Gio trnh sinh hc i cng

Bin tp bi:
Nguyn Hi

Gio trnh sinh hc i cng

Bin tp bi:
Nguyn Hi
Cc tc gi:
Nguyn Hi

Phin bn trc tuyn:

http://voer.edu.vn/c/3207fce7

MC LC
1. S hnh thnh tri t v kh quyn
2. Ngun gc ca s sng
3. S tin ha ca t bo
4. Hc thuyt t bo v cc phng php nghin cu t bo hc
5. Thnh phn ha hc ca t bo
6. T bo Eukaryote
7. Cu to ca t bo Prokaryote
8. Cc qu trnh sinh hc trong t bo
9. S a dng ca t bo
10. Khi nim v cu trc Enzyme
11. C ch hot ng,phn loi v cc yu t nh hng n hot tnh Enzyme
12. H hp t bo
13. Quang hp
14. Lch s pht trin ca di truyn hc
15. i tng v phng php nghin cu di truyn hc
16. Quan h gia di truyn hc vi cc khoa hc khc v vi thc tin
17. Acid Nucleic l vt cht di truyn cp phn t
18. Ti bn DNA
19. Nhim sc th
20. S phn bo
21. Gene v m di truyn
Tham gia ng gp

1/132

S hnh thnh tri t v kh quyn

Thuyt v tr hin nay c nhiu ngi cng nhn l thuyt i bng n (Big Bang) .
Theo thuyt ny mt khi nguyn t s khai khng l n v vt cht pht tn thnh
cc m my bi v kh v tr nhit rt cao cch nay 13 t nm.

Hnh 1.1: Thuyt Big Bang

Mt tri v cc hnh tinh ca n c hnh thnh t nhng m my bi v kh v tr
ny. Phn ln vt cht c c thnh khi rt nng gi l mt tri. Phn cn li hnh
thnh
cc hnh tinh trong c tri t quay quanh mt tri, cch nay khong 4-5 t nm. Khi
qu t c c, cc phn t nng nh Fe, Zn, Ni di chuyn vo tm, cc cht nh tp
2/132

trung gn b mt. Cc cht kh nh He, H2 hnh thnh nn kh quyn tri t u tin.

Tuy nhin, qu t nh nn trng lc yu, cc cht kh bay vo v tr li qu t
khng c kh quyn.
Sc nn ca lc hp dn, s tan r phng x l nguyn nhn lm trong lng tri t nng
chy hnh thnh li ch yu l Fe, Ni. Li nng c bao bc bi Manti (Silicat v Mg)
lng v ngui hn. Lp ngoi cng hay v tri t rn li to thnh lc a v i dng.

Hnh 1.2: Cu to ca Tri t.

Qu t ngui dn qua nhiu giai on. Cc kh nng bn trong thot ra ngoi qua ni
la hnh thnh nn kh quyn th hai. Bu kh quyn c xa c tnh kh mnh khng
c oxygen t do. Theo Oparin, kh quyn c xa bao gm: NH3, H2O, CH4. Mt s gi
thuyt khc cho rng kh quyn c xa cn c thm CO, CO2, H2 , N2, H2S v HCN.

3/132

Hnh 1.3: S hnh thnh kh quyn th hai (theo Oparin)

Trong thi gian , hi nc ngng t to ra nhng trn ma dm. Nc tp trung vo
cc ch trng hnh thnh nn i dng u tin. Cc dng nc mang mui khong
tch ly bin.

4/132

Ngun gc ca s sng
Theo nhiu gi thuyt, sinh vt u tin c to ra t mt qu trnh tin ho ha hc
trong 4 giai on: Tng hp v tch lu cc cht hu c c phn t lng nh t cc
cht v c c sn; Polymer ho cc cht hu c phn t lng thp thnh cht hu c
c phn t lng cao; S kt hp cc cht hu c tng hp bng con ng v c thnh
cc t bo (protobions) c nhng tnh cht ho hc khc vi nhng cht quanh chng;
ngun gc di truyn.

Tng hp v tch lu cc cht hu c c phn t lng nh t cc cht v c

c sn
Nm 1920, Oparin a ra gi thuyt, cc cht hu c c th c tng hp t nhng
cht v c c sn trong kh quyn v i dng. Cc cht hu c ny l cc amino acid,
ng t NH3, CH4 v hi nc trong kh quyn c xa. Cc sinh vt u tin xut hin
ngu nhin t dung dch m c nng ca cc cht . Tuy nhin, gi thuyt ny khng
c cng nhn v khng c thc nghim.
Nm 1953, Stand Miller v Harold Urey bng thc nghim chng minh cht hu c
n gin c th hnh thnh t cht v c theo con ng ho hc trong iu kin tri t
c
xa. Trong m hnh th nghim, Miller to ra iu kin tng t nh trn tri t c xa.
H thng ny gm: mt bnh nc un nng 80oC; bnh cu kh quyn gm: CH4,
NH3, H2; in cc pht tia la in (tia chp); h thng lm lnh (tri t ngui dn).
Sau khi Miller cho h thng ny hot ng mt tun, thu dung dch th nghim v phn
tch thnh phn. Kt qu cho thy, s c mt ca nhiu cht hu c cn cho qu trnh
tng hp cc i phn t sinh hc nh amino acid, lactate, acid hu c Th nghim
ca Miler chng minh mt s bc trong gi thuyt ca Operin. iu ny m ra
bc ngot mi trong tm hiu ngun gc ca s sng.

5/132

Hnh 1.4: M hnh tng hp cht hu c bng con ng ha hc

Nhiu phng th nghim lp li th nghim ca Miller nhng thay i thnh phn kh
quyn, dng cc tc nhn nh nh sng thng, tia X, tia phng x thay th cho tia
la in. Kt qu thu c 20 amino acids, purin (A & G), pyrimindin (C, T & U) v
ATP nu thm phosphate.

S hnh thnh cc cht hu c phc tp t cht hu c n gin

Cc cht hu c n gin tch ly trong mi trng nc polymer ha hnh thnh cc
cht hu c phc tp nh protein, nucleic acidTrong t bo cc phn ng ny c

6/132

enzyme xc tc, nhng trong qu trnh tng hp ha hc khng c enzyme v nng

cc
monomer trong nc thp. Nh vy lm th no cc phn ng polymer ho xy ra? Vn
nay khng n gin v c nhiu gi thuyt. Mt s cho rng nng cc cht hu c
trong trong bin nguyn thy l rt cao nn c kh nng gn kt vi nhau to thnh cc
polimer. Nhng ngi ny thm ch cho rng khng c enzyme xc tc, cc phn ng
to thnh cng c th xy ra trong thi gian di.
Tuy nhin, nhiu nh khoa hc cho rng, nng cc cht trong cc i dng c xa
khng khng m c thc hin polimer. Theo h phi c c ch c hc lm tng
nng . Mt trong gi thuyt cho rng di sc nng ca mt tri nc bc hi
cht hu c phn t lng nh tp trung trong h nc nh c c c. Mt gi thuyt
khc cho rng t st c kh nng huy ng cc monomer hu c do cc monomer ny
bm trn ht st tch in. Cc monomer hu c polymer ho to thnh cc hp cht
hu c cao phn t. Nhng polymer hu c c sng, ma cun tri tr li ao h v
i dng. Qu trnh c lp li nhiu ln lm cho nng cc cht hu c cao phn
t tng cao. Hai gi thuyt ny dc Sidney Fox (Clay theory: thuyt t st) v
Guterwachtershauser (ion pyrite theory: thuyt pirit st) chng minh bng thc nghim.

S hnh thnh t bo u tin

Sau khi cc polimer hnh thnh chng phi gn vi nhau to thnh cc i phn t lm
tng tnh phc tp ca t chc.
Nhng tnh cht ca s sng xut hin t s tng tc ca cc phn t c t chc
thnh nhng mc cao hn. Nhng t bo sng c th bt ngun gc t t bo
(protobionts: mt khi kt ca cc phn t) c to ra bng con ng ha hc. Nhng
t bo ny cha c kh nng sinh sn nhng chng duy tr mi trng ho hc bn trong
khc vi mi trng xung quanh v c biu hin mt vi c im ca s sng chng
hn nh trao i cht (metabolism), d b kch thch (excitability).
Mt trong nhng loi t bo c oparin gi l coacervate c th t lp rp khi
lc dung dch c cha cc phn t lipids, proteins, nucleic acid v polysaccharides.
Coacervate tch bit vi mi trng ngoi bi mng k nc. Cc ht coacervate c
th hp th enzymes v cc cht khc t mi trng v gii phng cc sn phm ca
phn ng enzymes. Khi hp th cc cht, coacervate sinh trng v phn chia thnh cc
coacervate nh. Cc coacervate c
thnh phn tt hn to ra v phn chia tip. Theo Operin, chn lc t nhin s gi li v
hon thin cc git tt hn to thnh t bo.

7/132

Mt t bo khc c Fox(1960) gi l tiu cu (microphere), c th c to ra khi

trn proteinoid vi nc ri un nng n 130-180 oC ri lm lnh dn qua 1-2 tun
trong pH v nng mui nht nh. Mt vi tiu cu c mng thm chn lc, c kh
nng xc tc mt vi phn ng nh thy phn glucose v c kh nng phng in (ging
t bo thn kinh). Cc tiu cu c kh nng ny chi v to ra cc tiu cu khc.
Mt t bo khc na l liposome c th hnh thnh trong t nhin khi thnh phn
dung dch c lipids. Mng ca liposome l lp lipid i ging mng t bo. Liposome
c kh nng sinh trng bng cch ha nhp cc liposome nh v sinh sn bng cc tch
liposome ln thnh nhng liposome nh.
Khng ging nh cc m hnh th nghim, t bo khng c cc enzyme tinh nh trong
t bo. Mt vi cht c tng hp bng con ng ho hc c kh nng xc tc yu
cho php t bo bin i cc cht hp th qua mng. Kh nng sng st ca cc t
bo tng ln theo hng hon thin cu trc bn trong, tng cng b mt ngn cch
vi mi trng, s phc tp v tnh hiu qu ca qu trnh trao i chat. Chn lc t
nhin s chn lc v hon thin cc t bo c nhiu u im to nn cc t bo u
tin v tip tc tin ha cho n ngy nay.

RNA c th l nguyn liu di truyn u tin

Cc t bo a dng v tnh thm, kh nng xc tc, sinh sn, sinh trng mi trng
s chn lc nhng t bo thch nghi v o thi nhng t bo khng thch nghi. Cc c
tnh ca t bo khng th duy tr v tin ha qua cc th h cho n khi xut hin mt
vi c ch di truyn.
Trong t bo thng tin di truyn c m ha trong nucleic acid (DNA & RNA). Nhiu
gi thuyt cho rng gen xut hin trc:
Nm 1929, G. Muller mt nh di truyn hc nu ra gi thuyt s sng bt u t mt
hoc mt vi gen c to thnh khng do cc sinh vt. trong mt thi gian di gi
thuyt ny khng c ch . Tuy nhin, cc dn liu t sinh hc phn t cho thy gii
thuyt trn ngy cng c l v nhng l do sau.
Th nht: Cu trc phn t v s ti sinh ca virus. Chng ta bit rng khi xm nhp
vo vi khun ch co1ADN hoc ARN c bm vo v t n sao chp ri to ra cc ht
virus mi.
Th hai: Trong qu trnh tng hp protein, ngai AND v mARN cn c s tham gia
ca tARN v rARN iu ny cho thy nucleic acid c trc.
Th ba: Nhiu nucleotide gi vai tr a dng v quan trng ca t bo tt c cc cc
sinh vt.
8/132

Hin nay cha c m hnh c th no cho thy qu trnh xut hin s sng l t nucleic
acid chng minh bng thc nghim. Nhng theo thuyt ny cc vt sng u tin l cc
i phn t c kh nng sau chp. Cc t bo u tin ny tch ly mt cch chm , v
bao bn ngoi bi cc cht khc. Mt bng chng minh ha r cho c ch ny l cc
virus cha ANR or RNA c cu to n gin.
Nhiu gi thuyt cho rng RNA xut hin trc DNA bi v:
RNA bn hn, ti bn.
RNA c kh nng nhn i t mch khun mu nhanh hn v t li hn cc
trnh t khc.
Vd: Mt trnh t RNA c 40 ribonucleotide c th t nhn i trong mi trng c km
lm xc tc vi sai st thp hn1%
RNA (ribozyme) c kh nng xc tc (Thomas Cech, 1980s): T bo hin i
s dng ribozyme xc tc tng hp cc RNA mi (rRNA, tRNA v mRNA.
Nh vy, RNA l cht t xc tc v trong th gii tin sinh hc trc khi c
protein v DNA, RNA c kh nng t ti bn.
RNA d tng hp hn DNA
S sai st trong qu ti bn cng vi tc ng ca chn lc t nhin to ra s
a dng ca RNA
Nh vy, trong mt thi gian di, vt liu di truyn ca cc tin sinh vt l RNA v s
tin ha dn n ch DNA mch kp n nh hn mang thng tin di truyn. v kh nng
xc tc c chuyn cho protein lm chc nng chuyn ha hu hiu hn.
Theo gi thuyt hin nay, s sng c hnh thnh qua cc bc:
S hnh thnh cc phn t RNA
C ch sao chp RNA
Hon thin h thng nh mng bao
Cc t bo tin ha theo 3 bc:

9/132

Hnh1.5 : S ti ha ca t bo
C th tham kho thm ngun g ca s sng ti: http://en.wikipedia.org/wiki/
Origin_of_life

Qu trnh hnh thnh s sng khng th xy trong iu kin hin ti v:

Oxy trong kh quyn c tch lu do hai qu trnh. Qu trnh phn li nc do nh sng
cc tm tc ng ln hi nc v qu trnh quang phn li nc trong quang hp.
-Oxy c trong kh quyn s phn hy cc cht hu c va tng hp.
-Kh quyn ngy nay c tnh oxy ha. Kh quyn c tnh kh tng cng phn ng kt
hp nhng cht n gin thnh cht phc tp.

10/132

-Tng hp cht hu c cn nng lng, UV (mt tri true to nhiu UV). iu kin hin
ti khng p ng c bi v tng Ozon kh quyn ngn cn cc tia UV.
-Sinh vt tn ti tiu th cc cht hu c to ra.

11/132

S tin ha ca t bo
Cc dn liu ho thch v ng thc vt
Tui nin i a cht c th xc nh thng qua thng qua tui ca lp t trm tch
c hnh thnh t ct, bn ca y h v i dng. Trong lp trm tch rt giu cc
ho thch sinh vt. C th dng phng php ng vi phng x xc nh tui ca
v ha thch. Cac ng vi phng x phn hy rt chm. Bng nhng k thut thch hp,
ngi ta tnh c tng lng cht phn r c th nh gi tui ca v cc ho
thch.
Vd: U238 chu k bn r 4.5 t nm, C14 chu k bn r 5600 nm

Hnh 1.6: Dng C14 xc nh tui ca v trai (clam shell)

Cn c vo c vo tui ca v cc ho thch, ngi ta chia s sng thnh 5 i: i
thi c, nguyn sinh, c sinh, trung sinh, tn sinh. Hnh bn di m t c th sng t
dng ban u sm nht v thi gian tng ng vi cc i a cht t tin Cambri.
Hnh 1.7: Cc i a cht v lch s s sng trn tri t.
12/132

Trc y cc nh a cht xem k Cambri l mt trong nhng im mc quan trng

trong nghin cu tin ho v khng thy mu ha thch no c hn. Tuy nhin mi
y, bng k thut mi cc nh a cht pht hin cc vi ho thch ging nh vi khun
trc k Cambri c tui khong 3100 triu nm trong cng Chert en.
Nh vy trong mt thi gian di, cch nay khong 3 t nm, trn tri t ch tn ti cc
sinh vt nh b, n gin tng t nh cc vi khun ngy nay.

