb5
nblu
gpl lo
klcb
thlcb
dllu
khllni
klcb tblch tbu
hoi Epl
lo
kich
thlcb
b5no Hal
klcb tblch coch
cb^v
nhtfnt:
kboon^
thitfl gian nbot dj.nb
gpl
la
tbSl glao
gl*^

bol ^^ch
chlch*
t^gir&l
dau
tlen dung
pbtfcfng
phop noy lo
Eruca
vo
c^nt;
sy
(bO) v5l snyc dlcb tlo bllu
trtf^n^
thy coo cuo
ncfrtKi
vm^^
cam giac
(y
oao,
l^lch
tblcb
htfng
duyc
dyt
voo
truct cOo
cua
trtfo'nfr chp
cotr,
klcb thlcb

diau kblln
dtfyc
dyt tpi
coc
diC'm
khoc nhau
tronF crtf^)g
thy
Coo*
Inb btfotsg
5c chl do klcb tblch dllu
khiSn gSy
ro da
lao
eiim
sw'
dop
tfn^^ cua ncfron dll
v5l
klcb chlch
Lu^> lya
voo
kit
quo noy tac gia da pbSn
lopi 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
dl
n^lSn
ci/u anli
but^
cua 5c chl
del
v5l dap tfng
•^51^^"
cua ncfron kbl klcb thtcb dllu kblln nam
0 coc
vJ.
trl
kbac nhau trong
trtf^nc
thy
casi
vo
tbSi gion gltfa boi h kich
thich
cwu^
khac
nhau*

C^a
phan tlch coc toe
^us tbIy
anb
htfc^
5c chl
couf;
oynb
kbl
kich cblch dieu
kblln
con?;
nao
gin
trung tao
trtfStig
thy cao va
bigu
qua 5o chl
l5n nbIt
kbl
th^l
glon
Gliiia
b*«l
klcb tblch
khdnj^
qua
20
moac*

Tuy
vSy
traa:
cnc tbl
ngbl|o trSn cbtfo
co
odt ch&xg
olnh
tryc tllp noo
kb^nf:
d}.nh
5c chl
htftfnfv
turn co ban
chit
sau
xlnr^p,
i>l
glal
quyli;
van dl noy, coc nho
nghldn
c5u da
onus
9§t
trong hai pbtfcfng phop sou
t phucfnir
pbap gbl
dlgn
thl

trong
vi
boo vo
pbtftyr^
pbap
dtfa
coe
chit del
khong cua
chit
trung
(3^1on
5c cbl voo trong tl boo bang vl
dign
di,
Kit
quo gbl
dlgn
thl
trone ^e
boo da chi ro,
dign
tbl
5c cbl sau xlnap
*•
Inhibitory Postsynaptic
i'ouentlal
(XP^
)
dc^c

too
3?o
tren ncfron kbl co
kich
tblch tryc tilp
ISn
bl
D;^t
vo
noo
(55), len cac
syl tbSn
klnh
vholomus - vd
noo(l02)
va do
bit
ky
oyt kich
thlcb
htf5nfi tao noo
(53,
5^*,
55,
56
,
62,
63,
€>4,
65, 102).

«
23
MSo
1977
iiellweg (102)
kbl klcb tblcb
d§y tbSn
klnh
Introorbltolls
(trorv^
do co nbonb di
ttf
thy
can;
thl vibriss)
do quon sot
thIy
bo
109!
phon
5ng
cue ncrron
VUJH'
COO gioc
aooQ
t
L09I
1 s
Slyn
tbl bung

phIn
sou
xlnop
-
xung—Sxcltatoxy
Posta^^naptlc
potential - spike
(Ei^SP-splke),
L09I
2
% Blgn
thl
5c
chl sau xlnop
ilx'^'^
ijopl
3 t
do CO dlgn
cbl hung
pbIn
sou xlnop
•*
xung
—
dlyn
thl 5c cbl sou xlnop
(iiPSP ~
spike -
IPtJP)
Lopl 3 thtfmig ggp bcfn

