V i e t n a m J o u m a l of Physiology 18(1), 5 / 2 0 1 4 ISSN: 1 8 5 9 - 2 3 7 6

CO C H t P H A N Tfp VA XU HU'DNG NGHIEN CLTU V ^ HOC TAP

V A T R l N H ^ r HIEN NAY
TS. BS. Le Oinh TCing^-^
^Chu nhi$m Bg mon Sinh ly hgc, Truvng Oai hgc Y Ha N6i
'Tong thv ky Hgi Sinh ly hgc Vi^t Nam

Tdm tat: Td nhd dwge djnh nghTa nhw la qua trinh ma hda (encoding), eit glw
(storing) vi tii hign (retrieval) thdng tin. Ba qui trinh nay Hen quan den 3 hg thing nhd
(memory system) khie nhau. Tri nhd eim giic (sensory memory), tri nhd ngin han
(shod-term memory) trong do ein Iwu y khai ni$m tn nhd lam vigc (working memory) vi
td nhd dii hgn (long-term memory). Hgc tgp vi tn nhd la hai khai niem cd lien quan chit
ehe. Niu nhw hge tip la sw finh hdi dwge (acquiring) kiin thdc va ky nang thi td nhd diin
ti n0i dung mi chdng ta finh hdi dwge.
Phin tich ea sd glai phiu va ea che phan td cOa hge tip va tri nhd la m^t trong
nhung thinh cdng Idn nhit eua khoa hgc than kinh hien dgi. Ba mwal nam trwdc diy,
ehung ta chi hiiu biit rit it vi tri nhd, nhwng nay ehung ta da hiiu biit khi rd vi nd.
Trong bii bio cao nay chung tdi se di cap din bin ndi dung chinh Hin quan din hgc tgp
va td nhd. Trwdc hit, chung tdi di cap din phin loai tri nhd theo quan dtem khoa hge
than kinh hign dgi Tiip din chung tdi se di cap den cac vung eiu true eua bg nao lien
quan din hgc t0p va tri nhd. Tri nhd dwge Itru trw vio bd nao cua chimg ta cd tuang tt/
nhw Iwu trw vao chip nhd cua miy tinh ci nhin (PC) hay khdng? Iwu trir toin bd tri nhd
vio cOng m0t vung cau true nao hay phin bi va cit giir a cac vung nio khic nhau? Tri
nhd dwge sw dgng nhw thi nio? Nhu-ng thay doi nao dien ra trong hi thing than kinh
trong qui trinh hlnh thanh tri nhd? Liiu cd gen hay protein dac biit nio liin quan den tn
nhd hay khdng? Lam thi nao di td nhd cd the duy tri dwac suit ca quang ddi? se dwge
trtnh biy d phin tiip theo. Cuii cung, mgt vin di hi trgng, va d$c bigt li khi ehiing ta
gii: lim thi nio di duy tri va cil thign dwge td nhd, lam thi nao cd thi "vi" Igi dwge tri
nhd da biphin manh?

I. D j n h nghta v d phan loai t r i n h a
Nhin Chung tri nhd la he t h i n g gom cdc
qud trinh ma hda (encoding), Iuu giu
(storing) vd ldi hidn (retrieving) thdng tin.
Ba qud trinh ndy tuang tac bdi cac hd
t h i n g nhd khdc nhau. Tri nhd dugc cho Id
b i t d i u bang qua trinh md hda hogc
chuyin d l i Ihdng tin Ihdnh dgng cd I h l e l l
giu d u g c d nao. Qua Irinh ma hda cung cd
I h l dugc h i l u Id ghi thdng lin vdo bp nhd
(registering information in memory). Cac hg
t h i n g trf nhd lidn quan d i n md hda hay ghi
thdng tin vdo vdo bp n h d Id tri nhd cam
gidc {sensory memory) vd Irf nhd ngan hgn
(shortterm memory).Tri nhd ngdn hgn, Iri
nhd Idm vide (working memory) va In nhd
ddl hgn (longtenn memory) ddng vai trd
quan trpng doi vdi hogi ddng hpc tdp,
gido dye - dao tao.
1. Tri nhd cam giac (sensory memory)
Thdng tin vao bp nao Ihdng qua cdc
ca quan cdm giac dugc giu Igi Irong mpl
khodng thdi gian rat n g i n trong giai dogn
ghi nhan cam gidc (sensory register). Ghi
nhdn cam giac lien quan d i n 5 gidc quan
Id thj gidc, thfnh giac, xuc gidc, khuu gidc
vd vj giac. Tuy nhien, Ihdng lin cam giac
dugc quan ldm chu y, d l cgp den nhilu
n h i t Irong cac cdng trinh nghien cuu id
ghi nhdn Ihj gidc va ghi nhgn Ihfnh giac.
Thdng tin thj giac hay "hinh anh" cd the
dugc xem nhu bdn sao chinh xdc nhiing
thd Chung ta nhin I h l y vd m d d i n di Iheo
thdi gian (Pashler and Carrier, 1996).
 Vietnam Joumal of P h y s i o l o g y 18(1), 5 / 2 0 1 4
Thong lin thj gidc d u g c luu giO' d dgng Iri
nhd thi giac ton tgi trong khoang thdi gian
vd cung ngan c d vai Iram miii gidy. Ghi
nhdn thfnh gidc cd the Iuu g i u thdng tin dai
han, c d l u 1 d i n 2 giay vd khdc vdt ghi
nhan thj gidc, Ihdng lin ghi nhdn thinh gidc
khdng m d dan di nhu hinh anh ma trong
s u l l khoang thai gian Ion tai r l i ngdn l y
npi dung thdng lin thinh gidc van d u g c bao
tdn nguydn ven.
2. Tri nhv ngan han (short-term memory)
Hdu h i t thdng tin trang ghi nhan cam
gidc khdng d u g c x u ly sdu (khdng d u g c
chu y den). Vi vdy, hoai ddng chd y d u g c
cho la cdng cu d i l u khien de luong thdng
tin di t i i ghi nhdn cam giac sang tri n h d
ngdn hgn (Gaddes & Edgell. 1994). Thdng
tin Irong In nhd ngdn han d u g c luu g i i i ldu
n h l i cung chf d i n khi chung d u g c lyc Igi
(rehearsed), va nguyen nhdn chd y l u lam
mdt trf nhd n g i n hgn chinh la Ihdng tin cu
bj thay t h l bang thdng tin vua mdi d u g c
chu y d i n (Pashler & Carrier, 1996). He
t h i n g In nhd ngdn hgn nhin chung d u g c
xem nhu Id nai luu giu Ihdng lin tam Ihdi,
keo ddi khodng 30 gidy d i n 2 phdt. L u g n g
thdng lin d u g c luu giu Irong Iri n h d n g i n
han khd hgn c h l . Chf cd the luu giii d u g c
7+/-2 dogn (chunk) hoac bii Ihdng tin khdng
lien quan vdi nhau vd Ihudng bang s l cac
con s l cda cdc so dien thoai. Oe cd Ihe
nhd nhieu han 7+/-2 con so thl cdc sd
trong day s l phai d u g c ghdp nhdm vdi
nhau sao cho mpl vai con so lao thdnh mol
nhdm cd t h l ITnh hgi d u g c (conceptual
chunk). Vf dy: so 6, 3, 9 ghep Ihdnh nhdm
639. Vdi each ghep nhu vgy, khi 1 day
n h i l u con so d u g c dua ra chiing la cd t h l
nhd lot han. De luu g i u ldu thdng tin Irong
In' nhd ngan hgn, can phai lien luc vd chu
dpng s u dung Ihdng lin Irong tri n h d ngan
han.Tri nhd ngan hgn d u g c chia thanh ft
nhat 2 nhdm (Pashler & Carrier, 1996). Hai
nhdm tri nhd ngdn hgn: trf nhd ngdn hgn
Ihj giac vd Iri nhd ngan han thfnh giac (Idi
ndi hogc am thanh) d u g c nghien cuu nhieu
ISSN: 1 8 5 9 - 2 3 7 6
n h i t (Baddeley, 1996; Vallar & Papagno,
1995). Theo do, trf n h d Ihi gidc Id loai Iri
n h d v l nhu-ng gi chung ta nhin I h l y va Iri
n h d thinh giac la tri n h d ve nhung dieu
chung ta nghe t h i y . Nhigm vy d trudng
hgc ddi hdi n h i l u tri n h d ngdn han, dac
biet Irf n h d ngdn han Idi ndi hoac am
Ihanh. Hpc sinh phai n h d ndi dung giao
vien hudng d i n , nhiing ndi dung ma
Chung nghe d u g c li> bdi giang, thao ludn,
hgi thoai vd nhung npi dung chung dpc
d u g c l u sdch giao khoa, Iren bang hay
man c h i l u . T l l ca Ihdng tin lijc dau di vdo
Iri nhd ngan han sau d d d u g c chuyen
Ihdnh tri nhd ddi han.
3. Tri nhd lam viec (working memory)
K i t qua nghidn c d u ve Irf nhd
(Baddeley, 1996, 1998; Vallar &
Papagno, 1995) chia tri n h d ngdn hgn
cung n h u tri n h d lam vigc thanh 2 nhdm;
(1) Iri n h d Idi ndi vd (2) Irf n h d thj giac.