T prokaryote n eukaryote, gun gc t bo nhn chun-Thuyt ni cng

sinh
Trong sut qu trnh hnh thnh eukaryotes, cu trc t bo v nhng qu trnh c trng
cho eukaryote xut hin: nhn c bao bc bi mng, ti th, lp th, h thng ni
mng, a nhim sc th (nhim sc th gm DNA v protein).
Prokaryote hnh thnh, tin ha v thch nghi t khi s sng xut hin v tr nn ph
bin nht ngy nay. Mt hng tin ha ca prokaryote l hnh thnh cc prokaryote a
bo v d nh vi khun lam. Hng th hai l hnh thnh tp hp t bo mi loi t bo
c li t vic chuyn bit ha trao i cht ca t bo khc. Hng th ba l phn
cch chc nng khc nhau trong cc t bo n. Hng tin ho ny to ra nhng t bo
eukaryote u tin.
Lm th no m s t chc cc bung ca eukaryote tin ha t prokaryote? Mt qu
trnh m trong h thng ni mng ca eukaryote: mng nhn mng li ni cht
nhm, Golgi c th tin ho t gp np mng prokaryote. Mt tin trnh khc c gi
l ni cng sinh to ti th v lp th trong eukaryotes.
Theo thuyt ni cng sinh, ti th ca Eukaryote c ngun gc t prokaryote t dng
hiu kh, lp th ca Eukaryote c ngun gc t vi khun quang hp c th l khun lam
(cyanobacteria).
C nhiu bng chng ng h thuyt ni cng sinh:
Cu trc ca ti th, lp th tng t nh vi khun
Mng trong ca ti th, lp th c h thng cc enzymes vn chuyn in t
trong mng vi khun.
Ti th, lp th nhn i tng t nh trc phn vi khun
Ti th, lp th c DNA vng ging prokaryotes
Mt s khng sinh km hm sinh trng ca prokaryote cn tr tng hp
protein bi ribosome ca ti th v lp th nhng khng cn tr tng hp protein
ca ribosome t bo cht. Khng sinh ngn cn s tng hp protein ca t bo
cht khng nh hng tng hp protein ca cc bo quan.

13/132

Hnh 1.8: Mt m hnh ngun gc ca eukaryote

T n bo n a bo
S pht trin theo hng phc tp ha t chc dn ti s hnh thnh v pht trin cc
sinh vt a bo
-Cu trc phc tp
-B my sinh sn phc tp
-S bit ha t bo
-Mt s h thng chuyn bt

S pht trin ca sinh vt a bo

Bng 1: Cc i a cht v lch s s sng trn tri t

14/132

15/132

Hnh 1.9: S tin ha ca sinh vt a bo

16/132

Hc thuyt t bo v cc phng php

nghin cu t bo hc
Hc thuyt t bo
Lch s pht hin t bo
Nm 1665, khi quan st lt ct g si (oak tree) di knh hin vi c phng i 30
ln (30X), Robert Hooke pht hin nhng hp nh v t tn chng l t bo.
Antoni Van Leeuwenhock pht hin gii vi sinh bng knh hin vi c phng i 300
ln (300X).
Nm 1839, Mathias Scheiden v Theodor Schwann tm tt nhng nhng kt qu nghin
cu di knh hin vi ca h: Tt c sinh vt u cu to t t bo, t bo mi c hnh
thnh t s phn chia ca t bo trc .
Nm 1862, Louis Pasteur bng thc nghim chng minh s sng khng t ngu sinh.
Nhng tuyn b ny l nn tng cho hc thuyt t bo.
Ni dung c bn ca hc thuyt t bo
(1) T bo l n v cu trc v chc nng ca c th sng
(2) Tt c c th sinh vt u c cu to t t bo.
(3) T bo c kh nng phn chia hnh thnh cc t bo mi.
(4) T bo c bao bc bi mng c vai tr iu ha hot ng trao i cht gia t
bo v mi trng.
(5) Tt c t bo c s ging nhau cn bn v thnh phn ha hc v cc hot tnh trao
i cht gia tt c cc loi t bo.
(6) T bo cha DNA mang thng tin di truyn iu ha hot ng ca t bo mt s
giai on trong i sng ca n.
(7) Hot ng ca c th l s tch hp hot tnh ca cc n v t bo c lp

17/132

(8) C hai loi t bo: prokaryote v eukaryote. Chng khc nhau trong t chc cu trc
t bo, hnh dng v kch thc nhng cng c mt s c im ging nhau, chng hn
nh tt c u l nhng cu trc mc cao, thc hin cc qu trnh phc tp cn thit
duy tr s sng.
Classical interpretation
1.
2.
3.
4.
5.

All organisms are made up of one or more cells.

Cells are the fundamental functional and structural unit of life.
All cells come from pre-existing cells.
The cell is the unit of structure, physiology, and organization in living things.
The cell retains a dual existence as a distinct entity and a building block in the
construction of organisms.

Modern interpretation
The generally accepted parts of modern cell theory include:
1.
2.
3.
4.
5.
6.
7.
8.
9.

The cell is the fundamental unit of structure and function in living things.
All cells come from pre-existing cells by division.
Energy flow (metabolism and biochemistry) occurs within cells.
Cells contain hereditary information (DNA) which is passed from cell to cell
during cell division
All cells are basically the same in chemical composition.
All known living things are made up of cells.
Some organisms are unicellular, i.e., made up of only one cell.
Others are multicellular, composed of a number of cells.
The activity of an organism depends on the total activity of independent cells.

Exceptions
See also: Origin of life
1. Viruses are considered by some to be alive, yet they are not made up of cells.
Viruses have many of the features of life, but by definition of life, they are not
alive.
2. The first cell did not originate from a pre-existing cell. There was no exact first
cell since the definition of cell is not that precise. This is an intellectual game
that comes from making strict logical symbols out of the biological definitions.
3. Mitochondria and chloroplasts have their own genetic material, and reproduce
independently from the rest of the cell.
Types of cells

18/132

Cells can be subdivided into the following subcategories:

1. Prokaryotes: Prokaryotes lack a nucleus (though they do have circular DNA)
and other membrane-bound organelles (though they do contain ribosomes).
Bacteria and Archaea are two divisions of prokaryotes.
2. Eukaryotes: Eukaryotes, on the other hand, have distinct nuclei and membranebound organelles (mitochondria, chloroplasts, lysosomes, rough and smooth
endoplasmic reticulum, vacuoles). In addition, they possess organized
chromosomes which store genetic material.
{accessed from http://en.wikipedia.org/wiki/Cell_theory )

Cc phng php nghin cu t bo hc

Hin vi
T bo c kch thc rt nh nn khng th nhn thy bng mt thng. S pht hin
knh hin vi gip nghin cu t bo nhng kha cnh khc nhau. iu quan trng i
vi knh hin vi khng ch phng i m cn gii hn phn gii.
Knh hin vi quang hc phng i khong 2000 ln, c th phn bit c khong
cch nh nht l 0.2m.
Knh hin vi in t c phng i khong 250.000 ln, c th phn bit n .
Gn y nhiu ci tin knh hin vi c thc hin v nhiu loi knh hin vi mi ra
i phc v tt hn cho nghin cu t bo nh knh hin vi hunh quang, knh hin vi
ni.
Tch v nui cy t bo
Trong nhiu trng hp vic nghin cu tng loi t bo l cn thit, tin hnh nhiu th
nghim , do cn mt s lng ln t bo . Cc phng php tch chit v nui cy
t bo ngy cng c ci tin v hon thin p ng nhu cu ny.
Phn on cc thnh phn t bo
Cc thnh tu khoa hc cung cp nhiu phng php cho vic tch ring cc bo quan
v cc i phn t sinh hc nghin cu thnh phn sinh ha v vai tr ca chng
trong t bo.
Cc phng php thng c p dng: Phng php siu ly tm, phng php sc k.

19/132

Ngoi ra, trong nghin cu t bo hc cn s dng nhiu phng php hin i khc
nh: in di, nh du bng ng vi phng x v khng th

20/132

Thnh phn ha hc ca t bo
Thnh phn nguyn t:
Trong t bo c th c mt hu ht cc nguyn t trong t nhin (92 nguyn t). Trong
, 25 nguyn t c nghin cu k l C, H, O, N, S, P, K, Ca, Mg, Fe, Mn, Cu, Zn,
Mo, B, Cl, Na, Si, Co l cn thit cho s sng. Trong , C, H, O v N chim 96%
trng lng cht kh ca t bo, cc nguyn t cn li chim 5%.
Cc nguyn t trong t bo c th c chia thnh 3 nhm: cu to cht hu c, cc ion
v nguyn t vi lng (i lng, vi lng v siu vi lng)
T l v vai tr ca tng nguyn t trong t bo cng khc nhau.

Thnh phn hp cht

Cc hp cht c bn ca t bo: Nc, cc hp cht v c, cc hp cht hu c, kh ha
tan.
Nc
Nc l thnh phn bt buc ca t bo v chim mt t l cao chng hn sa l
70-80%, ngi l 60%.
Tnh cht ca nc:
L mt cht phn cc: H+ - O- - H+
C th to lin kt hydrogen gi cc phn t nc v vi cc phn t cht khc.
Nhit dung ring ln (1cal/g/OC)
Vai tr ca nc:
L mt dung mi tt.
Tham gia trc tip vo cc phn ng ha hc (phn ng thy phn).
L nguyn liu cho hot ng ca t bo (cung cp proton H+ cho cc phn ng)
iu ha trng thi ca nguyn sinh cht: (sol, coacevate v gel).
21/132

To sc cng b mt t bo.
n nh cu trc t bo.
iu ha nhit t bo.
Cc cht v c
Trong t bo ngoi nc cn c nhiu cht v c khc: acid, base, cc ion c vai tr
quan trng trong cu trc v hot ng sinh l ca t bo.
V d: Fe. Cu, Mn l thnh phn ca cc enzyme; Cc mui ha tan l y t quan trng
trong s hp thu nc ca t bo.
Cc kh ha tan
Trong nguyn sinh ct ca t bo c cha cc kh hoa tan: CO2, O2,
Cc hp cht hu c
Carbohydrate
Bao gm c cc ng v polymer ca chng: monosaccharide, disaccharide v
polysaccharide.
Cng thc chung: CnH2nOn
Carbohydrate l nguyn liu cu trc v nhin liu ca t bo.
Monosaccharide
Monosaccharide l nhng ng c 3 n 7
Tiu ch phn loi: Phn loi da vo:
V tr gc carbonyl (C=O): ng c th l aldose (aldehyde) hay ketose (ketone).
S lng nguyn t carbon (hnh 2.1 a).
V tr ca nhm OH gn vo C1 trong cu trc vng: -glucose hay -glucose (hnh 2.1
b).
S sp xp trong khng gian ca 4 loi nhm th quanh carbon bt i xng (chiral
carbon): ng dng D hay L (ng phn quang hc).
22/132

Trong dung dch, glucose v hu ht cc monosacchride khc hnh thnh dng vng.
thun tin ngi ta nh s carbon trong vng bt u t carbon (C1) lin kt vi
oxygen gn vi carbon. (hnh 2.2)
Tt c ng n l ng kh (C5).
Disaccharide
Mt disaccharide c cu thnh t hai monosaccharide nh lin kt glycoside - mt
lin kt cng ha tr bi phn ng dehydrate ha gia hai monosaccharide (hnh 2.3).
Cc n phn ca sucrose v maltose c xp nh nhng ng tin nga.
Cc n phn ca lactose (-1,4 glycoside) c xp nh nhng ng tin nga-sp.
Polysaccharide
Polysaccharide l nhng polymer ca monosaccharide.
Cu trc v chc nng ca polysaccharide ph thuc vo cc monomer ca n v v tr
ca lin kt glycoside.
Tinh bt (starch)
Tinh bt l cht d tr thc vt, c tm thy trong lc lp (chloroplast), c (khoai
ty), mm.
n phn cu trc ca tinh bt l alpha-glucose lin kt vi nhau bi lin kt alpha-1,4
hay 1,6 glycoside.
Mt phn t tinh bt c khong 280-300 phn t glucose.
Tinh bt c hai dng: amylose v amylopectin.
Amylose
Mch thng.
Trong amylose cc phn t glucose lin kit vi nhau bng kin kt alpha-1,4 glycoside.
Amylopectin
Mch phn nhnh.