co*
Trong nbtfng tints
1979-19^^
Pbaooskl
(62, 63,
6^,
65)
gbl
dlyn
thl trong tl boo vung cats gioc khi klcb tblch
d^c
bl^u
voo coc thy coo
bne
vibriss cunc
QUon
sot dtfyc pbon
5ng ttfcfng
ty,
%^ol
3^0
Coc
glo
con ghi
dtfyc
logl
EPS**
^ IPSP
(kbdng;
co

xung),
Coc
n^hlGO
c5u tren
ncyron
vtaif:;
tblnh
^lac
vo
vunc
tb}.
glac
(53,
3^,
56)
oun^:
deu
gbl
di^
coc
logl
phon 5ng nol
tren,
Kbu
v^y
trons chonh phIn dlgn
thl
tron;:
te
boo

cua
pbIn
l5n
neuron dIu
co
dlgn
thl 5c chl sou
xlnop*
So
sanli tbtfl
(^lon
tilo tone
cuo
^SP
va
1P£5P 0
ncfron
kbl CO
kich
tblch
htf5nc
tSo,
nga^i
ta
nh§n tbIy
IFt^
xult
hl§n cb^is bcfn
i-PSi'
-vhoong

0,fc
-
1,2 osec (53, 55,
65,
IO2
,
15^).
sl llgu
noy thu dtfyc
khonc nhling 6
tl boo vung coo gioc
kbl klcb thich
ngogl
vl,
00
con
^
vuni,
tht
gloC
kbl
kich
tblcb doi
tht
vo
kbdnc tboy
ddl kbl klcb thlcb tlo
ubj
(55 )
Dllu do

cb5c£
to
r^n^;
sy
n^tfn^-
trg tron^r d2n truyin dl^n
tbl
5c chl sou xlnop xoy ra
cggy chlnh tronc
vo
coo
ch5 kbdng
pbal
trSn citrSnrr dSn truyen
u^:ool
vo*
libtf
v^'
len
tron^
vd noo
a
24
ar
pbol
CO
Q^c
nban
tl noo do
IOD

tfc chl quo trlnh din
truyin
btf5ng
tfio*
Rb§n
tl
noy
cd thl lo tl
boo
trun^:;
gion
do
nb^c
t5l 5 vrStim IPSP xuau hlgn
trong
trtfStog byj>
nay lo
li^i^P
bal
xlnop, tronr do xlnap 5c chl
gltfa
tl bao
trunc iilon
vo tl
bao tbap da co toe
dynt ban
chl sy
Ion truy!n btfng pbIn
Qgay
trong te boo

thap*
IrSn
coc vung vd nao cd
l|p,
boon toon
lopl trtf opl
dtfSn^
ll§n bf btf5n0
uSo*
Si"^
xulc bl^n
kbl klcb thlcb tryc
tllp len
Boo
(59, 60),
Dl5u
do
kbIng dj-nb thoo EPSSP
do
cblnb coc te boo 5c cbl trung glon
tronc
vd noo gay
ro*
Co
i;al
llgu
cho
ran^::
iPbP tarong
vo noo chu ylu do

5c
cbl
qu^t ngtfyc tpo
ro(130)*
Song
uheo Searkov
(55,
56)t thi
gltfo
5c cbl
btf5n=^
tStn
va 5c chl
qu§t ngtf^'c
co
nbimc^
dllo
kbnc
nbou.
t5c
chl
htf^m taa yu:1t
hlfn
ngoy tronr
l*n
klcb
cblch dau*
IFaP
cuo
^1

boo
vun^^
thlnti glac
co
tiial gian
tllo
vam:
thoy
ddl tuy
thuyc
voo
vJ
trl cua klcb thlcb
btf^nf-
t6o» Tb$l gian tleo
ton^:
a
an
nbIt
kbl klcb thlcb vao
tholaous (0,? -
1,3 osec), dal
bcfn
kbl klcb tblch voo hoi
nhon sou cya cu noo sinb tu
(I96 -
2,5 osec) va doo
dyng
l5n
khi