Tri nhd Idm vide chiia d y n g mpl bd phdn
kiem soat chfnh (main controller) hay bp
phdn x u ly trung tdm (central executive)
giup phien giai Ihdng lin mdi d u g c t i l p
nhdn vd tfch hgp thdng tin ndy vdi nhO'ng
thdng tin von da d u g c luu g i i i Irong Iri
n h d ddi han (Baddeley, 1996). Vi vdy. In
nhd lam viec n h u la nhd lal cdi ma chijng
la dang thyc hign trong khi lam viec. Trf
nhd lam viec can cho rat n h i l u nhiem vy
d trudng hgc vd Irong cupc song, chang
hgn de h i l u d u g c ndi dung doan van
phai n h d d u g c npi dung d d i u doan van
vdo thdi d i l m dpc den doan k i t ; Ghi
chep khi nghe giang phdi n h d dugc
nhdng ndi dung ma gidng vidn da vd
dang ndi Irong khi v i i l ; n h d cac hudng
dan khi t h y c hien cdng vigc, dac bigt vdi
thao lac d u g c t h y c hign qua nhieu budc;
tim k i l m mdt t u trong t d dien phdi n h d I d
trong khi lim k i l m ; n h d cdu hdi trong khi
lim kiem cdu Ira ldl. Khi thdo ludn ve m | t
chd d l nao dd, chdng la phdi n h d nhung
gi chijng ta ndi. Khi v l l l phan trd ldl cho
mpl cdu hdi bai ludn vdo bdi kiem tra,
 V i e t n a m J o u m a l of Physiology 18(1), 5 / 2 0 1 4
phdi n h d n$i dung cdu hdi de dua ra nhung
thdng tin quan trpng n h i t c h i i khdng phai Id
thdng tin lien quan d i n chu de. Tri nhd Idm
vide cung c i n t h i l t cho nhung nhidm vy d
d p dO cao han chang han khi phan tich,
t i n g hgp, va giai q u y l t v i n d l . Trong khi
gidi quyet v i n d l , chung ta phai n h d Igi v i n
d l l y , dua ra n h i l u gidi phdp va di d i n
q u y l t djnh giai phap nao cd I h l mang Igi
k l l qud mong muon. K l l qud nghien cuu
v l tri nhd cung chf ra khi s u dung tri nhd
Idm vigc cho cdc nhidm vy d u g c Ihyc hien
d miic nhgn Ihdc cao c h i n g han nhu giai
q u y l t v i n d l , lap ludn (reasoning) va dpc
mang lai k i t qud chi'nh xac han so vdt thyc
hign nhidm vy l y bang tri nhd ngdn han
(Kail & Hall, 2001).
4. Tri nhd dii h^n (long-term memory)
Hinh 1. Hd thong Iri nhd (Squire va Knowllon, 1994)
4.1. Phin I09I trl nhv dii h^n (Long-term
Memory Subsystems)
Chung la da tdng nghT rang Iri nhd n g i n
hgn Id m | l hd t h i n g , tri n h d ddi hgn cung la
mdt hd t h i n g . Song, cho d i n nay chung ta
d i u b i l t r i n g Irl nhd ddi hgn d u g c chia
thdnh n h i l u nhdm. Tdm t h i n hgc vd khoa
ISSN: 1 8 5 9 - 2 3 7 6
N l u nhu he t h i n g tri nhd cam gidc va
Irf nhd ngdn hgn lien quan d i n cdc qud
Irinh ma hda va ghi nhd thdng lin vdo bp
nhd thi he t h i n g trf nhd ddi hgn Igi lien
quan ddn cac qua Irinh luu giu va ldi hien
thdng lin. Luu giu Irf nhd ddi hgn d u g c
xem nhu khd on djnh. Mdt Ihdng lin l u Iri
nhd dai hgn dugc dugc ggi Id quen
(forgetting). Vao nhiing nam 50 cua t h l
ky trudc, da cd nhung nghien cuu h l l
sue ly Ihu lap tmng vdo viec xem xet
quen thdng tin dan gian chi Id do Ihdi
gian Irdi qua hay quen la vi nhung d i l u
chung la dang hpc vao thdi d i l m hien tai
anh hudng d i n kha nang nhd Igi Ihdng
lin ma chung la da hpc d thdi d i l m trudc
do (Baddeley, 1996).
hpc than kinh chia hg t h i n g Irf nhd dai
hgn thdnh 2 nhdm chinh: In nhd hign
(explicit memory, lam djch) hay cdn gpi la
Iri nhd Ihual lai d u g c (declarative
memory, lgm djch) vd khdng Ihudt lgi
dugc (nondeclarative memory, tgm dich)
(hinh 1). Hg thong trf nhd cd I h l Ihudt lgi
 V i e t n a m J o u m a l of Physiology 18(1), 5 / 2 0 1 4
la hg thong Iri n h d cd y thuc, p h i bien
n h l i , vd bao gom tri n h d v l t h y c te (fact)
va s y kign (event). Vf du: Ha Npi thu do
ctJa Vigl Nam la mgi Ihyc le. hay s y kign
n h u ky nghf Irudc d Paris. Tri n h d khdng
thugl Igi d u g c cdn gpi laferfn h d an (Implicit
memory, tgm djch), gdm cdc he thong tri
nhd khdng cd y thuc, cung rat quan trpng.
Trf nhd khdng thudt lai d u g c gdm: tri n h d
ve ky nang, thdi quen (habit) nhu di xe dap.
Idi d td, danh golf, tennis, chai piano), tri
nhd m l i (priming memory, tgm djch), hpc
lgp k l l hgp (associative learning) vf du d i l u
kidn hda kinh dien ( d i l u kien hda kieu
Pavlop), vd cudi cung la hgc tap khdng k l l
hgp dan gian, chang hgn quen thudc hda
(habilualion) vd cam giac hda
(sensilizalion) (hlnh 1). Tri nhd cd the thudt
Igi giong nhu " b i l l cai gi" (know what) va tri
nhd khdng Ihudt lai d u g c l y a n h u " b i l t I h l
nao" (know how). Nhdn b i l l dgc diem vd
chdc nang cCia mdi logi tri nhd ddng vai Ird
quan trong trong ting dyng cdc phuang
phap giang dgy vd hpc lap.
4.1.1. Trinh&cd the thuat lai (declarative
memory): Cdn gpi Id Irf nhd cd y thuc hay
Iri nhd hidn (explicit memory), thupc tri n h d
ddi hgn, luu g i u thdng tin v l thyc t l , trai
nghiem ca nhdn. Tri nhd cd the Ihugl lgi
chia Idm hai loai. N l u tri nhd hoi uc
(episodic memory) la Irf n h d ve cdc s y kidn
cd nhdn da trai nghiem, Ihdng lin t y Ihual
cua ban than, nhd s y kien xay ra d dau, khi
ndo; hoat dpng d ban cdu trai nhieu han
bdn c l u phdi (nghien ciiu bang hinh anh)
Ihi trt nhd kien thuc - khai niem (semantic
memory) lai Id tri nhd v l cac thdng tin
chung, giong nhu eudn bach khoa t h u hay
tir dien. Cd the s u dung tri nhd hoi dc de
nhd va hpc tgp khi Iri n h d k i l n thiic - khai
niem m i l .
- Tri nhd hoi dc (Episodic memory) Id
he thong tri n h d luu g i u cdc Ihdng tin ve
cac thdi ky (episodes) hay cdc s y kign
(events) trong cudc ddi cua mdi chung ta,
Chang han n h d v l nhdng viec chting ta dd
ISSN: 1 8 5 9 - 2 3 7 6
I h y c hien trong ky nghf he toi'dc, vj giao
s u md chung ta n g u d n g mp khi cdn di
hgc. Mdt tri n h d h l l l i e la l'jf do chfnh cua
benh vong tri kinh dien (Baddeley, 1996).
Tri nhv kiin thdc-khai niim
(SemanUc memory, tam dfch) la he
Ihong tri n h d luu giu Ihdng lin k i l n thuc
(knowledge) vd khai nidm (concepts).
Giao due pho thdng d u g c cho Id b l sung
vd lam ddy "kho" n h d k i l n thiic, khai
nigm chang han nhu n h d nguyen lac v i l t
hoa, c h i m cdu, Columbus tim ra Chau
My nam nao, hay Kari Marx la ai chfnh la
cac v i dy cy the ve loai Ihdng lin dugc
luu g i u trong he t h i n g tri n h d k i l n thuc,
khai niem.
Nhiem vu Irong Irudng hgc ddi hdi
n h i l u Irf n h d k i l n Ihdc-khai niem. Khdc
bidt giiia cac cd nhdn ndm d kha nang
s u dung hai logi he thong trf n h d nay
(Bors & McLeod, 1996). N h i l u dua tre
gap khd khan d Irudng hgc do Irf nhd
k i l n thuc, khdi niem t u a n g doi kem. S y
khac bidl vd tri nhd k i l n Ihiic-khdi nidm
vd hdi uc giua cdc cd t h l Id can c i i d l
nhiing ngudi lam cdng lac s u phgm dp
dung nhieu loai hinh hoat ddng dgy hgc
khac nhau chdng hgn n h u cung lam de
dn, lam viec nhdm, da ngogi. Vide dp
dung n h i l u hinh thuc day hpc giup cho
vide tao ra cdc hoi uc hay boi canh t i i dd
cd the dua cac Ihdng tin k i l n thiic-khai
niem vdo bp nhd.
4.1.2. Tri nhd khong the thu$t lai
(nondeclarative memory) hay tri nhd
khong y thdc, tri nhv tiem an (implicit
memory): Thudc vi tri nhv dai han,
gom ky nang, thoi quen, diiu kiin hoa
kinh diin.
Tri nhv phuvng thuv (procedural
memory) la hd t h i n g tri n h d luu g i i i cdc
ky nang t u finh h0i van dpng
(perceptualmolor). Ky ndng vgn dOng
nhu Idi xe hai, dgp xe dgp, ddnh golf.
Trong khi Iri n h d k i l n thuc-khdi nidm
d u g c cho la luu g i d k i l n Ihiic nd la cdi gi
 V i e t n a m J o u m a l of Physiology 18(1), 5 / 2 0 1 4
"what", thi hpc tap phuang thiic d u g c cho
Id luu g i u cdc kien thdc Idm t h l ndo "how"
(Baddely 1996).