23/132

Cu trc tng t nh amylose. Tuy nhin, bt u ca mi nhnh l lin kt alpha-1,6

glycoside.
C 24-30 n phn glucose li c mt nhnh.
Glycogen
Glycogen l cht d tr ng vt, c tm thy gan, c.
n phn ca glycogen l alpha-glucose.
Cu trc ca glycogen tng t nh amylopectin nhng phn nhnh sau 8-12 phn n
phn glucose
Cellulose
Cellulose l nguyn liu chnh ca vch t bo thc vt, c n phn cu trc l glucose.
Cc monomer lin kt vi nhau bng lin kt -1,4 glycoside.
S khc nhau trong lin kt ca cc monomer trong cellulose, tinh bt dn n s khc
nhau trong cu trc khng gian 3 chiu (3D) ca chng:
Tinh bt c cu trc xon, mi bc xon c 6 n v glucose c c n nh nh
lin kt hydrogen gia cc vng xon k nhau.
Cellulose mch thng. Cc phn t cellulose xp song song nhau, lin kt vi nhau bng
cc lin kt hydrogen gia cc gc OH t do lm thnh cc vi si (microfibrill) rt chc,
l nguyn liu cho xy dng t bo.
Chitin:
Chitin l polysaccharide, c cu trc tng t nh cellulose nhng gc OH ca C2 c
thay th bi NH - CO-CH3 (acetyl-glucosamine)
Chitin l nguyn liu xy dng v ca chn khp, vch t bo nm.
Lipid
Lipid l nhng i phn t sinh hc nhng khng phi l mt polymer. Cc lipid c
chung mt tnh cht l khng hoc t c i lc vi nc bi v chng cha phn ln gc
hydrocarbon v ch mt mt vi lin kt phn cc vi oxygen. Lipid khng tan trong
nc nhng tan trong dung mi hu c.
24/132

Lipid gi vai tr quan trng trong t bo: phospholid l thnh phn quan trng ca t
bo; l cht d tr nng lng , l cht cch nhit; l dung mi ha tan cc cht nh
vitamin (A,D, E,K)
Du m v sp
Du m c cu thnh t glycerol v cc acid bo (fatty acid).
Acid bo c mt u COOH v mt ui hydrocarbon no (saturated) hay khng no
(unsaturated). S nguyn t carbon ca acid bo vo khong 16-18.
M ng vt, triacylglycerol cha acid bo no d ng nhit phng. Ngc li, du
c v du thc vt thng cha cc acid bo khng no nn ng c nhit thp hn.
Du v m c th l tri, di hay mono-acylglycerol.
Cc acid bo trong du v m c th ging hay khc nhau.
Sp: mt lng nh acid bo lin kt vi ru mch di thay v glycerol.
Du m cha nhiu nng lng.
V d: Mt gram du m cha nng lng gp i mt gram carbohydrate.
Phosphol ipid
Hai gc -OH trn glycerol lin kt vi acid bo v gc -OH th ba lin kt vi acid
phosphoric.
Gc phosphate c th lin kt vi nhng gc phn cc khc to nn s da dng ca
phospholipid.
Phospholipid c mt u phn cc (-) a nc (hydropholic) v mt ui khng phn
cc k nc (hydrophobic).
Trong mi trng nc, phospholipid hnh thnh cu trc ht micelle vi u a nc
quay ra ngoi v u k nc quay vo trong. b mt mng t bo, phospholipid hnh
thnh lp i vi u a nc quay ra ngoi v u k nc quay vo nhau.
Phospholypid l thnh phn chnh ca mng t bo.
Steroid

25/132

Steroid l nhng lipid c xc nh bi mt sn carbon c bn vng lin kt. Cc

steroid khc nhau do s khc nhau cc nhm th.
Cholesterol l mt steroid tham gia thnh phn cu trc mng t bo ng vt v l tin
cht ca cc steroid khc. Tuy nhin nng cholesteron trong mu cao gy va x
ng mch (atherosclerosis).
Protein
Protein l mt polymer c cu thnh t 20 loi amino acid. Mt protein c th c
hnh thnh t mt hay nhiu polypeptide cun thnh cc cu hnh c trng.
Polypeptide l polymer ca cc amino acid. Cc amino acid lin kt vi amino acid k
tip bng lin kt peptide. Mt u ca polypetide l NH2 (N-terminus), u kia tn
cng bng COOH (C-terminus). Polypeptide c trng bi trnh t amino acid.
Cc amino acid khc nhau ch yu nhnh bn R v trong sinh vt ch tn ti cc Lamino acid.
Bn mc cu trc ca protein
Chc nng ca protein ph thuc vo cu hnh c trng ca n (conformation).
Trnh t ca polypeptide c th xc nh cu hnh khng gian ba chiu (3D) ca protein.
Cu trc bc mt (primary structure)
Cu trc bc mt ca protein l trnh t amino xc nh ca n. Trnh t amino acid
c xc nh bi thng tin di truyn. S thay i trong cu trc bc mt c th nh
hng n cu hnh v chc nng ca protein.
Cu trc bc hai
Chui polypeptide c th gp li thnh mt s cu trc u n trong khng gian. Cu
trc bc hai c bit n nhiu nht l xon (-helix). Sn polypeptide hnh thnh
mt cu trc xon phi vi 3,6 acid amin trn mt vng xon; nh vy nhm N-H trong
lin kt peptid th n to lin kt hydro vi nhm C=O trong lin kt peptide th (n+3)
ca chui. Nhng phn c cu trc xon thng tm thy trong cc protein hnh cu
v mt s protein hnh si.
Cu trc tm gp np (-pleated sheet), thng gi l gp , c n nh bi cc lin
kt hydro hnh thnh gia cc nhm N-H v C=O ca cc phn khc nhau trong chui
polypeptide. Mt vi on ca chui polypeptide c th c xp cnh nhau to nn
cu trc tm, trong cc nhnh bn R c th hng ln pha trn hoc pha di tm.
26/132

Nu cc on ni trn chy cng chiu (v d t u tn cng N n C), ta c tm song

song (parallel), nu chng xp khc chiu (N n C v C n N) ta c tm isong song
(antiparallel). Cc tm rn chc, ng vai tr quan trng trong cc protein cu trc, v
d nh trong si fibroin.
Protein collagen trong m lin kt cn c cu trc xon ba (triple helix), trong ba
chui polypeptide c bn vo nhau khin cho n rt chc.
Cu trc bc ba
Cu trc bc ba l cch thc m chui polypeptide vi nhng on c cu trc xon ,
gp hay cc cu trc bc hai khc cng vi cc vng ni (connecting loop) gp li
trong khng gian ba chiu.
Bn cht ca cu trc bc ba vn c nh hnh sn t cu trc bc mt. Khi t vo
iu kin thch hp, hu ht cc chui polypeptide t ng gp li thnh mt cu trc
bc ba ng bi v cu trc ny c nng lng thp nht c ngha l bn nht. Trong cu
trc bc ba, cc on cu trc bc hai v cc on ni gp li sao cho hu ht cc acid
amin a nc th quay ra b mt cn cc acid amin k nc th nm bn trong protein.
iu ny mang li s n nh cho ton b cu trc.
Cc yu t gip n nh cu trc bc ba gm c cc lin kt yu nh lc van der Waals,
lin kt hydro, lin kt ion, tng tc k nc xy ra gia cc nhnh bn trong chui
polypeptide. i khi cng c s tham gia ca mt dng lin kt cng ha tr: lin kt
disulfide hnh thnh gia hai cysteine.
Cu trc bc bn
Cu trc bc bn l s t chc nhiu chui polypeptide ging hoc khc nhau thnh mt
phn t protein. V d nh phn t hemoglobin c hai chui globin v hai chui globin
. Nhng lc lin kt gip n nh cu trc bc ba k trn cng l lc gip cho cc chui
polypeptide ny gn li vi nhau.
Chc nng ca protein
Cc cht xc tc: cc enzyme ribonuclease, cytochrome, trypsine (thy gii peptide)
Protein cu trc: glycoprotein, keratin
Protein vn chuyn: hemoglobin
Protein vn ng: myosin, actin
Protein bo v: khng th
27/132

Cc cht c hot tnh sinh hc: insulin, hormone

Nucleic acid
Nucleic cha v truyn thng tin thng tin di truyn
C hai loi nucleic acid: DNA v RNA, y l nhng i phn t gip sinh vt to nn
nhng thnh phn phc tp ca sinh vt t th h ny n th h k tip. DNA iu
khin qu trnh ti bn ca n, tng hp RNA v qua RNA iu ha tng hp protein.
DNA l vt cht di truyn ca sinh vt c nhn t b m. Khi t bo phn chia, DNA
c sao chp v truyn t th h t bo n th h t bo k tip.
Thng tin di truyn m ha trong DNA chng trnh ha hot ng ca t bo. Tuy
nhin, khng phi DNA m l protein iu khin trc tip hot ng ca t bo chng,
l cng c cho hu ht chc nng sinh hc.
Mi gene iu khin tng hp mt mRNA, sau tng tc vi b my tng hp protein
iu khin tng hp mt polypeptide. Ni tng hp protein thc s l ribosome. Nh
vy, thng tin di truyn c chuyn t nhn ra t bo cht.
Nucleotide n phn ca acid nucleic
Nitrogenous (amine) base
Cc base ca ADN v ARN c cu trc d vng (heterocyclic) thm. Cc purine c cu
trc hai vng, gm adenine (A) v guanine (G). Cc pyrimidine c cu trc mt vng,
gm cytosine (C), thymine (T) v uracil (U).
A, G. C tm thy trong c hai loi nucleic acid, thymine tm thy trong DNA v uracil
c tm thy trong ARN. Thymine khc vi uracil ch c thm mt nhm methyl
v tr s 5, v vy thymine chnh l 5-methyluracil.
Nucleoside
Trong cc acid nucleic, mi base lin kt ha tr vi v tr s 1 (C-1) ca mt ng
pentose to nn mt nucleoside. V tr ca lin kt ny trn base l v tr 9 (N-9) i
vi cc purine v v tr 1 (N-1) i vi cc pyrimidine.
ARN, ng pentose l ribose. Cn ADN, l 2-deoxyribose, trong nhm
hydroxyl v tr s 2 b thay bi mt nguyn t hydro.

28/132

Lin kt gia base v ng c gi l lin ktglycoside. ARN, cc nucleoside

gm adenosine, guanosine, cytidine v uridine. i vi ADN, tn cc nucleoside gm
deoxyadenosine, deoxyguanosine, deoxycytidine v deoxythymidine (hay thymidine).
Nucleotide
Mt nucleotide gm mt nucleoside cng vi mt hay nhiu nhm phosphate ni ha tr
ti v tr 3, 5 hoc 2 ca ng pentose (v tr 2 ch c ng ribose). Nu l ng
ribose, ngi ta gi hp cht l ribosenucleotide; cn nu l ng deoxyribose th
gi l deoxynucleotide. V mt ha hc, cc hp cht ny l nhng phosphate ester.
Trong trng hp c t mt n ba nhm phosphate gn vo v tr 5, ta c cc
hp cht 5-monophosphate, -diphosphate v -triphosphate tng ng; v d nh AMP
(adenosine 5-monophosphate), dGDP (deoxyguanosine 5-diphosphate), dCTP
(deoxycytidine 5-triphosphate), TTP (thymidine 5-triphosphate) hay ATP (adenosine
5-triphosphate)
Cc nucleoside 5-triphosphate (NTPs) hay cc deoxynucleoside 5-triphosphate
(dNTPs) l vt liu cu thnh nn phn t acid nucleic a phn. Trong qu trnh tng
hp ARN hay ADN, hai phosphate c tch ra, ch li mt nhm phosphtate cho
mi n phn tham gia vo chui nucleotide. Nucleotide chnh l n phn ca acid
nucleic.
Lin kt phosphodiester
Trong mt chui nucleotide ca ADN hoc ARN, mi phosphate lin kt ha tr vi
mt pentose v tr 5 v mt pentose k tip v tr 3 to thnh mt lin kt 3, 5phosphodiester. C th hnh dung mi chui nucleotide bao gm mt sn chnh gm
cc ng pentose xen k vi cc phosphat; trong , c mi ng li c mt base gn
vo v tr 1.
pH trung tnh, mi phosphate u cha in tch m v vy cc acid nucleic l nhng
polymer mang in tch m.
Trnh t ADN/ARN
Mi chui nucleotide (tr chui nucleotide dng vng) u c mt u 5 t do c
th gn hoc khng gn vi cc nhm phosphate v mt u 3 t do c mt nhm
hydroxyl. nh hng ca mi chui nucleotide l 53 v theo quy c vit trnh
t th u 5 nm pha bn tri. V d khng c vit trnh t mt on ADN mt
mch l ATACGTA, m phi vit l 5-ATACGTA-3. Mt trnh t ARN c th vit l
5-AUGCUUGA-3. iu ny c ngha l hng ca chui l xc nh, AUGCUUGA
khc hn vi AGUUCGUA.

29/132

Chui xon kp ADN

Cu trc chui xon kp
Cu trc ph bin nht ca ADN l cu trc chui xon kp (double helix). Cc c tnh
c bn ca cu trc ny c James Watson v Franois Crick xut vo nm 1953.
Phn t ADN l mt chui xon kp gm hai mch n xon u quanh mt trc, mi
mch n l mt chui nucleotide. Mi chui c nh hng 53; hng ca hai
mch trong chui xon kp l ngc chiu nhau nn ngi ta gi chng l hai mch i
song song. Mi chu k xon ca ADN gm 10 bp (base pair cp base) di khong
3,4nm, ng knh vng xon khong 2nm.
Sn phosphate-ng ca mi mch n hng ra ngoi, cn cc base ca chui xon
kp hng vo trong. Hai mch n kt hp vi nhau nh cc lin kt hydro hnh thnh
gia cc cp base b sung nm trn hai mch. A lin kt vi T bng hai lin kt hydro
v gia G v C l ba lin kt hydro.
Trn y ch l m hnh ca ADN dng B theo Watson v Crick. Ngy nay, ngi ta
bit rng ADN c nhiu dng h cu trc khc nhau mt vi ch s.
Dng B thng tn ti trong iu kin sinh l bnh thng, cn cc dng khc tn ti
trong nhng iu kin m v ion khc nhau. c bit, dng Z c chiu xon ngc
v pha bn tri v theo hnh zigzac.
Bng 1.1: Mt s dng cu trc ca ADN
S cp base
Chiu v gc xonso
Dng
trong mtchu k vi mt phngca
ADN
xon
base

Khong cch thng

ng gia hai base k
nhau

ng
knhvng
xon

11

Xon phi 32,7o

2,56

23

10

Xon phi 36,0o

3,38

19

9 v1/3

Xon phi 38,6o

3,32

19

12

Xon tri 30, 0o

3,71

18

T cc d kin v cu trc chui xon kp ca ADN cho ta hai khi nim c bn:

30/132

 Mi mch n l mt trnh t base khc nhau. Nh vy mi mch n mang

thng tin khc vi mch kia.
Hai mch n lin kt vi nhau bi mt quan h b sung. Chnh quan h ny
gip gii thch c cu trc cht ch ca phn t ADN, c bit l phng
cch t ti bn to ra hai phn t con ging ht nhau t mt phn t mban
u.
ngha ca cu trc chui xon kp
Phn t ADN thng c cu trc chui xon kp. Cu trc ny l mt cu trc n nh:
Trong chui xon kp, cc ng pentose v cc nhm phosphate xoay ra
ngoi, hnh thnh lin kt hydro vi nc m bo tnh n nh cho phn t.
Chui xon kp cho php cc base purine v pyrimidine c cu trc phng xp
chng kht ln nhau bn trong phn t ADN, hn ch s tip xc ca chng vi
nc. Nu hai n tch ri nhau, cc base k nc s phi tip xc vi nc,
iu ny s t chng vo mt tnh th bt li, khng n nh.
Hai mch n bt cp vi nhau nh cc lin kt b sung gia mt bn l purine
(A v G cng kch thc ln) v bn kia l pyrimidine (T v C cng kch thc
b hn). iu ny m bo cho hai mch n lun i song song.
Mi phn t ADN c mt s lng lin kt hydro rt ln nn d chuyn ng
nhit c lm ph v cc lin kt nm hai u phn t th hai mch n vn
c gn vi nhau bi cc lin kt vng gia. Ch trong nhng iu kin rt
khc nghit, v d nhit cao hn nhit sinh l nhiu ln, th mi c s ph
v ng thi qu nhiu lin kt hydro khin phn t b bin tnh, khng cn gi
c cu hnh ban u.