kich
t:ilch
ban^;
§m tbog
(7
- 23
msec),
IPBP glao
bin xuinc khi
klcb tblch
btfSng
too lo
nbimij lopc
klcb tblcb
llSn
tyc,
%tfyc Ipl IPiiP ^
tl boo
vunc
vd noo
v$n dyng
cbl xulc
blyn
kbl klcb thlcb
n^tfyc
cbllu voo bo chop
bloe
nbtfnci;
loat
klcb thlcb

llSn
tpc,
i#oat
klcb chlch co
tSn si
cof^i^
l5n
,
tbl
I?SP cang
tan£;
vo
coni'
keo del*
Tbi^l glen cleo tong
cua
li'SP
tron^
5c cbl
qu|t nguyc
doo
dgng
kbodng
2,6-6,2
©see
trunc
blnb 3, 9 osec
(55)*
Tv^
sy

khoc
blyt
t:dn,
tie
gla dl
din kit lugn
rang 5c
cbl
btf5ng tSo
kbdng pby
thuyc
voo 5c cbl
qu§t
ngtfyc,
Cac
tl boo trung
^lan thoo
gla voo
5c
cbl
qu|t ngijpyc
kbdn^^
a^a
trong
khSu dBn truyln btf5n(r
tlio*
«
25
*
lihtf v4y tronc

vd
olo tSn tpl bol bg
thing tl
bSo^trung
glon 5c chl,
ddo
boo
hoi qu5
trlnh 5c chl
kbic
nbou lo
ac
cbl
htf5npj too
vo 5c chl
qu^t
ngtfyc,
Tac dyuf
5c c
I
l3n nhou
cda
tl
bSo
vd
noo
con
dtxyc
ngblen
c5u quo sy xult

bL^n ^^ vS
l^^
^ tl bio
kbl
klcb
tblch
vSo
coc
dilo rSt nbd
ben
trorig
noo
ban^:
vl
dl§n eye
.
Pbtftfnc
phop noy
L^?1
tit l5 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 osec, klcb tblcb vno coc
dieo 0 Icfp
II,
III,
iv vung vd noo
vyn
dyng
a
oeo do gbl
di^yc
coc
dl^n
tbl
oou xlnop
cuo coc tl
boo
quanb
diem klcb
tblch,
Tr§n
kboon^:
coch
50>

oco tbeo
htfc'ng
nao
ngaog
vo
xucn^:;
sSu phio
dtf5l
dlgn tbl gbl dtfyc
tbuye loyl n>dt
xlnap, con tren khoang
each
100 oco
dl^n
tal cbuyc
logl
do
xlnop*
irong
do
HSP
g§p
nbleu
htfn
ti^^'^
Loal
I?^
oyt xlnop co
ch^l
{-Ion

tllo
tong
O^tj
-
1,4
osec.
Vol pbtrcfng
phop klcb thlcb trong nao ,
Cac tl 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
(29,
46,

9^.,
133), Phon
clcb ttf<?nc
quon cheo
bo^t
tlnb
tpo
xun^T:
cuo
hoi ho^c
vol tl boo cung
o§t
luc, coc tac
glo cbo
thIy oah
hn'cna^;,
tfc
ebl gltfo cac tl
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*
te boo
nbay hcfn
so
vSx
pbtfo^t^;
pbap gbl
nijOoi
tl bao,
Tv^
nhlSn
phtfcfng phop
kich
tblcb trong nao cd
nh(?ic dleo
lo
cdng dign
klcb thlcb tac
dj^ng
kbdni^
cbl
l§n
tb£.n tl bdo, co con co thl len ca nhtfng
syl quanb do,
ida thai
ddl
ph2n
ndo dnh
htfqrng
co

tbyc
cua
qua trlnh 5c chl trong nao,
Pht^jfng
pbap
dtfa
cac
chit
vao tl bao
bSng
vl dlgn dl la
pbtf(A3g
pbap
thuyn lyl bc^n
cd*
V5l oyt
Itfyai;:
nbd coc
chit
,
di^
quo vl dlgn
eye
tryc
tlop voo
te bdo do cho phep
quon