- Tri n h d m i l (priming memory): nam 1992
Squire de cdp den mgt hd t h i n g tri nhd
thudc nhdm tri n h d khdng Ihuat Igi d u g c
ggi id tri n h d moi. Tri nhd moi thugc loai tri
nhv can thong tin "moi" de nhv 1^1 d
nhung thdi diem sau dd. Tri nhd m l i
d u g c d l cdp d i n n h u la khi mdt vdi d u g c
tri gidc (perceived) hay x i i ly Irudc dd thi
de ddng d u g c Iri gidc hay x u ly khi ggp d
lan sau (Baddeley, 1996). Tri nhd moi cd
lidn quan vdi mdi Irudng gido dye dgc biet
Id vigc to chiic trudc khi l i l p can k i l n Ihuc
mdi.
T y u chung igi, Iri nhd dai han bao gom
hai hg t h i n g trf nhd cd I h l thual lai vd Iri
nhd khdng I h l thudt Igi. Ddc diem vd chuc
ndng cda m l i loai tri nhd ddng vai Ird quan
Irpng trong dng dung cdc phuang phdp
gidng dgy vd hpc tap.
4.3. Til hi^n tri nhd dii han (Long-term
Memory Retrieval)
Tdi hign d u g c d l cgp d i n nhu truy x u l l
Ihdng tin d u g c luu giu trong bd nhd.Cdch
thiic ma hda vd luu gid thdng tin se quylt
djnh vigc truy xuat vd ldi hidn Ihdng lin.
Han nua, Ihdng tin va s y kign dugc luu giu'
Id nhu-ng Ihdng tin mdi Ihudng d u g c d l
cdp d i n nhu Ihdng tin ggi (cues), dnh
hudng d i n s y de dang khi truy x u l t Ihdng
lin (Roediger and Guynn, 1996). Hidn nay,
nhieu nhd tdm t h i n hpc lin tudng rang qua
trinh tdi hidn ddng vai Ird quan trgng Irong
vide q u y l t dinh thdng tin d u g c nhd i l l hay
khdng (Roediger and Guynn, 1996). Mpt
vdi nghidn cdu cho t h i y thdng tin d l ddng
ldl hidn t i i Iri n h d ddi hgn n l u nhu chiing
dugc luu giO d u d i dgng Ihdng lin ggi nhd
(retrieval cue) vd thdng tin nay d u g c s u
dyng vdo Ihdi d i l m dugc nhd Igi (Roediger
and Guynn, 1996). Nghidn ciiu cda
Roediger vd Guynn ndm 1996 neu bdt s y
khdc bi$t giOa luu giu vd ldi hign tri nhd dai
hgn. Khi d i l u kidn d n cho luu giu Ihdng lin
ISSN: 1 8 5 9 - 2 3 7 6
Id n h u nhau, cac "ggi y nhd" lam tang
gap doi so t i i cd I h l nhd dugc. O'ng
dyng cdc "gdi y nhd" cho cdc nhd gido
due dd la cac thdng lin mdi nen dugc
trinh bay bang nhilu °ggi y nhd" d l Idm
tang kha ndng tdi hign thdng lin. Cdc "ggi
y nhd" cd ttil d d u d i nhilu dgng hogt
ddng da dugc Ihao lugn Irudc day,
Chang han hogt hda (khdi dgng) Irudc
k i l n Ihuc lien quan, ddl thdng tin vdo
nhdm nhdn thuc (conceptual categories),
tgo cdc thdng lin lien quan ca nhan,
va/hoac dua thdng lin vao bdi cdnh kinh
nghidm. Vi vay, khdng chi ma hda vd luu
gid Ihdng tin ddng vai Ird quan Irgng, md
chinh cdc "ggi y nhd" cung ddng vai trd
quan Irpng lam i l l u u hda kha nang tai
hien cac Ihdng tin da hpc. Theo Baddeley
(1996), nhgn thuc thudng giup vigc ldi
hign Ihdng lin chinh xdc han Id nhd Igi.
Bdi vi l u hay thdng lin dugc hpc Ihyc s y
Id cdc "ggi y nhd", Ihdc day hogc ldng
cudng kha nang Iruy xuat den cdc d i u
v l l nhd. Vi du, mpt sinh vien dugc dgi
cau hdi, "Vung ndo cua nao k i l m sodi thj
giac?", d day dan t h u l n chf Id nhidm vy
nhd Igi. N l u "ngdn hdng" tti ngu dugc
cung d p va Irong danh myc t u cd l u
"IhCiy c h i m " , t i i ndy ddng vat trd nhu "ggi
y nhd" cd ve nhu Ihuc day vigc Iruy x u l t
t u cac v l l tich nhd (memory trace) chda
dyng Ihdng lin, "vdng nao k i l m sodt thj
gidc Id Ihdy cham". Vi vay, s y khdc bigt
giua Ihyc hign nhigm vy nhgn Ihiic vd
nhd cd I h l d u g c s u dyng giup nhgn ra
lidu khoanh khac m i l nhd dgc bidl ndo
do la do qud Irinh luu giu thdng lin bj gdy
hay khd nang lai hidn Ihdng lin I h i t bgi, vl
kha nang lai hign thdng tin thudng m i l di
khi s u dyng cac test ddnh gia nhdn Ihuc
(Baddeley, 1996). Han the nua, nghidn
cdu cOng chf ra d l cdc "ggi y nhd" ddng
vai trd hdu dyng trong vigc Ihuc d i y nhd
Igi Ihdng tin Ihl chdng phdi d u g c x u l t
hidn trong qud trinh hpc.
 V i e t n a m J o u m a l o f P h y s i o l o g y 18(1), 5 / 2 0 1 4
5. Phin loai va each tiep can khac vi tri
nhv
Dya tren md hinh m i i c do x i i ly' thdng
tin (levels of processing model) Cralk vd
Lockharl chia ra 3 muc: (1) m u c luu giO'
Ihdng tin phy thudc m i i c do sau cua thdng
lin d u g c x u ly. (2) m i i c ndng (y thuc d u g c
thdng tin cam giac); (3) m i i c sau (Ihidt lap
lien he, lien k i t thdng tin, gan y nghTa cho
cac Ihdng tin cam gidc I n t u g n g , gan k i t
thdng tin cu vdi Ihdng lin mdi). S d dung
test cda Tulving va Craik d l danh gid cac
m i i c do nay.
Nghidn cdu v l tri n h d t i l p d n t i i ban
c h i t cua nhd chia ra cdc hinh Ihdc n h u tri
nhd bdng den (flash bulb memory), Ihdng
lin nhd vyl xudt hien rdi vut b i l n mat; tri
nhd hinh anh (eidelic memory): kha ndng
luu g i i i hinh anh sau khi kfch thfch b i l n
mat, 5% Ire em cd tri nhd hlnh hpa tot, vd
hau h i t cd tri nhd dai han l i t han Ire khdc.
Trf nhd vd van hda (Sir Frederick Bartlett),
II. D j n h khu t r i nhd-
Tri nhd d u g c d t giii d ddu? Hien lgi cd
ba cdch t i l p cdn c a bdn de nghien ciiu d u
trdc t h i n kinh lien quan d i n Irf nhd vd hgc
tap: (1) Ky thual ghi hinh hidn dgi chang
hgn n h u fMRI (functional magnetic
Hinh 2. Hlnh anh chyp cat ldp nao benh nhdn H.M sau khi phau thual lay bd hoi hai ma
(trai) vd hinh anh chyp d t Idp nao ngudi binh t h u d n g (phai). (dnh chyp cua Suzann
Corkin. 1997)
ISSN: 1 8 5 9 - 2 3 7 6
resonance imaging) hay PET (posilran
emission tomography) cho phep nhin
thay cac vdng nao hoat ddng khi nao
t h y c hidn nhiem vy chuyen biet. N l u mpt
ngudi d u g c dat trong may quel fMRI khi
t h y c hidn test Iri n h d cd t h l xdc djnh
d u g c vdng nao ndo hoat dgng Iren c a s d
dd cd t h l xac djnh d u g c vung nao lien
quan d i n x u ly va cat giu- tri nhd. (2) Gay
t i n t h u a n g nao t h y c nghiem, vdi phuang
phap nay tdng phdn nhd cua nao chudt
nhat hay chupt cong d u g c logi bd hay
gay bat hoat bang hda chat, chudt thyc
nghiem d u g c kiem tra mpt each he t h i n g
de danh gid xem cdc ton t h u a n g nay cd
anh h u d n g d i n he Ihdng Iri n h d ndo
khdng. (3) Nghien c u u t u cac trudng hgp
ton t h u a n g va benh iy nao: bang phuang
phap nay cdc nha khoa hpc tdn dyng Igi
t h l I d cdc bdnh nhan khdng may mdn bi
ton t h u a n g nao, v i dy do ddl quy ndo,
hay u nao d nhung vung nao dac bigl,
n l u k h i l m khuyet tri n h d chung td vdng
nao bi ton t h u a n g cd lien quan d i n loai
tri n h d bi m i l .
 V i e t n a m J o u m a l of Physiology 18(1). 5 / 2 0 1 4
Nghidn a i u kinh d i l n djnh khu tri nhd
chfnh la k i t qud dieu til bang p h l u thugt
cho bgnh nhdn Henry Molaison. ngudi
bgnh nay I h u d n g d u g c cac nhd khoa hpc
d l d p den d u d i ldn gpi H.M cho den khi
qua ddi vdo nam 2008. H.M n l i l i i n g Irong
cdc y van khoa hpc t h i n kinh bdi vi t d
chfnh nao bp cua bdnh nhdn ndy cac nhd
khoa hpc dd xdc dinh d u g c d u Irdc ndo
lidn quan d i n tri nhd. Vdo nhung nam
1950, H.M d u g c chan dodn m i c chung
dpng kinh khd kiem sodi, trong khi mgt so
Irudng hgp cd t h l k i l m sodi bang Ihuoc.
thl mdt so trudng hgp can phdi dieu Iri
bang phau thudt d l lpai bd nhung phan
nao gdy ra ddng kinh. Bdnh nhdn H.M
dugc p h i u thudt lay bd hoi hdi ma cd hai
bdn (hinh 2). T r u d c p h i u Ihudl. bdnh nhdn
H.M cd Iri nhd lot, sau p h l u Ihugt H.M
k h i l m khuyet tri nhd rat ngng n l . Khd ndng
hinh thdnh Irf nhd mdi ve thyc l l (facts) vd
s y kign (events) bj suy gidm nghidm trpng,
ggp khd khan khi hgc nhung I d vyng mdi;
khdng nhd dugc nhiing gi xdy ra cda ngay
hdm Irudc. Vi vdy, neu hdi H.M v l cudc
phdng v i n trudc dd, benh nhdn H.M se
nhd r i t ft hogc khdng nhd gi v l cupc
phdng van hay cdc s y kign xdy ra Irong dd.