31/132

T bo Eukaryote
Nhn
Phn ln t bo c mt nhn. Mt s trng hp, t bo c nhiu nhn nh Paramecium
(2 nhn), mt s t bo gan v t bo nc bt ca ng vt c v (2-3 nhn), mt s t
bo a nhn c n hng chc nhn (megacaryocyte trong ty xng). Tuy nhin, cng
c t bo khng nhn chng hn nh t bo hng cu.
Hnh dng ca nhn ty thuc vo hnh dng t bo: t bo hnh cu (lymphocyte, t
bo nhu m, neuron), nhn thng c hnh cu; trong cc t bo hnh tr, hoc hnh ko
di theo mt trc (t bo c, t bo biu b), nhn c hnh bu dc; mt s nhn c hnh
phc tp (nhn phn thy ca t bo bch cu).
Kch thc ca nhn ph thuc loi v trng thi chc nng ca t bo, trung bnh
khong 5m.
Cu trc nhn kh phc tp:
Mng nhn l mng kp (double membrane) tch bit nhau khong 20-40nm. Mi mng
dy khong 10nm v c cu trc bi lp phospholipid kp tng tc vi cc protein.
Mng ngoi ni vi h thng mng li ni cht (endoplasmic recticulum-ER) bi cc
khe b cha v hnh thnh h thng khe. Trong mt s trng hp, h thng khe ny m
ra khong gian bo lin h trc tip vi mi trng ngoi.
Mng nhn c cu trc khng lin tc. Trn mng nhn c cc l nhn (nuclear pore),
c ng knh khong 100nm. Ti mp ca l ny, mng ngoi v mng trong ni vi
nhau.
L nhn c cu to t mt phc hp protein gi l phc hp l. Phc hp ny ni cc
l vi nhau c vai tr iu ha kch thc l v vn chuyn cc cht c kch thc ln
(large macromolecules and particles) qua l.
Ngoi l mng nhn, trn mt ca mng nhn c cc phin mng (lamina) lt mt trong
ca mng, m bo n nh hnh dng ca nhn.
Ngoi vic tch bit nhn v t bo cht, mng nhn c chc nng trong trao i cht,
tng hp protein v n nh hnh dng nhn.
Nhn cha hu ht gene ca t bo. Cc DNA t chc vi protein lm thnh cht nhim
sc. Cht nhim sc (chromatin) bt mu c th nhn thy di knh hin vi nh mt
32/132

khi ln xn. Khi t bo chun b phn chia, chromatin xon li v dy ln gi l nhim

sc th (chromosome) c s lng c trng cho mi loi.
Hch nhn (nucleolus) c tm thy trong nhn ca t bo khi khng phn chia. y l
t chc tng hp cc thnh phn ca ribosome.
Nhn c vai tr cha v truyn thng tin di truyn t t bo ny sang t bo khc. Nhn
cha DNA, nhim sc th. Khi t bo phn chia, DNA v nhim sc th nhn i v
phn chia v hai t bo con.
Nhn iu khin, iu ha hot ng sng ca t bo. Ct trng Stentor, thnh hai phn
mt na c nhn v mt na khng c nhn. Mt na c nhn nhanh chng phc hi.
Na kia s cht sau . i vi Foraminifera, na c nhn phc hi v cng b mt,
na kia khng c kh nng ny.

Ribosome
Ribosome l trung tm tng hp protein. Nhng t bo c t l tng hp protein cao
thng c s lng ribosome ln chng hn nh t bo gan ca ngi (vi triu
ribosome), nhng t bo c t l tng hp protein thp c t ribosome.
Ribosome c hnh cu kch thc nh (20-35nm).
Trong t bo, ribosome bm trn h thng mng mt ngoi ca li ni cht, mng nhn
(bound ribosome) v nm t do trong cytosol.
Ribosome t do tng hp protein c chc nng trong t bo cht, chng hn nh cc
enzyme tham gia vo qu trnh chuyn ha cc cht (metabolitic process).
Ribosome bm tng hp cc protein c xut ra khi t bo. Nhng t bo chuyn ha
tit protein nh t bo cc tuyn chng hn nh tuyn ty (pancreas) c t l ribosome
bm cao.
Ribosome c cu to ch yu t RNA v protein. Ribosome ca prokaryote c hng
s lng l 70S gm hai hai tiu phn 30S v 50S. t bo eukaryote, ribosome c hng
s lng l 80S gm 2 tiu phn 40S v 60S.

Mng li ni cht (endoplasmic recticulum-ER)

Mng li ni cht l mt h thng mng chng cht (labyrinth) chim hn 50% tng
mng trong t bo eukaryote. Mng li ni cht bao gm cc ng, ti v b thng vi
nhau phn b khp t bo cht.

33/132

C hai loi mng li ni cht khc nhau. Mng li ni cht nhm (rough ER) v mng
li ni cht trn (smooth ER)
Mng li ni cht trn (smooth endoplasmic recticulum)
Mng li ni cht trn c chc nng trong chuyn ha cht nh tng hp lipid, chuyn
ha carbohydrate, phn gii ha cht (drug) v c t v co c.
Vd: Mng li ni cht trn ca t bo c (muscle cell) bm Ca2+ t cytosol vo trong
ti. Khi t bo c b kch thch bi cc xung thn kinh, Ca2+ c a ngc li vo
cytosol gy co c.
Mng li ni cht nhm (rough endoplasmic recticulum)
Mng li ni cht nhm mt ngoi c cc ribosome. nhiu loi t bo, protein tit c
tnh chuyn ha c tng hp ribosome ca mng li ni cht nhm chng hn nh
insulin ca tuyn ty. Mt polypeptide c tng hp c ng gi vo ti hay bng
mng (vesticle) thng qua l, c hnh thnh bi protein trn mng ca mng li ni
cht. Khi i vo ti, polypeptide xon thnh cu hnh khng gian c trng. Hu ht cc
protein tit l glycoprotein. Cc ti protein tit (transfort vesicle) sau tch khi mng
li ni cht v c vn chuyn n ni cn thit.

B my Golgi (Golgi apparatus)

Sau khi ri khi mng li ni cht, cc bng vn chuyn (transport vesicle) c a
n b my Golgi. y cc sn phm ca mng li ni cht c bin i, phn loi
sau a n ni cn thit.
B my Golgi gm nhng ti nh dp c mng kp xp song song nhau ta nh chng
a. Cc ti dp gn nhau ni vi nhau bi cc ng.
Cc ti dp c cu trc phn cc: mt cis v trans. Mt cis nm gn mng li ni cht
v tip nhn cc bng vn chuyn t mng li ni cht. Cc sn phm ny c bin
i trong Golgi t mt cis ti trans. Nhng bin i c th l loi b mt s monomer
trn gc ng ca protein, gn gc phosphate. Cc sn phm ca Golgi sau tch
khi mt trans n ni cn thit.

Lysosome
Lysosome l cc ti enzyme thy phn c bao bc bi mng. Cc enzyme ny c th
hot ng tt pH khong 5 v c th thy phn hu ht cc i phn t nh protein,
polysaccharide, nucleic acid v lipid. Mng ca lysosome m bo pH thp bng cch

34/132

bm H+ t cytosol vo. Cu hnh protein ca mt trong mng lysosome v enzyme tiu

ha m bo trnh s t phn hy lysosome.
Lysosome c vai tr trong:
- Tiu ha ni bo: amip, khng bo tiu ha (food vacuole) ha mng vi lysosome
tiu ha thc n.
- Ti ch nguyn liu thng qua qu trnh t tiu (autophagy) cc bo quan gi.
- Hy t bo theo chng trnh. Trong qu trnh pht trin ca nng nc thnh cc,
lysosome phn hy cc t bo ui. Trong qu trnh pht trin phi ngi, lysosome
phn hy mng ngn tay phi.
- Nhiu bnh di truyn gi l d tr lysosome (lysosomal storage diseases) nh hng
n chuyn ha ca lysosome. Mt ngi b bnh ny thiu active lysosome enzyme
hot ha. Cc lysosome b ng c cht nh hng n hot ng t bo.
Vd: Bnh Pompe: gan b tn hi do tch ly nhiu glycogen. Bnh Tay-Sarch, tch ly
lipid no.
Cc bnh ny c th cha tr bng cch tim enzyme c lin quan hay bng liu php
gene.

Khng bo (vacuole)
Khng bo (vacuole) v bng mng (vesicle) u c bao bc mi mng nhng khng
bo ln hn. Khng bo c nhiu chc nng. Khng bo tiu ha (hnh thnh do thc
bo phagocyte), khng bo co bp (contractile vacuole), khng bo trung tm (central
vacuole t bo thc vt trng thnh) c bao bc bi mng (tonoplast). Khng bo
t bo thc vt cha protein, cc cht v c (kali, chloride), sn phm trung gian ca
ha trnh chuyn ha cc cht, sc t, c t.
Khng bo c vai tr trong s ln ln ca t bo thc vt: do khng bo ko di khi ht
nc.
Khng bo ln c hnh thnh t vic kt hp nhiu khng bo nh hn.

Ti th (mitochondria)
Trong t bo ca eukaryote thng c nhiu ti th, mt s trng hp c mt ti th ln.
nhng t bo hot ng trao i i cht mnh, s lng ti th nhiu hn t bo hot
ng t.
35/132

Ti th c hnh ht hay hnh si ng knh trung bnh 0.1-0.5m, di1-10m.

Quay phim hin vi cho thy ti th lun chuyn ng, thay i hnh dng v phn chia.
Ti th c mng i, mng ngoi trn, mang trong gp np thnh mo c hnh rng lc
(cristae) lm tng thm b mt ca mng trong nhm tng cng hiu qu chc nng ca
n. Trn b mt ca rng lc c cc ht l cc protein xuyn mng (integral protein) v
ra mng (peripheral protein) c vai tr trong chuyn in t v tng hp ATP.
Mng trong chia ti th thnh hai bung: bung th nht (internal compartment) l vng
hp gia hai mng; bung th hai cht nn ca ti th (mitochondrial matrix) c bao
quanh bi mng trong cha enzyme thc hin mt vi bc chuyn ha cht trong h
hp t bo. Ngoi ra, cht nn cn c cc DNA. V vy, ti th c kh nng t nhn i
v tng hp mt s protein.
Ti th l ni thc hin h hp t bo, mt qu trnh d ha (catabolism) to ra ATP t
vic oxy ha ng, lipid v nhng nhin liu khc trong s c mt ca oxy.

Lp th (chloplast)
thc vt khng b nh sng t trc tip, lc lp c hnh cc hnh sao, hnh bn, hnh
chung
thc vt trn cn, lc lp c hnh bu dc. Vi hnh bu dc, lc lp c xoay b mt
iu chnh mc tip xc vi nh sng v s dng nh sng hiu qu nht. y cng
l mt c im tin ha ca gii thc vt.
S lng lc lp khc nhau cc loi thc vt. Mi t bo to c mt lc lp. Thc vt
bc cao trung bnh c 20-100 lc lp.
ng knh lc lp khong 4-6 micrometer, dy 2-3 micrometer.
Nhng cy a bng c s lng, kch thc lc lp v hm lng sc t trong lc lp
nhiu hn cy a sng.
Lc lp c bao bc bi mng kp, chia lc lp thnh hai bung: khong gia hai mng
v cht nn (stroma).
Stroma l ni xy ra cc phn ng ca pha ti trong quang hp. Thnh phn ha hc ca
stroma bao gm: Cc enzyme, protein, acid nucleic, ribosome, lipid v cc git du...
Grana l tp hp cc thylakoid xp chng cht ln nhau l ni xy ra cc phn ng
pha sng trong quang hp. Mng thylakoid c phc h enzyme tng hp ATP (ATPsynthetase). Trong mng cha h thng sc t v h thng chuyn in t. Mt grana
36/132

c 5-30 thylakoid c gi l thylakoid grana. Mi lc lp c khong 40-50 grana c

lin kt vi nhau bi thylakoid stroma.
Lc lp c vai tr trong chuyn i nng lng nh sng thnh nng lng ha hc v
dng n trong vic tng hp cht hu c t CO2 v nc. Lc lp c cha DNA, nh
vy chng c th t sinh sn v tng hp mt s protein.

Peroxisome
Peroxisome l cc bng mng, c mng n. Bn trong l cht nn cha cc enzyme
oxy ha c trng: catalase phn gii hydrogen peroxide, enzyme chuyn ha acid bo.
Peroxisome gan c vai tr trong gii c bng cch chuyn H t cht c cho oxygen.
Mt bo quan chuyn bit ca peroxisome l glyoxisome khng c ng vt c xng
sng bc cao. Bo quan ny cha enzyme chuyn ha lipid thnh carbonhydrate.

B xng t bo (cytoskeleton)
B xng t bo l mng li vi si, vi ng m rng khp nguyn sinh cht (cytoplasm)
v c vai tr chnh trong t chc cu trc v hot ng ca t bo.
Chc nng:
B xng t bo c chc nng nng v duy tr hnh dng ca t bo. Kin trc ca
b xng chc chn v n hi cao; b xng cung cp im t cho cc bo quan v rt
linh ng do c th b phn hy v ti thit lp.
B xng t bo lin quan n cc dng vn ng ca t bo. S chuyn ng i hi
s tng tc gia b xng v cc protein vn ng (motor) chng hn nh cc bng
vn chuyn trn cc ng ray (monorail) t b xng; co c
B xng cng tham gia vo iu ha hot ng sinh ha ca t bo. B xng c th
chuyn lc c hc t b mt t bo vo bn trong v n nhn. Trong th nghim, ngi
ta ko protein ca mng sinh cht lin kt vi b xng. Mt video hin vi cho thy c
s sp xp li nhn v cc t chc khc ca hch nhn (nucleoli).
B xng t bo c cu trc bi: vi ng (microtubule), vi si (microfilament) v vi
si trung gian (intermediate filament).
Vi ng
Vi ng c tm thy tt c eukaryote, l nhng ng thng rng gia di khong
200nm -25m, ng knh khong 25nn.

37/132

Vch vi ng c cu trc bi protein hnh cu (globular protein) gi l tubulin. Mi

tubulin c hai thnh phn: alpha v beta tubulin. Cc thnh phn ca tubulin c th tch
ri v ti s dng.
Vi ng cng c th tp hp thnh lng, roi, cc vi ng thn kinh ca si trc (axon), thoi
phn bo.
Vi ng c vai tr:
Ko nhim sc th v hai cc, nh cc vi ng ca thoi v sc.
Vn chuyn cc bo quan t ni ny n ni khc: chuyn ng ca ti th, bng mng
xut nhp bo .
Duy tr hnh dng t bo: do s sp xp c trng ca vi ng (bit ha t bo)
Vi si
Vi si ng knh khong 7 nm, cu trc bi hai si actin xon li, c tm thy trong
tt c eukaryote. Vi si dng ny phn b khp cytoplasm nhng thng tp trung st
ngay mng sinh cht. Cu trc ca chng chu lc cng (tension forces). Cc vi si tng
tc vi cc protein khc to thnh mng li c vai tr nng v duy tr hnh dng t
bo. t bo ng vt chuyn ha cho vn chuyn cc cht, cc b microfilament lm
thnh li ca lng t lm tng din tch b mt t bo.
Dng vi si dy hn l myosin tng tc vi actin c vai tr trong vn ng ca t bo.
S co c l kt qu ca vic trt ln nhau ca actin v myosin lm t bo ngn li.
S tng tc ca actin v myosin trong t bo ng vt ang phn chia hnh thnh rnh
phn ct chia t bo m thnh hai t bo con.
Vi si trung gian
Vi si trung gian ph bin trong eukaryote c ng knh 8-12nm. Cc si ny chuyn
bit cho vic chu lc cng v rt a dng. Si trung gian l thnh phn c nh ca
t bo v c vai tr trong vic m bo hnh dng t bo. Khi x l loi b microtube
v microfilament, t bo vn gia nguyn hnh dng. Nh vy, mng li cc si trung
gian m bo hnh dng nguyn thy ca t bo.

Trung th (centriosome)
Trung th c cc t bo ng vt a bo cng nh n bo v nm cnh nhn.
Trung th t bo ng vt c mt cp trung t (centriole) v cht quanh trung t
(pericentriole). Ngy nay, ngi ta gi trung th l MTOC (microtubule organizing
centre). T bo thc vt khng c trung t.
38/132

Trung t c hnh tr ng knh khong 0.15-0.25 m di 0.7 m. Thnh tr c cu

to t 9 b ba vi ng. Cc vi si bao quanh tm tr. B ba vi ng gm tubilin A, B v C
v c ni vi nhau bi protein nexin. Khi t bo phn chia trung t nhn i.
Trc y cho rng, trung t c vai tr trong to thoi v sc. Nhng nhiu bng chng
gn y cho thy nhng t bo b ph trung t vn hnh thnh thoi v sc.
Cht quanh trung th c nhiu cu trc hnh cu kch thc khong 40-70nm c cung
nh vi cc vi ng ca trung t v cc vi ng t do xp phng x quanh trung t (hnh
2.37)

Lng v roi
Lng v roi l nhng phn li ca t bo cht c bao bi mng c ng knh khong
0.2m. Ngi ta phn bit lng v roi da vo chiu di, s lng v cch thc hot
ng (beating patern). Roi thng di hn lng (10-200m so vi 2-20 m). T bo c
khong vi roi trong khi c ti khong 300 lng. Roi chuyn ng un sng to lc cng
hng vi trc ca roi. Lng chuyn ng nh mi cho (oar) to lc vung gc vi
trc ca lng.
Mc d c chiu di, s lng v chuyn ng khc nhau nhng lng v roi c chung
cu trc siu hin vi (ultrastructure). Lng v roi c bao bc bi mng sinh cht v
gm 9 i microtubule xp song quanh 2 microtubule gia (9+2). i vi ng gm
alpha v beta microtubule lin kt vi nhau bi protein nexin. Alpha microtubule c hai
tay l protein dynein hng ti beta microtubule ca cp k. Alpha microtubule ni vi
mng bao hai microtubule trung tm bi ni phng x. Cu trc lng v roi c neo
vo t bo nh th nn (basal bodies) tng t nh trung t c vai tr trong ti sinh lng
v roi. Dynein c vai tr trong chuyn ng ca lng v roi. C ch chuyn ng ca
DYNEIN ging nh mo leo cy v cn nng lng (ATP).
Lng v roi gip t bo chuyn ng v dn cc cht lng trn b mt m. Lng t bo
m (lining: nim mc) ca kh qun qut nhng mnh vn ra khi phi.

Vch t bo (cell wall)

Vch t bo thc vt bn chc v quy nh hnh dng t bo. Mt trong cua vch dnh
vi mng nguyn sinh cht.
Vch t bo bao gm cc thnh phn cu trc sau: Cellulose chim 30%, hemicellulose
v pectin chim 60%, protein, lipid, peroxydase, ATPase v phosphatase. t bo gi,
vch t bo c thm lignin (ha g), cht sp, cutin(cuticle), tannin v suberin

39/132

Cc si microfibil ca cellulose an kt trong mt khi c cht v nh hnh ca

polysaccharide (hemicellulose v pectin) v cc protein.
Khi t bo cn non, chng c bo bc bi vch s cp. Gia vch s cp ca cc t
bo ln cn l cc phin gi (middle lamina) thnh phn chnh l pectin, cht kt dnh
cc t bo li vi nhau thnh khi m. Khi t bo trng thnh, mt s t bo thm thm
cc cht lm vch s cp cng hn, mt s khc hnh thnh vch th cp (secondary cell
wall) nm gia mng sinh cht v vch s cp, c nhiu lp phin mng hn.
Vch t bo c cc chc nng:
Trao i ion nh nhm carboxyl c trong pectin vi H+, K+ v cc cation khc.
Cht suberin tham gia iu tit ch nc v nhit ca cy.
Vch t bo cnh nhau lin kt li thnh h thng gian bo, c vai tr trong
vn chuyn vt cht.
Vch t bo c cc l, ni m cc si lin bo xuyn qua, m bo cho s
thng thng, s vn chuyn cc cht v truyn thng tin gia cc t bo.
Vch t bo duy tr hnh dng t bo nh p sut nc v quyt nh kch thc
ca t bo.

Mng nguyn sinh (plasma membrane)

Cu trc
Mng sinh cht l mng khm lng, l lp sinh cht ngoi cng ca t bo cht, c cu
to c th ca mng sinh hc bao gm:
Lp phospholipid kp: Cc phn t phospholipid sp xp vi u k nc quay vo nhau
v u a nc quay ra mi trng to nn lp phospholipid kp. Cc hydrocarbon
u k nc ca phn t phospholipid c th chuyn ng theo nhiu hng to nn
tnh linh ng ca mng. Cc phn t phosphilid c th di chuyn t ni ny sang
ni khc trong cng mt lp phospholipid hoc chuyn t lp phospholipid ny sang
phospholipid khc.
Cc protein :
Mt s protein xuyn qua lp phospholipid kp mt phn hay hon ton. Phn khng
phn cc ca protein tng tc vi u k nc ca phospholip v thng c cu trc
xon alpha. Phn a nc ca protein quay ra pha ngoi mng.
Mt s protein khng xuyn mng: Cc protein neo trc tip vo phospholipid kp hay
thng qua phn t ng hay phn t lipid.

40/132

Cc protein gn vo mng thng qua s tng tc vi protein khc ca mng.

Cc protein trn c th l cc enzyme, cc cht nhn hay cht mang, cc knh vn
chuyn qua mng, protein iu ha v protein cu trc.
Cng ging nh phospholipid, mt s protein c th di chuyn trong mt lp hoc
gia hai lp phospholipid. Tuy nhin, do protein ln hn cc phn t phospholipid nn
chuyn ng chm hn nhiu.
Nhiu phn t protein mt ngoi ca mng gn vo b xng t bo tht s bt ng.
S gn xen ca cc protein vo mng c vai tr iu ha tnh cht c hc ca mng.
Cholsterol (hnh 2.40)
Cc phn t cholesterol xp xen k vo gia cc phn t phospholip theo cch nhm
phn cc xp mc cc u a nc ca phospholipids v nhn steroid xen k vo ui
hydrocarbon ca phospholipids. Cholesterol c vai tr c nh c hc cho mng.
Cc glycoprotein v glycolipid (hnh 2.40)
Mt s protein v phospholipid mt ngoi mng gn vi cc oligosaccharide bng lin
kt ha tr to nn cc hp cht glycoprotein v glycolipid.
Cc glycoprotein v glycolipid l im nhn bit cc tn hiu v quan h gia cc t bo.
Tnh cht ca mng sinh cht
Mng c tnh lng (fluidity): Ph thuc v thnh phn cu trc ca mng (phospholipid
c dui l cc hydrocarbon khng no lm tng tnh lng ca mng v ngc li;
Cholesterol lm gim tnh lng ca mng).
Mng c cu trc khm: Cc protein khm vo lp phospholipid kp.
Mng nguyn sinh l mng bn thm c chn lc, cho nc v mt s cht ha tan i
qua nhng khng cho cc cht ha tan khc i qua.
Chc nng
Mng nguyn sinh cht ngn cch t bo vi mi trng: iu ny to cho t bo mt h
thng ring bit; qua mng, t bo trao i cc cht v thng tin vi mi trng ngoi
cn thit cho s sinh trng v pht trin ca t bo.

41/132

Cu to ca t bo Prokaryote
Hu ht Prokaryotes l sinh vt n bo. Mt s loi sng thnh nhm, tp on gm
nhiu t bo ging nhau. Mt s khc hnh thnh nhng t chc a bo c s phn cng
chc nng.
Prokaryotes c nhiu hnh dng khc nhau: hnh cu (cocci), hnh que (bacilli), hnh
phy v hnh xon (helice) bao gm xon khun (Spirilla) v Spirochettes
Kch thc t bo t 1-5 ?m. Tuy nhin Prokaryotes ln nht c hnh roi di 0.5 mm (t
bo Eukaryote 10-100 ?m).

Vch t bo
Hu ht prokaryotes c vch t bo bn ngoi mng sinh cht (plasma membrane) v
duy tr hnh dng ca t bo bo v t bo khng bi v khi t trong mi trng nhc
trng v c lysozyme. Tuy nhin mi trng c p sut thm thu qu cao, Prokaryote
s cht.
Khng ging nh vch t bo thc vt c cu to t cellulose, vch t bo vi khun
c cu to t peptidoglycan gm cc phn t polysaccharides lin kt ngang vi cc
phn t petides ngn. Thnh phn cu to ca vch l khc nhau gia cc loi. mt
s loi, vch t cn c thm lipopolysaccharides (carbonhydrate lin kt vi lipids)
y l c im gip phn bit vi khun khi nhum Gram (Gram stain) vi thuc
nhum tm tinh th (crystal violet). Vi khun Gram dng (Gram-positive bacteria) bt
mu s c mu ta, vch t bo ca loi ny phn ln l peptidoglycan khng c
lipopolysaccharides. Vi khun Gram m (Gram-negative bacteria) khng bt mu, vch
t bo c t peptydoglycan v c thm lipopolysaccharides.
Phn ln vi khun gy bnh l vi khun Gram m bi v lypolysaccharides gip vi khun
chng li lysozyme c trong tuyn nc bt v tuyn mi ca ngi v c kh nng
khng khng sinh bng cch ngn cn ng vo ca khng sinh. Lysozyme tc ng
trc tip ln vch t bo. Khng sinh chng hn penicillin ngn cn s hnh thnh lin
kt cho gia polysaccharides v polypetides. Nhiu vi khun gy bnh cn c lp v
nhy bn ngoi vch t bo gi l capsule tng cng kh nng chng khng ca t
bo vt ch v dnh c cht ca chng. Mc khc, capsule gip kt dnh cc t bo ca
vi khun hnh thnh khun lc.
Mt s loi c khun mao gip kt dnh t bo vo gi th, gi cc t bo dnh lu
truyn DNA trong sut qu trnh tip hp (conjugation).

42/132

V d:Neissenria gonorrhoeae gy bnh lu (goorrhoaeae) dng khun mao bm vo

nim mc vt ch.

Lng v roi
Hn phn na prokaryotes c kh nng chuyn ng nh hng do c roi. Cc roi tp
trung trn ton b c th hoc ch mt hoc hai u ca t bo. C ba c ch trong
chuyn ng ca vi khun. Th nht l nh roi. Th hai l nh hai hay nhiu khun
mao bn di vch t bo c cu trc tng t nh roi. Mi si c motor gn vo t
bo. Khi motor quay t bo chuyn ng theo c ch xon nt chai. C ch ny l c
bit hiu qu trong mi trng c nht cao. C ch th ba l mt s vi khun c th
tit ra cht nhy gy ra chuyn ng trt nh khi thiu khun mao. Trong mi trng
ng nht Prokaryote chuyn ng theo hng ngu nhin. Trong mi trng khng
ng nht, Prokaryote chuyn ng hng kch thch chng hn nh hng sng, hng
ngun thc n, trnh cht c. Mt s loi vi khun c cha mt s phn t nh nh
hp cht st cho php chng phn bit trn, di nh hng bt mi.
6.3 Mng sinh cht
Mng ca prokaryotes c cu trc khm lng ging nh mng ca Eukaryotes. S gp
np ca mng to nn nhng mng c chc nng ring bit chng hn nh thylakoids,
mesosome v mng h hp (respiration membranes)

Vt cht di truyn
Vt cht di truyn ca prokaryote l DNA mch n vng. Nhng vng DNA ln tng
tc vi proteins hnh thnh nhim sc th ca Prokaryotes hay gi genes tp trung mt
s khu vc nh trong t bo gi l vng nhn (khng c mng nhn). So vi Eukaryotes,
b gene ca Prokaryotes nh hn v n gin hn.
Trong b gene ca Eukaryotes, DNA mch thng tng tc vi cc proteins hnh thnh
b nhim sc th c trng cho loi. Trung bnh b gene ca mt prokaryote ch bng 1/
1000 DNA trong mt t bo Eukaryote.
Ngoi DNA ca nhim sc th, Prokaryotes cn c nhng vng DNA nh hn gi
l plasmids cha vi genes. Trong hu ht mi trng, Prokaryotes tn ti khng cn
plasmids bi v cc chc nng quan trng u m ho trong DNA nhim sc th. Tuy
nhin, cc genes ca plasmids c th gip prokaryotes sng trong mi trng c khng
sinh, cc cht dinh dng l. Plasmid nhn i c lp vi nhim sc th v chuyn cho
t bo khc gii khi tip hp (conjugation).

43/132

Cc qu trnh sinh hc trong t bo

Trao i cht qua mng t bo
Tnh thm ca lp phospholipids kp
ui k nc ca phospholipids ngn cn s vn chuyn cc ion v cc phn t phn
cc nh H+, Na+. Cc phn t khng tan trong nc nh hydrocarbon, CO2, O2 qua
mng d dng.
Nhng phn t nh phn cc nhng khng tch in nh ethanol cng c th qua mng.
Lp phospholipids kp khng thm i vi nhng phn t khng tch in nhng kch
thc ln chng hn nh glucose v sucrose.
Tuy nhin, lp phospholipids kp ch l mt phn cu chuyn v tnh thm c chn lc
ca mng.
Protein vn chuyn
Nhng protein vn chuyn l protein xuyn mng (hnh 2.40). Mt s protein vn
chuyn lm thnh cc chanel a nc (hnh 2.40). S khc lin kt vi cc cht v vn
chuyn chng qua mng. Trong c hai trng hp, mi protein vn chuyn chuyn bit
cho mt hay mt s c cht.
Nh vy, tnh thm ca mng ph thuc vo c hai lp lipid kp v cc protein vn
chuyn chuyn bit.
Khuch tn (diffusion)
Khuch tn l c ch vn chuyn qua mng mt cch th ng khng tn nng lng.
Cc cht c vn chuyn qua mng theo chiu t ni c nng cao n ni c nng
thp.
Thm thu (osmosis)
Thm thu l mt hnh thc vn chuyn nc mt cch th ng qua mng. Nc vn
chuyn t mi trng nhc trng sang mi trng u trng.
So snh nng cc cht ha tan ca hai dung dch. Dung dch c nng cao l
dng dch u trng (hypertonic), dung dch c nng thp l dung dch nhc trng

44/132

(hypotonic). Nu hai dung dch c nng bng nhau l cc dung dch ng trng
(isotonic).
Qu trnh thm thu ca nc qua mng t bo c th c minh ha qua s co nguyn
sinh (plasmolysis) ca t bo khi c trong nhng mi trng c nng khc nhau
(hnh 2.46)
Khuch tn c lm d
Mt s phn t phn cc hoc ion khng thm qua lp lipid kp c th c vn chuyn
qua mng nh cc protein vn chuyn. Hin tng ny gi l khuch tn c lm d.
Trong nhiu trng hp, protein thay di hnh dng vn chuyn v tr gn c cht t
bn ngoi mng sang bn trong mng.
S vn chuyn ch ng
S hp th ch ng cc cht ha tan vo t bo i hi phi tiu tn nng lng. S
vn chuyn ch ng l s vn chuyn vt cht ngc chiu gradient nng , phi tn
nng lng (ATP).
Nhp bo (endocytosis)
T bo ly cc cht c phn t lng ln vo t bo bng cch hnh thnh bng mng
t mng sinh cht. C ba hnh thc nhp bo: thc bo-phagocytosis (c cht l cht
rn); m bo-pinocytosis (c cht l cht lng) v nhp bo c lm d nh receptor
(receptor-mediated endocytosis).
Xut bo (exocytosis)
T bo tit cc cht c phn t lng ln qua mng t bo bng cch ha cc bng mng
vi mng.
Trao i thng tin qua mng.
Qua mng t bo c th pht i v thu nhn thng tin iu ha cc hot ng sng.
Thng tin dng nhng tn hiu ha hc c kh nng lin kt vi cc th quan trn
mng. S truyn tn hiu c th l:
Ni tit tc ng xa (endocrine transmission): cht ni tit v trong mu tc ng n
cc t bo khc nhau trong c th.
Cn tit (paracrine transmission): tc ng ln t bo k tip (khong cch xung quanh
khong 1mm) bng cc cht ha hc trung gian cc b (local chemical messenger)

45/132

S truyn qua synapses (synatic transmission).