Bgnh nhan H.M mac dd ggp khd khdn Irong
vigc hinh ttidnh Irf n h d mdi nhung van iuu
giu tri nhd cu v l cdc s y kign vd thyc t l . Cy
the, bgnh nhan ndy v i n cdn luu giu lodn bp
Iri nhd v l thdi nidn I h i l u , t i t ca Iri nhd cd
dugc Irudc khi t i l n hdnh phdu thual. Bieu
hidn nhu I h l gpi Id qudn Ihudn c h i l u
(anterograde amnesia). Nhu vgy, k i t qud
nghien cuu t u bdnh nhdn H.M cho thay
vung hdi ma khdng phai Id nai luu giu trf
nhd cu. Cho d i n nay, chdng la cung b i l l
r i n g tri nhd cu d u g c luu giu d nhung vCing
khdc cda nao. c h i n g hgn nhu vd ndo thdy
trdn. Qud trinh c h u y i n doi Iri nhd t u dgng
khdng viing chac (labile) Ihdnh dgng vung
c h i c han (more enduring) dugc gpi Id "qud
Irinh cung c6" (consolidation). Qud trinh
ndy d i l n ra d mdt vdng khdc cda ndo chu
ISSN: 1 8 5 9 - 2 3 7 6
khdng phai d vung tri n h d dirge ma hda
luc dau. Benh nhan H.M Id mdi Irudng
hgp Ihu vj, mac du suy gidm nghidm
trpng khd nang hinh Ihanh Iri nhd mdi v l
s y kien vd Ihyc l l , nhung lal cd the hinh
Ihdnh d u g c Iri nhd mdi ve ky nang vd
Ihdi quen vd khdng he y Ihdc dugc minh
cd ky nang dd, khdng Ihual Igi du-gc. K l l
qua nghien CUTJ ndy chdng td Irl nhd ky
ndng vd thdi quen khdng dugc hinh Ihdnh
d hdi hai ma. T i n g hgp k i t qua nghien
cuu t u benh nhdn H.M vd nhung bgnh
nhan khac cd t h l k i t lugn tri nhd dugc
phan bd d hau khap cac khu vyc trong hg
than kinh, va cdc vung nao khdc nhau lien
quan d i n cac logi bi nhd khdc nhau.
Hinh 3 tong hgp k i t qud nghidn ciiu
v l cdc d u true vd vi Irf gidi phau lien
quan d i n cac he t h i n g trf nhd trong
nhilu Ihdp kJ khang djnh h l i thdi duang
giua va cac d u trdc hoi hdi ma lien quan
den Iri nhd s y kign vd thyc l l ; T h l vdn
lien quan d i n Iri nhd ky ndng vd thdi
quen; vd ndo mdi (neocortex) lidn quan
den Iri nhd moi; hanh nhdn lidn quan d i n
Iri nhd cam xuc (emotional memories); vd
l i l u ndo lidn quan d i n cdc dgng Ihuc dan
gidn cua hpc lgp lien hgp. Cdc vung ndo
I h i p han vd tuy song thgm chi chua
dyng nhu'ng dgng dan gidn han niia v l
hpc tgp. Tdm lai, Iri nhd khdng luu Iru d
mpl vdng ndo duy n h l i md dugc phan b l
d cac vdng nao khac nhau.
III. C a c h l te bdo va phan t u cua tri
nhd*
Hau h l l hieu biet ve ca c h l than kinh,
ca c h l phan tu cda iri nhd va hgc tgp cd
dugc bang each t i l p can nghien ctiu l u
hg t h i n g md hinh (model system). Vdi hg
Ihong nay, chung ta d l dang kiem chdng
dugc d miic te bao. Sen bien Aplysia
ealifomiea la smh vgl cd hg thong md
hinh (hinh 4) d u g c dng dyng nghidn cuu
n h i l u . Sinh vat nay dugc lim Ihay d cdc
vung dpc theo bd bien Nam
California, dai khoang 15 cm (6 inches)
 Vietnam Journal of Physiology 18(1), 5/2014 ISSN: 1859-2376

va nang 150 g. Cac nha khoa hpc Ihan kinh
da sdm nhdn ra cau Irdc cda Aplysia cd
nhilu Igi thl v l mat ky thual d l lim hieu ca
chl nln tang va ca che phdn tu- cda tri
nhd. Qua Ihyc, nhung kham pha md
dudng tu cac nghidn cd-u cua Eric
Kandel khi tiln hdnh thyc nghigm d lodi
ddng vai nay da dugc khang djnh bang
giai Nobel cho Sinh ly hgc va Y hpc vdo
nam 2000.

Hinh 3. He thong tri nhd va cac cau trtJc nao hen quan (ve lai tti mo hinh ctJa Squire va
Knowlton, 1994)

Hinh 4. Nghien cuu cam gidc hda ngan hgn Iren Aplysia (Eric Kandel, 2000).
Thyc nghiem tren Aplysia cd 3 lgi dang dan gian cua hgc ldp khdng lhu$l
Ihl: Trudc hll, dgng vdt nay cd day dd cdc lgi dugc: (1) dieu kign hda kinh diln cda
 V i e t n a m J o u m a l of Physiology 18(1), 5 / 2 0 1 4
Paplov; d i l u kign hda hdnh ddng (operant
conditioning) va cam gidc hda. (2) Aplysia
cd hd t h i n g I h i n kinh r i t dan gian. N l u so
vdi bg nao ngudi cd 100 IJ^ naron Ihi todn
bp hg t h i n g than kinh cda lodi dpng vdt ndy
chf cd 10.000 t l bao. Cac t l bdo nay phdn
b l d hgch t h i n kinh (hinh 5). Mdi hgch cd
khoang 2.000 te bdo, Idm Irung gian hodc
kiem seal mpt so hanh vi khac nhau. T u c
Id, moi hdnh vi cd t h l d u g c k i l m sodi chi
Hinh 5. Aplisya cariifornia va naron cua chdng
Hinh 5 cho thay hgch than kinh duac
ldch rdi quan sal d u d i kinh h i l n vi. Hach
cd dudng kinh c d 2 mm Cac d u tnic hinh
d u phd lodn bd hgch chfnh Id thdn cua cdc
t l bdo t h i n kinh. Naron cd I h l quan sal
dugc dudi kinh hien vl, d u g c phdn b l va
hogi ddng chdc nang ridng re. cd t h l lach
naron ra khdi hach va nudi d y song dugc
Irong r l i nhieu ngay. Tren thyc l l , da cd
nhilu nghidn cuu cho I h l y cdc naron ldch
ra khdi hgch than kinh, nudi trong mdi
Irudng Ihfch hgp da lai lgp Igi cdc k i t n l i
synap, cung d p mOl hd thong ihyc nghigm
vd cdng hiiu dyng d l nghidn cuu cac dgc
d i l m sinh ly hgc l l bdo I h i n kinh vd cdc
I S S N : 1859 - 2 3 7 6
bdi 100 naron hoac thdm chi il han. Cd
the ldch ra dugc mpl mach naron hoan
chfnh lam n l n ldng cho mpt hdnh vi, vd
sau do, qua qud trinh huan Iuygn d u g c
Ihyc hidn bdi mach naron ndy, k i l m tra
nhung thay doi Irong mgch de Hm ra c a
c h l tri nhd va hpc ldp. (3) hgch t h i n kinh
d u g c lao nen t u nhung naron cd kich
thudc rat Idn.
dgc Irung k i t noi cda chiing. Hlnh 5C
dua ra mot vi dy minh hpa l l bao than
kinh cam giac (TB nhd ndm ben phai) vd
t l bdo than kinh van dpng (TB Idn nam
ben Irdi) trong mdi Irudng nudt d y . Dudi
kinh h i l n vi didn tti cd I h l nhin t h i y vi
dign c y c dugc dat vao naron cam gidc
vd naron van dgng de Ihuc hien phep ghi
dien the ngi bdo.
1. C a c h l te bao cua cam giac hda -
mpt dang d a n gidn cua logi trl n h d
khong thudt lai d u g c
Thyc nghigm cam gidc hda d u g c t i l n
hdnh nhu sau Con vgl (Aplysia) b| kich
thich didn hogc co hgc y l u vdo
 V i e t n a m J o u m a l of P h y s i o l o g y 18(1), 5 / 2 0 1 4
dudi. Kfch Ihich nay gay ra mdt phan xg
bao vg co rut c a Ihe bao gdm co ca dudi va
cdc phdn quanh Ihdn n h u mang va
siphon. Sau m l i 5 phut mdi kfch thich d u g c
tgo ra, thdi gian cua moi kich Ihfch n h u
nhau, con vdt dap ting vdi kfch Ihich bang
phan xa rdt lui khd rd rang. Nhung n l u mpl
kfch thich mgnh (shock didn) vao phdn
thdn, sau dd kich Ihich vao phdn dudi thl
thay dap ung manh len ro ret. Tang dap
ung d u g c xac djnh la k l l qua cua kich
Ihich manh vao thdn. K i t lugn, con vat hpc
d u g c ddy Id kich thich "nguy h i l m " . Hien
tugng nay ggi la cam giac hda, hinh thiic
hpc lap cd d h l u h i t cdc ddng vgl k l ca
ngudi.
Mach nv ron va cv che cua cam giac
hda
B i n g each ghi dign I h l Id- ben trong
naron cd kich thudc Idn cda Aplysia cd the
bdc Id mgch naron lam n l n lang cho hidn
tugng cam giac hda. Kich Ihich da lam
hogt hda cdc naron cam gidc (SN), glai
phdng glutamate vdo khe synap vdi naron
vdn ddng (MN). N l u long kfch Ihfch l u cac
synap Iren naron van dgng du Idn. naron
van dgng se d u g c hogt hda. sau dd cac
didn the hoat dgng d u g c tao ra se lan
Iruyen den cdc hach gay co ca. Cac n a ron
van dpng cdng hung phan thi phan xa dap
dng cang mgnh. Phan xg nay Irdn Aplysia
l u a n g l y n h u phan xa duoi gdi hay phan xa
cdng c a d u g c Ihyc hidn thdng qua cdc
mgch naron l u a n g t y d luy s i n g .
Ca che phin ti> cua cam giac hda: Kfch
thfch cam giac dan den giai phdng c h i t
truyin dai I h i n kinh serotonin (5-HT) (hinh
4). 5-HT lam Ihay doi dan truyen synap
giua naron cam giac vdi naron vgn
dpng. T r u d c khi qud Irinh hpc tap dien ra.
mpl dien the hoat ddng l u naron cam giac
tao ra mpt dign the kich Ihich nhd sau
synap (EPSP) d naron van dpng. Nhung,
sau mdt kich thich gay cam gidc hda
(shock dien), mpl dien t h l hoat dOng d
naron cam gidc tao ra mpt dign the Idn sau
ISSN: 1 8 5 9 - 2 3 7 6
synap a naron vgn ddng. Oidn the sau
synap Idn d naron van dgng lam tang
kha nang hoat hda naron van dpng gdy
ra mpl co bdp c a mgnh han (cam gidc
hda). Nguyen ly cda hgc lap va Iri nhd
ndy cdn nguyen gid trj d l i vdi hpc tap vd
tri n h d trong bd nao cua ngudi, dd la hpc
lap lien quan vdi nhiing thay doi d i n
t r u y i n synap giu-a cdc naron. C a c h l tri
nhd vd hpc lap d day khdng phai tai s i p
xep lai he t h i n g Ihdn kinh, hay phai trien
cac naron mdi ma la thay doi ddn I r u y i n
synap ton tgi t r u d c dd. Vay c a c h l sinh
hda nen lang cho hgc tap vd tri nhd la
gi?
Cv che ngin han cOa cam giac hda:
C a che cua tri n h d ngan hgn cho cam
giac hda d u g c minh hpa d hinh 4. Kich
thfch cam gidc hda (shock didn) lam giai
phdng 5-HT. Sau do, 5-HT gan vdi hai
loai receptor trdn naron cam giac lam
hoat hda he Ihong DAG/PKC vd hd
thongcAMP/PKA. Protein kinase cd hai
tdc dung: Irudc h l l , chdng d i l u hda cdc
dgc linh kenh ion trdn mang t l bao Id ca
s d cho khdi phai vd ldi c y c cda dign t h l
hogt ddng. K i t qua cho thay, kich thich
gay cam giac hda lam tang tinh kich thich
cua naron cam giac, tang c u d n g dign I h l
hoai ddng d u g c tgo ra khi kfch thfch tren
da, khi dien t h l hoat ddng d u g c tao ra vd
lan den cue tan cung cua naron cam
gidc, kenh Ca m d , Ca di vdo ldn cdc Ign
cdng tang giai phdng c h i t I m y i n dgi than
kinh vdo khe synap giu-a naron cam gidc
vd naron vgn dpng. T h u hai, kinases dieu
hda cdc qud Irinh t l bao lien quan d i n
giai phdng chat Iruyen dai than kinh (tdng
kfch Ihudc cda b l synap san sang gidl
phdng chat Iruyen dgt than kinh, lang ddp
ling vdi ddng Ca dl vdo do dien I h l hogi
dpng). Cuoi cung, 5-HT Idm Ihay doi dgc
trung d u trdc cua cdc naron vgn d | n g
(5-HT Idm tang so lugng receptor t i l p
nhgn glutamate). N h u vgy, (1) hgc ldp
lidn quan d i n hd t h i n g c h i t truyen tin t h d
 Vietnam Joumal of Physiology 18(1), 5/2014
2. Trudng hgp ndy d protein kinase C
(PKC) vd hg t h i n g protein kinase A (PKA)
deu lien quan d i n . K i t qud ndy dd m d ra
xu hudng nghien cuu tien hdnh k i l m
chung vai trd cua he Ihong tmyen tin t h u
hai doi vdi cac he t h i n g trf nhd d ca ddng
vgt cd x u a n g song va ddng vdt khdng
xuang s i n g . (2) Iri nhd lien quan d i n dieu
bien kdnh ion Iren mang naron. C a c h l nay
bao gdm ca cdc kdnh lidn quan Iryc t i l p
d i n giai phdng c h l l Iruyin dgt than kinh
(kdnh Ca^* d cdc naron Irudc synap), cdc
kdnh k i l m sodi tinh kfch thich cua naron
sau s ^ a p , vd cdc kenh lam Imng gian cho
cdc dap ung synap cua naron sau
synap. (3) cAMP li chit truyen tin thw hai
quan trgng liin quan din td nhd. Vdi
nhung bang chting ndy cd t h l m d ra trien
vpng nghien cuu mdi Idm t h l ndo d l tang
Iri nhd. Vi vgy, Irf nhd ngan han "short-
term" cdn gpi la Id tri nhd thodng qua
(transient) va thodng qua Id vi nhung Ihay
d l i sinh hda nen tdng chf d i l n ra Irong
khodng Ihdi gian n g i n . Thdi gian cda Iri
nhd phu thudc vao Ihdi gian bj phosphoryl
hda cda cdc protein ca c h i t (substrate
proteins) (VD:cdc kenh Irdn mdng l l
bdo). PKA se chf hogt hda trong mpl
khoang thdi gian rai ngan sau mpl kfch
Ihich ngdn (brief stimulus) do cAMP sd bj
gidng hda va Idm PKA gidm x u i n g ; Protein
phosphatases Ioai bd cac g i c phosphate ra
khdi protein c a c h l l luu g i u tri nhd Idm cac
protein m i t tdc dyng.
Cv chi dai hgn cua cam giic hda: trf nhd
n g i n hgn vd tri n h d ddi hgn khdc nhau d 2
d i l m ca bdn. (1) Trf nhd ddi han Iten quan
d i n thay doi trong long hgp protein vd d i l u
hda b i l u hidn gen. (2) Irf nhd ddi hgn lidn
quan d i n b i l n dot v l d u trdc. Khi so sdnh
c l u true naron cam gidc cda ddng vdi chua
dugc h u l n luydn vd dOng vdt dd dugc
h u l n luydn cdm gidc hda. K l l qud cho thay
sgi tryc ddy Idn vd cd nhieu nhdnh
nhd. Dpc theo cdc nhdnh sgi Injc cd nhilu
c h i m nhd hogc nhdnh sgi tn^c c h l l Idn
ISSN: 1859-2376
(varicosities). Nhung dogn nhdnh sgi Iryc
chdi Idn nhu vdy chfnh la cac Ign cung
sgi Iryc cua cdc naron cam gidc Igo cdc
k l l n i l mdi vdi cac naron vdn ddng. K i t
qua nghien cuu d u true naron vdn ddng
d con vgl 24 gid sau h u l n luydn cam
giac hda cho t h i y s y khdc bidl Idn v l so
sgi Iryc: naron d ddng vdt dugc huan
luyen cd so sgi Iryc vd cue tan cdng
nhilu han so vdi naron cda nhung ddng
vat chua dugc h u l n luyen. Nhu vdy, Iri
nhd dai hgn lien quan d i n s y bien doi ve
d u tnic bao gdm phat Irien cac nhanh
sgi tryc va hinh thanh synap mdi. Do dd,
nhd mol d i l u gi dd den ngdy mat, tuan
ldi hay nam ldi Id do s y b i l n doi c l u tnic
d synap dang d i l n ra trong nao.
K l l qua t u cdc nghidn ctiu Ihyc
nghiem cung cho I h l y trf nhd dai hgn
dugc cho la lien quan d i n b i l u hign gen,
mpl Irong cdc hudng nghien cuu chfnh
cda cdc nha khoa hpc Ihan kinh Id Hm ra
logi protein va gen ddc hidu lidn quan d i n
trf nhd ddi han. K i t qud dugc minh hpa d
hinh 4 chi ra mpt vdi gen vd protein lien
quan d i n cam giac hda. Luu y, c h i t
truyen lin Ihti hai cAMP lidn quan d i n Irl
nhd ngdn hgn cung lien quan d i n Iri nhd
ddi hgn. Tuy nhidn, ngodi ldc dyng
phosphoryl hda cdc protein kenh tren
mang l l bao, cAMP Ihdng qua PKA gdy
ra lac dung phosphoryl hda y l u l l phidn
md. Cdc yeu lo phien ma (vl dy; CREB),
khi dugc phosphoryl hda cd khd ndng
dieu hda bieu hien gen d i n d i n nhung
thay doi trong long hgp protein giu vai trd
quan Irpng Irong vigc hlnh Ihdnh vd duy
tri nhu'ng bien doi dai hgn ve dan Iruyin
synap, nhd dd ddng vai Ird quan trpng
doi vdi tri nhd dai hgn. Tuy nhien, d n
luu y r i n g , khdng cd mpt gen nhd "than
ky" ndo ca, hay ndi mpt each khdc vide
hinh thanh vd duy tri tri nhd (Ihdm chf
trong m0t naron) lidn quan d i n s y g i n
k i t cda r i t nhilu gen vd protein, chdng
tdc dpng d i n g Ihdi n h l m Ihay d l i cdc
 V i e t n a m J o u r n a l of P h y s i o l o g y 18(1), 5 / 2 0 1 4
dgc tinh cua naron, d i l u hda cac dgc tinh
cua naron va ddn t m y i n synap. Cung d n
luu y s y b i l n doi trong b i l u hien gen khdng
xuat hign ngay lap l u c ma qua n h i l u giai
dogn. Mdt so b i l n doi xuat hien s d m , mpt
s l bien doi xuat hien mupn thdm chi 24 g i d
sau khi qua trinh hpc tgp dien ra.