Sinh trng v sinh sn ca t bo.

Sinh trng ca t bo
T bo con bng con ng trao i cht lm tng khi lng t bo cht v nhn. T
bo tng trng n mc no th phn chia cho cc t bo mi.
Sinh sn ca t bo
T bo prokaryote v mt s t bo eukaryote phn chia t bo bng hnh thc trc phn.
Trong trc phn ca prokaryote, DNA nhn i v dnh hai im khc nhau trn mng
sinh cht. Mng sinh cht mi v vch mi hnh thnh phn gia t bo chia t bo
m thnh hai t bo con. y khng c s xut hin nhim sc th cng nh thoi v
sc. Nhiu khi trong trc phn, nhn phn chia thnh hai na khng u nhau. T bo
cht phn chia cng vi nhn hoc khng phn chia hnh thnh t bo hai hay a nhn.
T bo gan, sn, bch cu phn bo theo kiu trc phn.
Phn bo c t l hnh thc phn bo ph bin t bo eukaryote gm hai giai on:
phn chia nhn c s hnh thnh thoi v sc, mt cu trc gip cho s phn chia u
nhim sc th v hai nhn; phn chia t bo cht.
S phn chia t bo cht t bo ng vt v thc vt c s khc bit. t bo ng
vt, mt rnh phn chia xut hin trn b mt t bo gn mt phng xch o, sau n
su vo do tc ng co rt ca vng vi si actin bn trong t bo cht ct t bo m thnh
hai t bo con ging nhau (daughter cells). Trong khi t bo thc vt, khng xut hin
rnh phn ct. Nhng bng mng (vesicle) cha nguyn liu xy dng vch t bo t
Golgi di chuyn ti gia t bo v t chc thnh a t bo (cell plate) chia t bo m
thnh hai t bo con (daughter cell).

Bit ha t bo (differentiation)
Bit ha t bo l mt qu trnh m mt t bo cha chuyn ha (undifferentiated cell)
tr nn chuyn ha thnh mt hay nhiu t bo cu trc nn c th. Trong sut qu trnh
bit ha, mt s gene c hot ha mt s gene khc b bt hot. Kt qu l t bo bit
ha c cu trc v chc nng c trng .

46/132

S a dng ca t bo
S a dng hnh thi ca t bo
Hnh dng t bo
Mt s t bo c hnh dng nht nh: t bo tinh trng, Paramescium, t bo thc vt,
t bo hng cu.
Mt s t bo lun thay i hnh dng: amip, bch cu.
Hnh dng t bo ph thuc vo chc nng, sc cng b mt, tc ng ca t bo bn
cnh cng nh tnh cht ca nguyn sinh cht.
Trong mi trng lng t bo c hnh cu: t bo bch cu trong mu c hnh cu.
Nhng khi ra khi mch mu, bch cu th chn gi v bin i hnh dng.
T bo ng thc vt c hnh khi do tng tc vi cc t bo trong m.
T bo thn kinh c hnh sao v phn nhnh ph hp vi chc nng dn truyn thng
tin. T bo c trn c hnh thoi ph hp vi chc nng co rt.
Kch thc ca t bo
Kch thc t bo rt a dng thng t 3-30m.
C nhng t bo rt to nh t bo trng chng hn nh trng iu c ng knh
khong 17 cm.
C nhng t bo rt nh nh Mycoplasma laidlawii ch khong 0.1m bng 10 ln b
hn t bo vi khun, 100 ln b hn t bo ng vt v 1000 ln b hn t bo amip.

Chc nng
T bo ng vt a dng v chc nng v c th c chia thnh 3 nhm: t bo sinh
sn (germ cell), t bo soma (somatic cell)-cu trc c th, t bo gc cha bit ha
(stem cell)

Phn bit t bo prokaryote v eukaryote

Tnh cht

prokaryote

Eukaryote
47/132

T chc vt cht di
truynMng
nhnHistonS lng
nhim sc thInstronTh
nhnPhn bo nguyn
nhim Ti t hp vt cht
di truynTi thLp
thMng c
sterolRoiMng li ni
chtGolgiVch t boS
khc bit trong cc bo
quan n
ginRibosomeLysosome
v
peroxisomeMicrotubuleB
xngBit ha

Khng KhngMt +
plasmidHimKhngKhngChuyn
mt phn theo mt
hngKhngKhngCh c
Mycoplasma v methanotrophNh
hn, mt si (one
fiber)KhngKhngPeptidoglycan
(tr Mycoplasma v
Archeaobacteria)70SKhngKhng
hoc rt himKhngKhng

ccnhiu hn mtph
binccPhn bo gim
nhim v th
tinhCCCLn hn, c
mng bao, 20 microtubule.
9+2C
CCellulose80SCCCM,
c quan

Phn bit t bo ng vt v thc vt

Tnh cht

T bo thc vt

T bo ng vt

Vch t boLc
lpCht d
trPhn
boKhng
boCentriole

CCTinh btPhn bo khng

c sao, phn t bo cht bng
vch ngn trung tmPht
trinKhng

KhngKhngGlycogenPhn bo
c xut hin sao, phn t bo cht
bng eo tht trung tmKhng
pht trinc

48/132

Khi nim v cu trc Enzyme

Khi nim Enzyme
Trao i cht hay chuyn ha vt cht l ton b cc ha trnh ha hc xy ra trong t
bo. Phn ln cc phn ng xy ra trong t bo c iu ha bi cht xc tc c bit
gi l emzyme.
Enzyme l cht xc tc sinh hc c cc tnh cht sau:
(1) Enzyme khng gy ra phn ng m lm gim nng lng hot ha (activation
energy) ca cc phn ng ha hc, do lm tng tc ca cc phn ng. iu ny l
bi v khi enzyme lin kt vi cc c cht to thnh phc hp enzyme-c cht lm cu
trc ni ti ca c cht thay i theo chiu hng gim nng lng hot ha.
(2) Enzyme khng b b bin i hay mt i v c th s dng nhiu ln trong cc phn
ng ca t bo. Sau phn ng enzyme c gii phng.
(3) Ch cn mt lng nh enzyme so vi c cht cng xc tc c phn ng.
(4) Enzyme c tnh c hiu cao. Tnh c hiu ca enzyme th hin cc cp khc
nhau. Mi ezyme ch xc tc cho mt kiu phn ng nht nh, mt c cht nht nh
hay mt loi ng phn nht nh ca c cht .
(5) Khng phi mi t bo cng cha mt h enzyme. iu gii thch ti sao c nhiu
hn mt loi t bo

Cu trc ca Enzyme
Bn cht ha hc ca enzyme l protein hnh cu c th chia enzyme thnh hai nhm
ln sau:
Enzyme n gin-enzyme mt thnh phn ngha l khi thy phn ch cho mt thnh
phn duy nht l amino acid. Hot tnh ca enzyme ny ch ph thuc vo cu trc ca
protein.
Enzyme phc tp: Ngoi thnh phn protein gi l apoezyme, enzyme cn cha thm
nhm khng phi l protein gi l cofactors. Nhm ngoi ny rt a dng. C th phn
bit ba nhm ngoi c bn:

49/132

(1) Nu nhm ngoi l cc phn t hu c c trng lng phn t thp th gi l

coenzyme, thng bt ngun t vitamin ch khng t amino acid. Coenzyme lin kt
thun nghch vi enzyme, tham gia trc tip trong cc phn ng m enzyme xc tc.
V d: NAD v FAD l hai coenzyme nh cht mang in t v H cho nhiu phn ng
oxi ha kh c enzyme xc tc trong t bo.
(2) Nhm ph gia (prosthetic group) tng t nh coezyme nhng gn cht vi protein
enzyme bng lin kt ha tr chng hn nh enzyme catalase. Ging nh coenzyme
nhm prosthetic thng l thnh phn c bn trong c ch xc tc ca enzyme.
(3) Nhm ngoi l cc ion kim loi nh Fe, Mn, Cu, Ca, Zn. Cc ion c vai tr n nh
cu trc ca enzyme v c cht hoc c th tham gia trc tip phn ng chng hn nh
cytochrome cha Fe tham gia vo chui chuyn in t.
Mt s enzyme i hi c hai thnh phn c hot tnh xc tc. Nhm ngoi thng
n nh nhit trong khi protein enzyme b bin tnh khi b un nng. Phc h enzymecofactor c hot tnh xc tc gi l holoenzyme. Khi cofactor b loi b th phn protein
(apoenzyme) cn li mt hot tnh xc tc.
Trn b mt ca cc enzyme c mt hoc mt s khe, ti- ch lm gi l v tr hot ng
(active site) ni xy ra hot ng xc tc ca enzyme.

50/132

C ch hot ng,phn loi v cc yu t

nh hng n hot tnh Enzyme
C ch hot ng ca Enzyme
M hnh hot ng ca enzyme
Nhin cu u tin v tnh c hiu ca enzyme vi c cht do Emil Fischer xng
(1894). Fischer cho rng khi enzyme xc tc phn ng th c cht lp kht vo v tr hot
n ca enzyme tng t nh cha kha lp vo kha. T Fischer nu gi thuyt m
hnh kha v cha kha (lock and key model) (hnh 3.1) cho hot ng ca enzyme.
Mi kha c mt cha kha. Do mi enzyme ch c mt ch xc tc cho mt loi c
cht nht nh.
Theo m hnh ny, c ch xc tc ca enzyme tri qua nhiu giai on: Giai on u
tin l s kt hp gia enzyme v c cht to thnh phc hp enzyme-c cht nh hnh
thnh nhiu lin kt c bit l lin kt hydrogen. S lin kt ny lm thay i cu hnh
khng gian ca c cht lm thay i ni nng, nng lng hot ha ca phn ng gim,
phn t tr nn linh ng d phn ng hn. Sau enzyme xc tc ln c cht to thnh
sn phm. cui phn ng enzyme c gii phng.
Tuy nhin, khi Daniel Koshland nghin cu ng hc ca enzyme, nhng bng chng
thc nghim cho thy rng cu hnh trung tm hot ng ca enzyme khng phi c
nh v c th bin i ph hp vi c cht khi lin kt vi c cht . T y, Daniel
Koshland a ra m hnh khp cm ng (induced-fit model) cho hot ng xc tc ca
enzyme (hnh 3.2). Theo m hnh ny th mt enzyme c th xc tc cho mt hoc mt
s phn ng ha hc tng t do chng c tnh c hiu trong vic la chn c cht. V
tr hot ng ca mi enzyme khc nhau c hnh dng sao cho ch khp vi mt hoc
mt s c cht nht nh. M hnh ny hin nay c chp nhn rng ri.
Tnh c hiu ca enzyme
Khc vi cht xc tc v c, enzyme ch c kh nng xc tc cho mt mt hay mt s
c cht hoc mt kiu phn ng nht nh. Tnh c hiu ca enzyme rt a dng. y
ch cp n hai kiu c hiu:
(1) c hiu cho phn ng
Mi enzyme ch xc tc cho mt kiu phn ng chng hn nh phn ng ca NAD
dehydrogenase trong h hp, lipase ct lin kt ester ni glycerol v acid bo ca nhiu
loi lipids.
51/132

(2) c hiu c cht

(a) c hiu tuyt i: enzyme ch c tc dng ln mt c cht nht nh chng hn nh
aspartase chuyn fumarate thnh L-aspartate.
(b) c hiu tng i: enzyme co th tc ng ln nhiu c cht c cu trc khc nhau
nhng tc phn ng khc nhau chng hn nh phosphatase thy phn nhiu ester ca
acid phosphoric, carboxyesterase thy phn ester ca cc acid carboxylic.

Phn loi Enzyme

Cc enzyme thng c tn tn cng l ase. C su nhm enzyme chnh:
Nhm 1. Oxidoreductases: xc tc cc phn ng oxi ha kh, chuyn H v in t t c
cht sang cht nhn.
Nhm 2. Transferases: chuyn nhm nh cc nghuyn t t c cht sang cht nhn.
Nhm 3. Hydrolases: Lm gy cc lin kt bng thy phn.
Nhm 4. Lyases:lm thay i chiu di mch carbon. Fructose-1,6-diphosphate aldolase
ct Fructose-1,6-diphosphate thnh ALPG v PDA; Decarboxylase: lai CO2
Nhm 5. Isomerases: chuyn i ng phn chng hn Glocose-P-isomerase chuyn
Glo-6-P thnh Fro.-6-P); triose-P-isomerase chuyn ALPG thnh PDA)
Nhm6. Ligases: to lin kt ha hc chn hn DNA ligase gn cc on okazaki trong
nhn i DNA mch chm.