2. Co- chi te bao va phan tir cua hien
tuvng quen thugc hda
Hinh 6. Mgch naron gian the cda hien l u g n g quen thudc hda va d i l u kien hda kinh dien
(Eric Kandel, 2000)
Hien l u g n g quen thudc hda lan dau tien
dugc d l cap den bdi Ivan Pavlov va
Charies Sherrington.Trong khi nghien c i i u
ve l u t h l vd vdn ddng khu Iru (locomotion),
Shemngton quan sal thdy hidn l u g n g gidm
c u d n g dp ddp iing mpt sd phan xa nhat
djnh, chang han nhu gap chi de ddp u n g
vdi nhung kich Ihich lap Igi. Dap ung phan
xg chf quay lai sau khi nghf mpt vdi giay.
Ong cho r i n g s y suy giam ndy chinh la s y
quen Ihudc hda, va la k i t qua cua vigc m i t
d i n truyen synap tren cdc d u d n g ddn
truyen d i n naron van ddng von da bj hoai
hda lgp di lap lai. Hien l u g n g ndy sau dd da
ISSN: 1 8 5 9 - 2 3 7 6
Quen thudc hda Id mpt trong nhiing
hinh t h u c hpc lap khdng y thdc d a n gian
nhdt, con vdt hpc d u g c dac d i l m cua
nhu'ng kich Ihich vd hai. D i u tien chung
dap ung v d i kich thich m d i bang mgt logt
cac dap i i n g cd tinh djnh h u d n g . N l u
kich thfch ndy la cd fch hay vd hgi con vat
se hpc d u g c vd lan sau neu kfch Ihfch
nay lgp lai thl chdng s§ Id di.
dugc Alden Spencer va Richard
Thompson nghien ctiu d muc t l bdo. Hg
Hm Ihay cd s y song hdnh g i n gui hanh
vi va t l bdo giiia quen thupc hda cdc
phan xa gap d meo va hign t u g n g quen
thupc hda phiic lgp han d ngudi. Ghi
dien t h l npi bdo I d cdc naron vdn ddng
tuy song d meo cho t h i y hien l u g n g
quen thudc hda lam gidm ddn tmyen
synap giua naron lidn hgp kich thich vdi
cac naron vdn dgng. K i t n l i giua naron
cam gidc vdi naron lidn hgp khdng bj dnh
hudng Do lo chiic naron lien hgp d tdy
s i n g cua dpng vdi cd x u a n g s i n g r l i
 V i e t n a m J o u m a l of Physiology 18(1), 5 / 2 0 1 4
phiic lgp vi vdy phan tich sdu c a c h l t l
bdo d l Chung minh s y quen Ihupc hda cdc
phdn xg g i p r i t khd khan. Nd lyc chiing
minh hign l u g n g ndy ddi hdi he t h i n g dan
gidn han. Sen bien Aplysia ealifomiea, la
d | n g vdt cd hd t h i n g t h i n kinh dan gian Id
ra phu hgp d l nghidn ciiu he t h i n g tri nhd
ndy. Aplysia l y vg bdng each co mang vd
siphon, phdn xg ndy l u a n g l y phdn xg gdp
chdn d u g c Spencer va Thompson nghien
cdu. Kich thfch nhg vdo siphon tam co ca
siphon vd mang (gill). Neu kfch thfch lgp di
lgp Igi con vgt se quen. Phan xg nay da
dugc nghidn cdu ky ludng. Mdt kfch thich
mdi Idn siphon Idm naron cam gidc chi p h l i
phan ndy phdi ra dign I h l hogi dpng va
c u l i cdng tgo ra dign I h l kich Ihich sau
synap d naron lidn hgp vd naron van ddng
(hinh 6). Didn t h l d u g c phai ra l i i naron
cdm gidc vd naron lidn hgp dugc ggp lai
theo khdng gian vd thdi gian lam cho naron
vdn ddng bj hung p h i n lien tyc Idm co
mang mgnh. N l u kich thich lgp lai didn t h l
kich Ihfch sau synap do naron cdm giac lao
ra d naron lidn hgp vd naron vgn dpng nhd
di ddng k l . Do dd, kich Ihich lgp Igi Ihl didn
I h l synap do mdi vdi naron lien hgp tao ra
Irdn naron vdn dOng cung yeu han, k l l cyc
Id naron vdn dpng phdt xung yeu d i n cuoi
cdng phdn xg ndy b i l n m i l . Vgy y l u l l nao
Idm gidm higu qud d i n Iruyen synap cua
naron d m gidc? K i t qua phdn tfch gidi
phdng ldi synap cho I h l y gidm d i n truyin
synap Id do gidm so lugng lui synap dugc
gidi phdng ra l u cac Ign cdng sgi Iryc cua
naron cdm gidc (hinh 6). C h l l truyin dgt
I h i n kinh do naron cam gidc gidi phdng Id
glutamate. Glutamate sau khi d u g c gidi
phdng d i n gdn vdi 2 logi receptor Irdn
mdng naron vdn dgng, mgi loai gan g i l n g
W-melhyl-d-aspartale (NMDA) receptor d
ddng vdi cd x u a n g song cdn logi kia Id
non-NMDA receptor. Cdc nghidn cdu cung
chf ra khdng cd s y Ihay d l i ve dp nhgy cua
receptor vdi glutamate Irong hidn lugng
quen thu|c hda. Nguydn nhdn Idm gidm
ISSN: 1 8 5 9 - 2 3 7 6
giai phdng chat truyin dat than kinh cho
d i n nay v i n chua dugc h i l u b i l t day du,
song cd gia t h i l l cho r i n g mot phan la do
giam van chuyin cdc tdi synap vdo vung
van dgng (active zone) cda mdng Irudc
synap. S y suy giam ndy kdo ddi nhilu
phdt. B i l n doi m i m deo v l chuc ndng
d i n I m y i n synap ddng gdp vdo ca c h l l l
bdo cho trf nhd ngdn hgn cua hign tugng
quen thudc hda. S y b i l n doi nay d i i n ra
lgi n h i l u vj trf tren mgch naron cua cung
phan xg, Irf nhd Irong Irudng hgp ndy
phan bd va luu giu tren loan bp mach
naron khdng khu tni d mpl vj Iri cu I h l
ndo. Gidm ddn tmydn synap d ca naron
cam gidc, naron lien hgp hoac ca hai la
ca c h l pho b i l n giai thich hifm lugng
quen Ihudc hda d mpt vai dap ung nil lui
da nghien cuu ky Iren ldm (crayfish) vd
gidn (cockroaches) cung nhu phdn xg
gidt minh d dpng vai cd xuang s i n g . C a
c h l synap cda hien tugng quen thugc
hda cd I h l thay d l i Iheo hai each. (1) vi
tri suy giam dan Iruyin synap xual hign d
bat ky vi Iri synap nao. Vf du trong phdn
xg gap khdng cd s y suy gidm d i n truyen
synap giua naron cam gidc va naron lien
hgp ma s y suy gidm lai dien ra lgi synap
giua naron lien hgp vd naron vgn dgng.
(2) tang lac dyng tic c h l synap gdy ra
hidn lugng quen thupc hda. K i t qud
nghien cuu cung cho Ihay hpc lgp cd the
lam thay doi ddn Iruyin synap vd thdi
gian luu giu tri nhd ngan hgn dugc q u y l l
djnh bdi khoang Ihdi gian dan truyen
synap b i l n doi. Lien quan d i n nghidn
cuu nay, cau hdi dugc dgi ra la, Ihay d l i
d i n I m y i n synap d mtic do ndo vd s y
Ihay doi dy keo dai bao lau? Mpt d g i kich
Ihich 10 ldn len siphon cd the Igo ra Iri
nhd ngan hgn cho hign lugng quen Ihupc
hda keo dai vdi phdt, 4 d g i kich thich
each nhau l u vai gid d i n 1 ngdy giua cdc
d g i cd the hinh Ihanh Iri nhd ddi hgn kdo
ddi d i n 3 t u i n . Aplysia chua dugc h u l n
Iuygn cd den 90% naron d m gidc Igo
 V i e t n a m J o u m a l of P h y s i o l o g y 18(1), 5 / 2 0 1 4
cac k i t n l i synap trdn naron van ddng
mang. Ngugc Igi, khi con vgt d u g c huan
luyen quen thudc hda s l k l l n i l giam chf
cdn 30%; ly lg nay keo dai 1 ludn vd hoi
phyc khdng hodn todn 3 tuan sau do. Trong
giai dogn b i t hoat ddn I r u y i n synap keo ddi
nay t h y c s y da cd bidn doi d u trdc naron
cam giac. Khdng phai t i t ca synap d i u cd
kha nang Ihfch nghi n h u nhau. Dan t m y i n
tgi mol sd synap cua Aplysia hiem khi Ihay
doi, ngay ca khi d u g c hogt hda lap di lap
Igi. Tuy nhien, mpt so synap chuyen biet
cho hpc tap vd luu giu Iri nhd, chang han
k l l noi giua cac naron cam giac vdi naron
van dpng va vdi naron lien hgp cua mgch
naron phan xa gap, chl mol lugng huan
luydn it di dgc bigl vdi khoang each thdi
gian huan Iuygn Ihich hgp cung cd t h l tgo
ra mpt s y Ihay doi Idn va keo dai trong d i n
Iruyin synap. Huan luyen lidn l i l p khdng cd
khoang thdi gian nghf giua cac d g i kfch
thfch tgo ra tri nhd n g i n hgn rdt manh
nhung tri n h d ddi hgn lai kem. K i t ludn nay
dua ra nguyen ly chung cho hgc lap: huan
luyen each doan thudng tao ra tri nhd dai
hgn hieu qua han la h u l n Iuygn lien lyc
trong mol dgt.