Cc yu t nh hng n hot tnh Enzyme

Bt k iu kin no lm thay i cu hnh ca enzyme u lm thay i hot tnh
enzyme
Nng enzyme v nng c cht
Tc phn ng ca phn ln cc phn ng bin i theo nng c ca c cht v nng
enzyme. Khi tng nng c cht th tc phn ng tng ch khi nng c cht
tng i thp. Khi nng c cht ln tc phn ng t ph thuc vo nng c
cht v c khuynh hng t cc i do nng enzyme c mt quyt nh.
nng c cht thp, nhiu phn t enzyme c trung tm hot ng cha lin kt vi
c cht. Nn vic tng hn ch c cht l tng tc phn ng. Tuy nhin, nng

52/132

c cht cao, hu ht cc trung tm phn ng lin kt vi c cht lm cho s phn t

enzyme tr thnh yu t gii hn. Khi s phn t ezyme tng tc phn ng c i
tng ln tng ng.
Cht km hm
Hot tnh ca enzyme b thay ikhi c mt ca cht c ch.
(a) Cht c ch cnh tranh
Cc cht km hm c cu trc tng t nh c cht. Cht c ch gn thun nghch vo
trung tm phn ng ca enzyme cnh tranh vi c cht, khin cho hot ng xc tc ca
enzyme chm li. Khi cht c ch c gii phng hot ng xc tc ca enzyme tr li
mnh thng.
V d: acid malonic hot ng nh cht cnh tranh trung tm hot ng ca enzyme
sucxinic dehydrogenase vi acid sucxinic. Trong iu kin bnh thng, sucxinic
dehydrogenase xc tc loi hai nguyn t H ca acid sucxinic to thnh acid fumaric.
Khi c mt ca acid maloic, enzyme lin kt vi trung tm hot ng to thnh phc
hp enzyme cht c ch v khng th chuyn ha c. Nh vy lm s lng trung
tm phn ng gim v tc phn ng cng gim theo.
(b) Cht c ch khng cnh tranh
Cht c ch khng cnh tranh khng gn vo v tr xc tc m gn thun nghch vo v
tr khc trn enzyme. iu ny s lm thay i cu hnh ca v tr hot ng khng cn
ph hp vi c cht. Vng m cht c ch cnh tranh gn vo enzyme gi l v tr di lp
th (allosteric site) v gy ra hiu ng d lp th (allosteric effect). Khi cht c ch gii
phng, hot ng xc tc ca enzyme tr li bnh thng.
Mt s cht km hm gy bin i hot tnh enzyme mt cch khng thun nghch. Cht
km hm dng ny lm bin i lin tc hay ph hy trung tm hot ng ca enzyme.
Phn ln cc cht c l nhng cht km hm khng thun nghch chng hnh nh
cyanide (CN-) gn vo cytochrome cn tr vic vn chuyn in t n O2, penicillin
c ch khng thun nghch enzyme transpeptidase ca vi khun cn tr to vch t bo.
Nhit
Mi mt enzyme hot ng tt nht nhit ti u nht nh. Khi nhit lch sang
hai bn nhit ti u hot ng ca enzyme gim v tc phn ng s gim theo.
S tng nhit lm cho lm cho ng nng ca enzyme v c cht tng, chng chuyn
ng nhanh hn, va chm nhiu hn. Cc phc cht emzyme-c cht hnh thnh nhiu
hn, phn ng xy ra nhanh hn. Tuy nhin nu nhit qu cao, enzyme b bin tnh.
Khi cu hnh v tr xc tc khng cn ph hp vi c cht, enzyme mt hot tnh xc
53/132

tc. Khi nhit h thp hn nhit ti u, c cht v phn t enzyme chuyn ng

chm. Tn s va chm gia chng thp> t phc hp enzyme-c cht hnh thnh v tc
phn ng gim.
pH
Mi mt enzyme hot ng ti u ti mt gii hn pH thch hp chng hn pepsin hot
ng ti u pH 2, trypsin hot ng ti u pH 8.5. Khi pH lch sang hai bn pha
pH ti thch, hot tnh ca enzyme gim xung.

54/132

H hp t bo
Khi nim h hp t bo
Cc hp cht hu c cha nng lng trong s sp xp cc nguyn t ca n. Vi s
xc tc ca cc enzyme, t bo phn gii cc hp cht hu c phc tp giu nng lng
thnh nhng cht n gin. Mt phn nng lng gii phng ra c s dng vo hot
ng ca t bo, phn cn li c gii phng di dng nhit. Qu trnh ny gi l
d ha. Mt qu trnh d ha phn gii mt phn nguyn nhin liu trong s vng mt
ca oxygen gi l ln men (fermentation) hay h hp ym kh (anarobic repiration). Con
ng d ha hiu qu v ph bin nht l h hp trong qu trnh ny c s tham gia ca
oxygen hay hiu kh (arobic repiration).
H hp l qu trnh phn gii cc hp cht hu c thnh cc sn phm v c cui cng
ngho nng lng nh CO2 v nc ng thi gii phng ra mt ngun nng lng ln.
C6H12O6 + O2 = 6CO2+ 6H2O + 674 kcal/mol glucose
Thc cht, h hp l mt h thng oxy ha kh phc tp trong xy ra cc phn ng
oxy ha kh: tch in t v H t nguyn liu h hp, chuyn ti oxy khng kh di
tc dng ca h enzyme. Nng lng gii phng t h hp c c nh trong cc cht
giu nng lng chng hn nh ATP.

Mi quan h gia h hp v ln men:

Nhiu nh sinh l hc ni ting th gii th k 19, u th k 20 cho rng: H hp l
s k tc ca ln men.
C6H12O6 = 2C2H5OH + 2CO2 (Ln men)
2C2H5OH + O2 = 4CO2 + 6H2O

Giai on chun b ca h hp v ln men (ng phn: glycolysis):

Glycolysis gm mt lot (10 phn ng) cc phn ng khng cn O2, xy ra nh mt
nhm enzyme trong cytosol.
c im ca c cht trong glycolysis l dng ng c hot ha thnh dng ester.
y l dng ng linh ng d dng tham gia phn ng. Trong glycosis, glucose c
xem l c cht v pyruvate l sn phm cui cng ca glycolysis.

55/132

Glycolysis gm hai phase:

*Pha khi u (initial phase):
C cht t nhng ngun khc nhau c chuyn thnh cc triose phosphate. Qu trnh
ny cn 2ATP
*Giai on bo tn nng lng (energy-conseving phase or energy pay-off phase)
Cc triose phosphate tri qua hng lot cc phn ng (5 phn ng) hnh thnh sn
phm pyruvate. NAD+ b kh thnh NADH bi glyceraldehyde-3 phosphate. ATP c
tng hp trong phn ng giai on ny c xc tc bi phosphoglycerate kinase,
pyruvate kinase.
Glycolysis ca mt phn t glucose cn 2 ATP, to 4 ATP, 2 NAD(P)H v 2 pyruvate.

H hp hiu kh (aerobic respiration)- crebs cycle citric acid cycle:

Trong sut th k 19, ngi ta bit rng trong s c mt ca O2, t bo to CO2 v H2O.
Krebs (1937) cng b pht hin chu trnh citric acid hay cn gi chu trnh Krebs.
Chu trnh ny gii thch c ch oxy ha pyruvate thnh CO2, H2O, NAD(P)H, FADH
v ATP v xy ra trong matrix ca ti th.
Chu trnh Krebs
u tin pyruvate to ra t glycolysis cytosol c vn chuyn qua mng trong ti th
v sau c oxy ha qua hng lot cc phn ng c tm tt trong hnh 3.10 .
-Oxy ha mt phn t pyruvate to ra: 4 NAD(P)H, 1 FADH, 1 ATP, CO2 v H2O.
ngha ca chu trnh Krebs
T suy lun

H hp ym kh (anaerobic respiration-fermentation):
Trong iu kin ym kh, chu trnh Krebs v phosphoryl ha khng th thc hin.
Glycolysis khng th xy ra bi v ngun NAD+. khc phc iu ny, pyruvate to
ra trong glycolysis c oxy ha bi NADH ti to NAD+. Qu trnh ny gi l qu
trnh ln men. C hai hnh thc ln men quan trng: ln men ru v ln men lactic.

56/132

Hiu qu nng lng trong ln men thp bi v hu ht nng lng trong glucose vn
cn lu tr trong cc sn phm ca qu trnh ln men chng hn nh ethnol, lactic acid.
Trong ln men ru, pyruvate bit i thnh ethanol theo hai bc: pyruvate gii phng
CO2 v hnh thnh acetaldehyde, sau b kh bi NADH thnh ethanol. Qu trnh ny
ti to NAD+. Hnh thc ln men ny xy ra nm men, vi khun
Trong ln men lactic acid, pyruvate b kh bi NADH thnh lactate. ngi, qu trnh
ln men ny xy ra t bo c lc chi th thao cng thng bi v thiu ngun oxy.
Lactate tch ly gy mi v au c. Sau , lactate theo mu n gan v bit i thnh
pyruvate.

Chui chuyn in t v qu trnh phosphoryl oxyha (electron transport

and oxidative phosphorylation)
Nh cp trn, glycolysis v chu trnh Krebs hnh thnh 4 ATP trn mt phn
t glucose bi phosphoryl ha mc c cht. Cc phn t NADH v FADH2 qua chui
chuyn in t to nn ng lc tng hp 34 ATP. Qu trnh ny gi l phosphoryl
oxygen ha.
Phosphoryl mc c cht
Nng lng to ra khi tch H t c cht c s dng tng hp ATP.
Trong giai on ng phn

Phosphoryl oxy ha
Con ng vn chuyn in t.

57/132

in t c tch ra t nhin liu trong glycolysis v chu trnh Krebs c NADH

chuyn cho mt flavoprotein l FAD (flavin mononucleotide) sau in t n ionsulfur protein (Fe.S) ubiquinone (Q) bn cht l lipid. Hu ht cc cht mang in
t c li gia Q v oxygen l cytochrom (cyt) nhm prothestic ca chng gi l hem c
bn vng hu c bao quanh nguyn t Fe, tng t nh hemoglobin nhng Fe ca cyt
tham gia chuyn in t ch khng chuyn oxygen. in t c chuyn qua nhiu loi
cyt v cyt cui cng l cyt a3. in t t cyt a3 sau c chuyn cho H2O.
Mt ngun in t khc l t FADH2 c to bi chu trnh Krebs. FADH2 chuyn in
t ca n vo chui in t mc nng lng thp hn NADH. Chnh v vy, nng
lng m FADH2 cung cp cho tng hp ATP ch bng hai phn ba so vi NADH.
Chui chuyn in t khng trc tip tng hp ATP. Vai tr chnh ca n l chuyn
in t t nhin liu n oxygen.
Ha thm thu (chemiosmosis): c ch tng hp ATP
Mng trong ca ti th tn ti mt phc hp protein l ATP synthase, enzyme tng hp
protein hot ng ngc li vi bm ATP. Bm ATP dng nng lng (ATP) vn
chuyn ion ngc chiu nng , ngc li ATP synthase dng nng lng do s chnh
lch nng ion (prton H+) gia hai bn mng trong ca ti th tng hp ATP.
S chnh lch nng gia hai bn mng trong ti th l bi v khi in t qua chui in
t gii phng nng lng bm H+ t matrix vo khong gia hai mng. H+ c xu hng
i mng gim s chnh lch ny. Mng trong khng thm i vi H+ nhng ATP
th thm. S chnh lch H+ to lc proton (proton-motive force) qua ATP synthase.
Nh vy, gradient H+ i km vi cc phn ng oxy ha kh chui in t tng hp
ATP. C ch cp i dn n s phosphoryl oxy ha ny c gi l ha thm thu.
Lm th no chui in t bm H+ vo trong khong gia hai mng? Cc nh nghin
cu tm thy rng mt s thnh vin ca chui in t nhn v gii phng H+ cng vi
in t, ngc li mt s khc ch chuyn in t. Nh vy, ti mt v tr no trn
chui in t, mt s H+ c nhn sau tr li mi trung xung quanh; s khc c
chuyn t matrix vo trong khong gia hai mng.
Lm th no ATP synthase dng dng i vo matrix ca H+ tng hp ATP? Phc
hp ATP synthase gm c ba thnh phn: mt ng hnh tr (cylinder) xuyn mng trong,
mt knob quay vo matrix v mt trc gia ni ng tr v knob. Khi H+ i vo ng hnh
tr lm cho ng hnh tr v trc quay. iu ny lm knob thay i hnh dng, hot ha
v tr xc tc ca ATP synthase, xc tc phn ng ADP v H3PO4 to ATP.

58/132

Hiu xut oxy ha hon ton mt phn t glucose

Giai on ng phn to: 2 ATP v 2 NADPH2 s to 6 ATP khi qua chui in t.
Nh vy giai on ng phn to 8 ATP
Giai on hiu kh to:2 NADPH2 ( t pyruvate) s to 6 ATP, 24 ATP trong chu trnh
Krebs. Nh vy, giai on hiu kh to 30 ATP
-Tng ATP to ra khi phn gii 1 phn t glucose l 38 ATP
H=

3810
674

100 =56

(1 ATP ~ 7.3 kcal; 1 glucose ~ 686 kcal)

Vai tr ATP trong hot ng ca t bo

- Cung cp nng lng cho vn ng ca t bo: co c, vn ng ca nhim sc th v
cc cc.
- Cung cp nng lng cho t bo vt chuyn cc cht: bm cc cht qua mng.
- Hot ha cc cht trong nhiu phn ng xy ra trong t bo.

59/132

Quang hp
Khi nim v quang hp:
AS, dip lc
6CO2 + 6H2O ------------------- > C6H12O6 + 6O2
Quang hp l qu trnh bin i nng lng nh sng mt tri thnh nng lng ha hc
trong di dng cc hp cht hu c.
Quang hp l qu trnh bin i cc cht v c n gin thnh cc cht hu c phc tp
c hot tnh cao trong c th thc vt, ng thi gii phng O2 nh nng lng nh sng
mt tri v h sc t.

ngha ca quang hp:

ng gi nng lng nh sng mt tri di dng nng lng ha hc cn thit cho sinh
vt.
Nng lng mt tri ri xung tri t 5.10 23 Kcal, 2.10 23 xung vng c cy ci.
Cy xanh chuyn 2% (5.10 21 Kcal) nng lng ny thnh nng lng ha hc.
Quang hp l ngun cung cp ch yu cc cht hu c trn tri t cho hot ng sng
ca sinh vt v loi ngi.
Quang hp to 4,5.10 11 tn cht hu c (trn cn 5.10 10 tn con gni s dng 3,5%,
di nc 4.10 11 tn con ngi s dng 2.10 -5 )
Lm sch khng kh.
Quang hp ng ha 170 t tn CO 2 gii phng 115 t tn O 2 .

L c quan lm nhim v quang hp

-Hnh thi:
Thng c dng bn mang c tnh hng r rt, c kh nng vn ng nhn nhiu
nht nng lng mt tri cng nh gim s t nng khi cng sng mnh.
-Cu trc:

60/132

Lp cutin (Cuticle): chng thm nc

Biu b (epidermis): c vai tr bo v l v gim thot hi nc kh khng (stoma)trn
lp biu b iu ha thot hi nc v trao i kh. (CO2)
M du (palisade mesophyll cell): Nm st lp biu b. Cc t bo m du xp st nhau
gn nh song song nhm hp thu nng lng nh sng cao nht. T bo m du c nhiu
lc lp, thc hin chc nng quang hp.
M khuyt (m xp: spongy mesophyll cell) : Gia t bo c nhiu khong trng gi l
gian bo, ni cha CO2, hi nc cung cp cho quang hp. Ngoi ra, t bo m khuyt
cn cha lc lp nhng t hn t bo m du.
Mch dn (vein) gm mch g (xylem), mch ry (phloem), t bo vng bao b mch
(bundle sheath cell): Dn nc v khong phc v cho quang hp, thc hin quang hp
v dn sn phm quang hp ra khi l.