3. Ca chi te bio va phan td cua
kien hoa kinh diin
Dieu kien hda kinh dien la hinh thuc hgc
tap phuc lap han so vdi cam giac hda. Con
vgl khdng chf hpc mol kfch thich ma la hpc
m l i lien he gida kich thfch nay vdi kfch
Ihich khac. Kich Ihfch cd d i l u kien y l u den
trudc cd I h l gay ra ddp ung cd hieu qua
cao neu d u g c ghep ddi vdi kich thich
khdng d i l u kien manh. Phan xa cd the
manh han do dieu kien hda kinh dien vd
cam gidc hda, tuy nhien d i l u kidn hda kinh
dien cd t h l lam phan xg tang mgnh va keo
dai han. Phan xg co siphon va mang
cua Aplysia la vf du cho ca dieu kien hda
kinh dien vd cam gidc hda. Phan xa co
mang cd the d u g c tao ra theo hai each:
kfch thfch tryc t i l p len siphon hoac kich
thich ldn d u trdc vd ngay canh do {mantle
ISSN: 1 8 5 9 - 2 3 7 6
shelf). Cam giac t u siphon va vd do hai
nhdm naron d m giac khac nhau I h y c
hien. Vi vay, cung phan xg cd d i l u kign
dpc lap v d i nhau bang each ghep ddi mgt
kfch Ihich cd dieu kien vdo vung Ihfch
hgp (hoac siphon hoac vd) vdi mgi kich
Ihfch khdng d i l u kien (shock dien manh
vdo dudi). Sau khi d u g c ghep cgp hogc
h u l n Iuygn iien h g p , ddp u n g mgch
naron vdi kfch thich cd dieu kidn mgnh
han ro rgt d cung phan xa cd ghdp cap
so vdi mgch naron cda dap ung vdi kich
thich cd d i l u kien khdng d u g c ghep cgp.
Doi vdi dieu kign hda kinh dien, khoang
thdi gian giu-a kich thfch cd dieu kidn vd
khdng dieu kien ddng vai Ird quan trpng.
Kich thfch cd d i l u kien phai d i n trudc
kich thfch khdng d i l u kign, thdng t h u d n g
khoang Ihdi gian giu-a 2 kfch thfch nay Id
0.5 giay (hinh 6). C a c h l t l bdo d n cho
ghep cap v l thdi gian 2 loai kich thich
nay la gi? Trong dieu kien hda kinh dien
cda phan xg co mang cua Aplysia, dgc
Irung quan Irpng chinh id Ihdi gian hpi ly
Ihdng Hn t u naron cam giac cho kich
thfch cd dieu kign (kich Ihich siphon) va
kich Ihfch khdng d i l u kien (shock dign
len phdn dudi). Kich thfch khdng d i l u
dieu
kidn len dudi Ihuc day hogt dpng naron
lien hgp Igo synap k i l u tryc-lryc vdi
naron cam giac mang thdng tin ti> siphon
va vd den, thuc day hogt dpng Irudc
synap cho hien t u g n g cam giac hda. Tuy
nhien, n l u kich Ihich khdng dieu kidn
(kfch thfch dudi) vd kfch thich cd d i l u
kien (kfch thfch siphon hoac vd) d u g c
ghep ddi sao cho kich Ihfch cd dieu kien
ngay t r u d c kich Ihich khdng d i l u kidn.
khi do cdc naron lien hgp dieu b i l n gan
k i t vdi kfch thfch khdng d i l u kign se hogt
hda naron cam giac ngay sau khi kfch
thich cd dieu kign hogt hda naron cdm
gidc. Hogi hda naron cam giac Iheo trinh
l y n h u vgy khi kfch thfch cd d i l u kidn vd
khdng d i l u kign each nhau vdi mgt
khoang Ihdi gian nhat djnh lam lang
 V i e t n a m J o u m a l of Physiology 18(1), 5 / 2 0 1 4
mgnh hoat d$ng trudc synap khi so sdnh
vdi Irudng hgp hai kich thich ndy khdng
ghep d p vdi nhau. Ddc d i l m ndy cua dieu
kidn hda kinh cHln d u g c gpi la phu thudc
vdo hogt ddng. Hidn tugng tang hogi dgng
synap nay phu thudc vdo hoat dgng cda cd
naron trudc vd sau synap. Kich thich cd
dieu kign se tgo ra cdc didn t h l hogi dpng
dan d i n ddng Ca^* di vao naron cam gidc
trudc synap, Ca^* hogt hda calmodulin.
Phuc hgp Ca^Vcalmodulin d u g c hogi hda
se gan vdi adenyl cyclase, k h u l c h dgi dap
ung vdi serotonin tang cudng san sinh
cAMP. Vi vdy, ca c h l te bdo cua d i l u kidn
hda kinh d i l n Iren cdc dudng d i n truyin
dan synap cda phan xg rdt lui Irdn
Aplysia mpl phan Id do ca c h l cdm gidc
hda trdn cdng mpl cung phdn xg. Tdc dyng
nay do adenyl cyclase cd lac dyng giong
nhu chat phdt hign dong thi {coincidence
detector). Cd mgt Adenyl cyclase chiing td
(1) cd mgt cdc phdn tti lien quan d i n kfch
thich cd d i l u kidn (dign t h l hoai dpng Irdn
naron cdm gidc vd ddng Ca^* di vao); (2)
kich thich khdng d i l u kign (kfch Ihfch dudi
Idm giai phdng serotonin), adenyl cyclase
ddp dng vdi cd kfch Ihfch cd d i l u kien ( g i n
vdo phdc hgp Ca^Vcalmodulin von dugc
hogt hda bdi Ca^* dl vao Irong naron cdm
gidc khi cd didn t h l hogt dgng) va kich
thfch khdng d i l u kidn (gan vao Gas dugc
hogt hda do serotonin gdn vao receptor).
Naron vdn ddng sau synap cua d i l u kien
hda kinh d i l n t r u y i n Ifn higu ngugc d i n
naron cam gidc. Trdn mang naron vdn
ddng cd hai logi receptor l i l p nhdn
glutamate' nonNMDA vd NMDA receptor
(NMDAR). Dau ngogi bdo cda kenh ion tao
ndn bdi NMDAR dugc Mg^*ddng kin khi
didn I h l mdng d gid trj dign the nghf. Trong
d i l u kign binh Ihudng vd khi da quen Ihupc
hda hogc cdm gidc hda Ihi chi cd logi non-
NMDA receptor d u g c hogi hda do NMDAR
bj ddng kin bdi ion Mg^*. Tuy nhidn. khi
kich thich cd d i l u kidn vd kich thich khdng
d i l u kidn d u g c ghdp ddi phu hgp gay ra
ISSN: 1 8 5 9 - 2 3 7 6
dieu kign hda kinh d i l n lam k h u cyc
mgnh mang t l bdo sau synap Idm bdt
Mg2* ra khdi kdnh gan vdi NMDAR vd
nhd dd Ca^* di vao trong l l bao. Ca^* di
vao trang naron van dpng dugc cho la
hogi hda con dudng lin higu glai phdng
cdc chdt tmydn dgt lin higu ngugc chieu
d u g c naron cam gidc tien synap l i l p
nhan. d ddy chung Idm tdng giai phdng
c h i t tmyen dat I h i n kinh Id- tan cdng
Irudc synap vao khe synap. Tdm Igi, 3 lin
higu ddng Ihdi d i n naron cam giac lam
tang cudng giai phdng c h i t i m y i n dgi
t h i n kinh trong d i l u kign hda kinh d i l n :
(1) hogt hda adenyl cyclase do Ca^+di
vdo trong naron cam giac do kfch Ihfch
cd d i l u kign gay ra; (2) hogt hda receptor
cda serotonin, receptor ndy gan vdi
adenyl cyclase, kel qud cda kich Ihich
khdng d i l u kidn; vd (3) Ifn higu ngugc
cho Ihay naron vdn dpng sau synap
dugc hoai hda do kich thfch khdng dieu
kidn.
Co- chi tri nhd dai h^n cua diiu ki^n
hda kinh dien
Ruoi giam dgc bigl huu higu cho cac
thao lac lien quan d i n gen. Seymour
Benzer vd cpng s y i l n dau lien cho I h l y
ruoi giam cd hien lugng dieu kien hda
kinh d i l n , vd dot b i l n rieng re 4 logi
gen: dunce, rutabaga, amnesiac, va PKA-
R1 lam mat kha nang hpc tdp. Nghien
ctiu cac ddt bien nay dua ra 2 k i t luan.
(1) i l l ca dpi b i l n deu lam m i t khd ndng
hlnh Ihanh d i l u kign hda kinh dien vd
cam gidc hda. (2) cd 4 logi dpi b i l n deu
t h i l u vang con dudng Ifn higu cAMP. Dpi
biln Dunce ddn din thieu hyt
phosphodiesterase, enzym giang hda
cAMP. Hau qua la, dpi b i l n ndy k h i l n
lugng cAMP cao bat thudng v u g l qua
ngudng dteu hda binh Ihudng. Dpi bien
Rutabaga d i n den I h i l u hyt adenylyl
cyclase phu Ihupc Ca^Vcalmodulin vd vi
vgy lam giam lugng cAMP xuong dudi
muc nen. DOl b i l n Amnesiac lam I h i l u
 V i e t n a m J o u m a l of P h y s i o l o g y 18(1), 5 / 2 0 1 4
hyl cdc c h l l tmyen dgt t h i n kinh bdn chat
peptid ldc ddng vao adenylyl cyclase, va
dot bien PKA-R1 d i n d i n thieu hut
PKA.Vgy, con dudng cAMP-PKA-CREB cd
vai trd the nao trong iuu giii tri n h d ddi
hgn? Cd the dp dyng de gidi Ihich cho cac
lodi va dang hpc lap khac nhau khdng?
Gan day, nghien cuu hpc tap va Iri n h d cd
I h l s u dyng n h i l u r u l i g i i m chuyen gen
hogc lang bpc lp gen, bat hogc i l l gen lidn
quan d i n linh nang khao sat Id cdng cu h l l
sue hO'u ich cho nghien cuu linh mdm deo
synap. B i n g Hip can nay William Quinn va
cpng s y Ihay rang chan lac dyng cua PKA,
Iham chi trong mot khoang thdi gian r i t
ngdn lam anh hudng d i n kha nang hpc vd
hinh Ihdnh tri nhd ngdn hgn cua ruoi gidm.
Thyc nghigm gdy dpi b i l n Hlu phan cd
hogt tinh catalytic cda PKA cho k i t qua
tuang t y . Hinh thanh tri nhd dai han d ruoi
giam ddi hdi tong hgp protein mdi. Rudi
g i i m bpc Id ca y l u to hogt hda (CREB) vd
uc c h l (CREB-2). Jerry Yin, Tim Tuily, vd
cpng s y cda hp t h i y rang bpc Id qua mtic
yeu to uc c h l (CREB-2), d u g c cho la ngan
chan bgc lp gen d u g c hoai hda bdi cAMP,
se lam mat tri n h d ddi han, khdng anh
hudng den hgc lap va tri nhd ngdn han.
Ngugc Igi, bdc Id v u g l muc y l u to hogt
hda (CREB) cd the tgo ra tri nhd ddi han
ngay ca vdi huan luydn tgo Iri nhd ngdn
han Iren ruoi g i i m binh t h u d n g .
4. Tang dien the sau synap keo dai
(long-term potentiation - LTP) c o le Id c c
c h l synap d o i vd'i t r i nhd- c d the t h u a t Igi
Tinh mim deo synap (synaptic
plasticity) keo dai d u d i dang lang dign t h l
sau synap keo ddi (LTP) lidn quan den r l i
nhieu he thong trf nhd thudt lgt dugc. Hien
lugng nay xay ra d hoi hai ma, d u true
nao lien quan vdi tri nhd thudt lai d u g c .
ISSN: 1 8 5 9 - 2 3 7 6
LTP cd the d u g c nghien c i i u Id- cac
lat d t nao. Kich thich didn vdo cac sgi
d i n tao ra didn t h l kich Ihfch sau synap
va gid tn dign the nay c d the ghi lai bang
didn c y c ghi dat d naron sau synap. Khi
khoang Ihdi gian giu-a cdc kJch thfch qua
ddi (phdt), bien do cua EPSP khdng thay
doi. N l u kfch Ihfch vdi lan s d cao,
khoang Ihdi gian giu-a cac kfch thich rat
n g i n (1 giay) (kfch thfch gay co cdng)
cac sgi den lao ra hai hinh Ihiic lang dien
t h l d naron sau synap. T r u d c h i t , giai
doan tang dien I h l ngan gpi Id tang dien
t h l hdu CO cung (post-tetanic potentiation
-PTP) hinh thuc nay nhanh chdng b i l n
mdt sau vai phul. T h u hai, hinh thuc tang
dien I h l kfch thfch sau synap kdo ddi hdu
CO cung ggi la LTP. LTP la c a c h l d n
Ihiet cho vigc luu g i i i Iri n h d dai han
(hinh 7). Receptor cua glutamate thugc
loai NMDA (NMDAR) ddng vai Ird quan
trgng de hlnh Ihanh LTP, dac bigl la LTP
x u l t hidn lai synap k i t n i l naron vung
CA3-CA1 d hoi hdi ma. Tgi cdc ny sau
synap (postsynapHc spines) cda cdc
naron vdng C A I cd hai logi receptor d l i
vdi glutamate: NMDAR va AMPAR (hinh
7). Ca hai logi nay d i u cho ion N a * v d K*
di qua nhung NMDAR cd them hai dac
Ifnh. (1) NMDAR ddng vai Ird chfnh cho
Ca^* dl vao Irong t l bdo. (2) binh thudng
kdnh nay bj ddng lai bdi ion Mg^*. Khi
glutamate gdn vdi NMDAR se lam thay
doi d u hinh cda receptor nay nhung ion
K* khdng di ra vd ion Na*, Ca^* khdng di
vdo d u g c bdi vi kdnh cho cdc ion nay bj
ion Mg2* bjt Igi. Vi vdy, mdt kich thich y l u
se khdng m d d u g c nhdng kenh ndy.
Kich thfch y l u lao ra mgt EPSP, nhung
EPSP nay Id do lac dyng vdo AMPAR
c h u khdng phdi NMDAR, t y a n h u
NMDAR khdng cd mgi d dd.
 N^etnam Joumal of Physiology 18(1), 5/2014 ISSN: 1859-2376

Hlnh 7. LTP dugc tgo ra d vdng CAI cda hot hai ma khi kfch thfch con dudng nhdnh
Schaffer vd md hlnh minh hpa ca c h l gay ra hign tugng LTP. A Trong d i l u kign blnh
thudng, t i n so dign t h l hoat dgng thap lam giai phdng glutamate tac dyng Idn ca
NMDAR vd AMPAR. Nhung ion Na vd K chi di qua kenh AMPAR chu khdng di qua
NMDAR do ion Mg bjt kin kdnh nay khi mang t l bao d gia tn dien the nghi
{Eric Kandel, 2000)

Kfch thich CO cung (tetanus) (hinh 7)
Idm EPSP xual hign Id l i n g k i t qud cCia lal
cd kfch Ihfch Iheo khdng gian va Ihdi gian
khi kfch thfch vao rat n h i l u sgi di den cung
mdt naron. K i t qud Id, mdng l l bdo bj khu
cyc mgnh, mgnh han r l i nhilu so vdi chi
mdi kfch Ihfch vdo sgi d i n . Ldc nay mat
Irong mdng t l bdo lich didn duang, ion
Mg2* bj bdl ra khdi kdnh (tich cfidn cung
diu). cdc kenh ion khdng cdn bj chan lai
nda. ion Ca^* cd I h l di vdo irong cac nu
dudi gai qua cdc kdnh ion gdn vdi NMDAR
da m d sSn. Ion Ca^* di vdo Irong l l bao
hogt hda m0l logi enzym protein kinase,
gdy ra hdng logt b i l n d l i . Nguyen nhdn ddn
d i n thay d l i ddi hgn Id hign tugng chen
mdi AMPAR vdo mdng sau synap (hinh
7). Do dd, sau kich Ihfch co cdng, cdc c h i t
truyen dai than kmh dugc giai phdng ra
t u cac naron trudc synap se g i n k l l vdi
mpl so lugng receptor nhilu han cd mgi
d mang sau synap. Khi nhieu receptor
g i n k i t vdi chat Iruyen dat than kinh se
tgo ra EPSP cd bien dp dign t h l ldn han
va LTP se dugc tao ra. Han nua, cung
cd bdng chung cho thay lugng c h l l
t m y i n dat Ihdn kinh dugc giai phdng ra
tti cac naron Irudc synap cung tang
len. Hieu ung trudc synap va sau synap
p h l i hgp, dien ra dong thdi cdng Idm
lang bien do EPSP. Luu y, ca c h l sau
synap d l i vdi tri nhd cd the thugt Igi cd
vai net tuang dong vdi co c h l synap ciia
Iri nhd khdng I h l thuat lgi Irong cam gidc
hda da dugc thao lugn d phan trudc
Mac du ve chi till thi khac, nhung cd hat
 Vietnam Joumal of Physiology 18(1). 5 / 2 0 1 4 ISSN: 1 8 5 9 - 2 3 7 6

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 Vietnam Joumal of Physiology 18(1). 5/2014 ISSN: 1859-2376

21. Kimble, GA (1961). Hiigard and learning is experience of what was

Marquis' Conditioning and Learning. acquired.
2^ Edition. New Yorit Appleton-Century- Anatomical basis of analysis and
Crofts. molecular mechanism of learning and
22. Nicoll, RA, JA Kauer and RC Malenka memory is one of the greatest successes
(1988). The cun^nl excitement In long-termof modem neuroscience. Thirty years ago
potentiation. Neuron 1:97-103. little was known at)out how memory
23. Sargent, SS and KR Stafford (1965). works, but now we know a great deal. In
Basic Teachings of the Greatthis report we will mention four main
Psychologists. Garden City, NY: Dolphin content related to learning and memory.
Books. First of all, we refer to a classification of
memory in view of modem neuroscience.
MOLECULAR MECHANISM AND TREND Next up well refer to areas of the brain
OF RESEARCH ON LEARNING AND associated with learning and memory.
Memory is stored in our brain are similar
MEMORY PRESENT
to stored on chip memory of personal
Le Dinh Tung MD, PhD^-^ computer or not? entire archive from
^Head of Department of Physiology, memory into the same region of the brain
Hanoi Medical University structure or distribution and stored In
'General Secretary of die Vietnam diffemnt brain regions? How does
Association of Physiology memory work? The changes would take
place in the nen/ous system in the
Memory is defined as the process of process of memory formation? Is there
encoding, storing and mtrieving any special proteins or genes related io
information. Three processes involve with memory or not? How to memory can
different memory system. Sensory memory,maintain throughout life? will be
short-term memory in which to note the presented in the next section. Finally, a
concept of woriiing memory and long-termcritical Issue, especially as we age: How
memory. Learning and memory are two can memory be maintained and
closely related concepts. Memory is impmved. how can It be fixed when it is
acquisition of knowledge or skill while brokened?