Lc lp (chloroplast)-bo quan thc hin quang hp

-Hnh dng
Thc vt khng b nh sng t trc tip, lc lp c hnh cc hnh sao, hnh bn, hnh
chung
Thc vt trn cn, lc lp c hnh bu dc. Vi hnh bu dc, lc lp c xoay b mt
iu chnh mc tip xc vi nh sng v s s dng nh sng hiu qu nht. y
cng l mt c im tin ha ca gii thc vt.
-S lng, kch thc
S lng lc lp khc nhau cc loi thc vt. Mi t bo to c mt lc lp. Thc vt
bc cao trung bnh 20-100 lc lp.
ng knh lc lp khong 4-6 micrometers, dy 2-3 micrometers
Nhng cy a bng c s lng, kch thc lc lp v hm lng sc t trong lc lp
nhiu hn cy a sng.
-Cu trc
*Mng:
Lc lp c bao bc bi mng kp, c cu trc ging nh mng c bn.

61/132

*Stroma:
L ni xy ra cc phn ng ca pha ti trong quang hp. Thnh phn ha hc ca stroma
bao gm: Cc enzyme, protein, acid nucleic, ribosome, lipid v cc git du...
Lc lp c kh nng tng hp DNA v protein cho chng. Mt s protein khc c
tng hp cytoplast v c vn chuyn ti lc lp.
*Grana:
Grana l tp hp cc thylakoid xp chng cht ln nhau v l ni xy ra cc phn ng
pha sng trong quang hp. Mt grana c 5-30 thylakoid c gi l thylakoid grana.
Mi lc lp c khong 40-50 grana c lin kt vi nhau bi thylakoid stroma.
*Cu trc ca thylakoid:
Thylakoid gm mng c bn, gia cc lp ca mng l h sc t v enzyme thc hin
cc phn ng ca pha ti v sng. S sp xp cc thnh phn trong thylakoid m bo
thc hin cc phn ng khc nhau trong quang hp.

H sc t
Chlorophyll, carrotenoid, phycobilin v anthocyan.
Chlorophyll
Bao gm chlorophyll a, b (c,d,e c trong vi sinh, rong v to)
Cng thc phn t ca chlorophyll a: C55H72O5N4Mg v b: C55H70O6N4Mg.
Cng thc cu to: Gm 4 vng pyrol ni vi nhau bi cu ni methyl (-CH=) to nn
cu trc porphyrin vi Mg gia. Vng pyrol s 4 gn nhm phytol v vng pyrol th
3 c vng cyclopental.
H thng ni i, n cch u trong cu trc l cho dip lc c hot tnh quang ha
mnh. Nng lng lin kt ni i nh, in t hnh thnh ni i d bc ra khi nhn
nng lng nh sng. ui phytol gm gc ru phytol c 20 carbon, a lipid c vai tr
nh v phn t dip lc trn thylakoid.
*Tnh cht vt l:
Chlorophyll khng tan trong nc nhng tan trong dung mi hu c. Chlorophyll hp
th quang ph nh sng trong khong 400-700 nm vi cc i hp thu 430 nm v 662

62/132

nm. Trong l cy, dip lc kt hp vi cc protein nn c cc i hp th sai khc t

nhiu; 700, 680, 685
Hin tng hunh quang ca chlorophyll: L hin tng pht ra nh sng khc c
di sng di hn di sng ca nh sng m chlorophyll hp th. y l trng thi
kch thch s cp ca dip lc trng thi singlet.
Hin tng ln quang: Cng tng t nh hunh quang nhng khi tt ngun sng mu
huyt d cn lu li mt thi gian ngn na. y l trng thi kch thch th cp, trng
thi triplet ca dip lc.
*Ha tnh:
Chlorophyll tc dng vi base to chlorophyllate c mu xanh.
Tc dng vi acid to pheophytin c mu nu do chlorophyll b kh mt Mg.
Phyophytin khng c kh nng hunh quang. iu cho thy Mg quyt nh tnh cht
ca dip lc. Phyophytin tc dng vi kim loi khc (Cu) to hp cht c mu xanh.
Dip lc c lp mt mu khi tip xc vi nh sng v c mt ca oxygen.
Chl + H ---------- > Chl*
Chl* + O2 ------------ > ChlO2 (mt mu xanh)
*Vai tr:
-Hp thu nng lng nh sng mt tri.
-Di tr nng lng vo trung tm phn ng.
-Tham gia bin i nng lng nh sng thnh nng lng ha hc ti cc trung tm
phn ng.
Carrotenoid:
Carrotenoid l nhm sc t c mu vng, cam, tm . Kh nng hp th nh sng ca
carrotenoid do cc ni i lin kt i n quyt nh crrotenoid s cp tham gia vo
quang hp, carroten th cp: trong qu, hoa, c quan gia khng tham gia quang hp.
*Carroten (C40H56):

63/132

Carroten c cu to t cc gc isopren, khng tan trong nc, tan trong dung mi

hu c, hp th nh sng c bc sng t 446 n 476 nm. Carroten c xem l tin
vitamin A.
*Xanthophyll (C40H56On vi n=1...6)
Xanthophyll hp th nh sng c bc sng trong khong 451 n 481nm.
*Vai tr ca carrotenoid
Nhn v truyn nng lng nh sng cho chlorophyll.
Bo v chlorophyll khi b phn hy cng nh sng cao.
Xanthophyll tham gia vo cc phn ng phn ly nc.
Cc protein tham gia chui in t
Pheophytine (Pheo), Plastoquinon A, B (QA, QB), Cytochrome b6f [Cytb, Cytf, FeSR
(Riesk-ion-sulfur proteins)], plastocyanin (QC), Cc protein cha lu hunh v sc
(FeSx, FeSA, FeSB), ferredoxin (Fd), Flavinprotein-NADP reductase (FNR)

C ch ca quang hp
Quang hp c hai pha: pha sng v pha ti. Pha sng xy ra thylakoid v pha ti xy
ra stroma.
Bn cht ca pha sng:
Bn cht ca nh sng:
Anh sng ca mt tri c tnh ht (photon) v sng.
E=h? =hc/
H: hng s planck =6,625 10-34js
: di sng nh sng
?: Tn s pht x
C: Vn tc nh sng

64/132

E: nng lng ca 1 photon.

E c th lm thay i trng thi bnh thng ca vt cht thnh trng thi kch ng
khng bn. Vt cht trng thi khng bn c xu hng tr v trng thi ban u bng
cc cch sau y: Pht nhit pht x hunh quang, truyn nng lng cho phn t ln
cn v truyn in t cho phn t khc.
Giai on quang l:
Chlorophyll hp th nng lng v tr thnh dng kch ng. Ty vo mc nng lng
E, chlorophyll c th tr thnh:
Singlet 2 c i sng 10-13-10-12 s
Singlet 1 c i sng t 10-9-10-8 s
Triplet c i sng 10-3-10-2 s
Singlet c th mt nng lng tr thnh triplet. Trng thi triplet ca chlorophyll c
vai tr quan trng trong vn chuyn in t v H kh CO2.
Cc sc t ph cng nhn nng lng nh sng v truyn cho chlorophyll.
Tm li: Trong giai on quang l, chlorophyll hp th nng lng nh sng v tr thnh
dng giu nng lng sn sng tham gia vo cc phn ng sau ny.
Quang ha:
Chlorophyll s dng nng lng hp th vo cc phn ng quang ha to cht d tr
nng lng v cc cht kh.
thu nhn nng lng nh sng, cc sc t t chc thnh trung tm gi l PSI trung
tm phn ng l P700 v PSII c trung tm phn ng l P680. Phn ng sng ti PSI v
PSII v con ng vn chuyn in t khng vng to O2, NADPH v ATP. P680 ti
trung tm PSII b kch ng khi hp th tia sng c bc sng 680 nm chuyn in t
cho Pheo, in t t Pheo sau c chuyn n QA, QB, phc hp cytob6f, PC v
n P700 ca trung tm PSI.
P700 ti trung tm PSI b kch ng khi hp th tia sng c bc sng 700nm chuyn
in t cho A0 (mt chlorophyll) sau n A1 (quinone), FeSX, FeSA, FeSB, Fd. Fd
trng thi kh cng vi FNR kh NADP+ trong stroma thnh NADPH.

65/132

Bng ng v phng x, cc nghin cu cho rng O2 thot ra t quang hp c ngun gc

t H2O.
Phn ng sng ti PSI v con ng vn chuyn in t vng to ATP nhng khng to
NADPH.
Thay v in t t P700 chyn n Fd kh NADP+ li chuyn n phc hp Cyto
b6f ri n PC v v P700. Trong con ng vn chuyn in t ny km theo tng hp
ATP.
S tng hp ATP
Qu trnh hnh thnh ATP do tc ng ca nh sng gi l qu trnh quang phosphoryl
ha. C hai kiu quang phosphoryl ha.: Phosphoryl ha vng: i km vi con ng
vn chuyn e- vng (xy thc vt khi gp iu kin bt li) (1-2ATP.); Phosphoryl
ha khng vng: i km vi con ng vn chuyn in t khng vng (khong 4 ATP)
Cng vi s vn chuyn in t t PQH2 n PC, phc hp Cytob6f gii phng H+ vo
pha trong ca thylakoid. H+ ny v H+ t phn ng phn li nc lm tng nng
H+ pha trong hai mng thylakoid. H+ pha trong mng sau khuch tn qua ATpase,
enzyme tng hp ATP nm trn mng thylakoid ra stroma. iu ny lm gim nng
H+ trong khong gia hai mng thylakoid v ATP c hnh thnh stroma.
Tm li: Pha sng ca quang hp to ATP v NADPH. Hai cht ny s c s dng
trong pha ti.
Quang phn ly nc:
Sau khi phng thch in t bi tc ng ca nh sng, P680 tr thnh dng oxy ha
mnh oxy ha Yz v Yz+ oxy ha nc gii phng O2 v H+ vo trong hai mng
thylakoid.

66/132

Bn cht ca pha ti:

C hai c ch chnh trong qu trnh c nh CO2 thc vt. S khc nhau ca hai c
ch ny l hp cht u tin v cht nhn CO2 u tin.
Chu trnh C3 (Calvin v cng s, 1950):
To Chlorella nui trong mi trng C14. Sau nhng khong thi gian nht nh, c lp
to bng CH3OH, ph v t bo, c lp cht hu c v phn tch thnh phn ca chng
bng sc k. Kt qu phn tch cho thy rng: Sn phm u tin l 3, phosphoglyceric
acid (PGA), sau l cc triose, hexose... cui cng l cc pentose v CO2. Nh vy
chu trnh C3 gm 3 giai on:
*Carboxyl ha (Carboxylation):
Cht nhn CO2 u tin l ribulose-1,5 biphosphate. Di tc dng ca ribulose-1,5
biphosphate carboxylase, phn ng to thnh cht trung gian 2-carboxy-3-keto Darabinitol 1,5-biphosphate, sau l 2 phn t PGA.
*Giai on kh (reductive phrase):
PGA b kh bi NADPH v ATP thnh glyceraldehyde-3 phosphate. y l giai on
then cht ca chu trnh, bin quang nng thnh ha nng, gm hai giai on:
PGA b bin i thnh 1,3-biphosphate-glycerate nh enzyme
phosphoglycerate kinase.
1,3-biphosphate-glycerate b kh bi NADPH thnh glyceraldehyde-3
phosphate nh enzyme glyceraldehyde phosphate dehydrogenase.
96% PGA bin i thnh DHAP (dihydroxyaceton-3 phosphate). 1 phn t PGA kt
hp vi DHAP to 1 phn t fructose-1,6 phosphate.
*Ti to ribulose-1, 5 diphosphate:
Hai triose to 1 phn t fructose-1,6 diphosphate. Phn cn li tham gia ti to
ribulose-1,5 diphosphate.
Chu trnh C4 (Hatch Slack, 1965):
Chu trnh C4 xy ra mt s thc vt nhit i nh ma, c lng vc, rau dn... Trong
chu trnh , cc cy ny dng PEP (CH2=COP-COOH) phosphoenol pyruvate nh l
cht u tin nhn CO2 to cc cht hu c c 4 carbon nn gi l chu trnh C4.

67/132

V mt gii phu, l ca cy C4 c hai loi t bo ng ha CO2: T bo tht l

(mesophyll) v t bo vng bao b mch (bundle sheath cells). T bo vng bao b mch
ca thc vt C4 pht trin mnh. Quang hp thc vt C4 xy ra hai ni: t bo tht l
(mesophyll cells) v t bo vng bao b mch. t bo tht l xy ra s c nh CO2 to
malate, aspartate. t bo vng bao b mch xy ra cc phn ng tip theo ging nh
chu trnh C3.
Do quang hp ca thc vt C4 mnh hn, hiu qu hn, hn na thc vt C4 khng c
h hp sng, nng sut sinh hc cy C4 cao hn cy C3.
Chu trnh CAM (Crassulacian Acid Metabolism):
Con ng c nh CO2 ny xy ra thc vt sng vng kh hay nhim mn nh cy
da, xng rng... hn ch b mt nc, nhng cy ny m khng vo ban m v
ng khng vo ban ngy, lm thay i con ng c nh CO2.
Ban m, CO2 khuch tn vo trong t bo, c c nh thnh acid c 4C v c tr
khng bo cho n sng mai. Vo ban ngy, cc acid 4C b kh to CO2 khuch tn
vo chu trnh C3.

68/132

Lch s pht trin ca di truyn hc

Trong qu trnh sinh sn, con ci lun ging cha m mt mc nht nh. S sao
chp li cc tnh trng ca c th qua cc th h c gi l hin tng di truyn. Song
s sao chp trn khng phi l tuyt i, con ci vn c nhng nt khc nhau v khc
vi cha m. l hin tng bin d.
T xa xa, con ngi quan tm n cc hin tng di truyn v bin d. Nhiu
phng php thun ha, chn lc v lai ging c cc dn tc c xa p dng.
Nhng do s hiu bit cn hn ch nn trong vic gii thch cc quy lut di truyn v
bin d vn cn rt nhiu quan nim ngy th, sai lm.

Giai on trc Mendel

Th k IV - V trc Cng nguyn, c hai lun thuyt v s di truyn ca cc tnh trng
c nu ra. Hippocrates theo thuyt di truyn trc tip, cho rng vt liu sinh sn c
thu thp t tt c cc phn ca c th, nh vy tt c cc c quan trong c th u c
nh hng n con chu. V sau, thuyt di truyn gin tip ca Aristotle bi bc quan
im ca Hippocrates, cho rng vt liu sinh sn c to ra t cht dinh dng m v
bn cht tin nh cho cu to ca cc phn khc nhau trong c th.

69/132

Lamarck, ngi u tin xy dng hc thuyt kh hon chnh v c h thng v s pht

trin lch s ca sinh gii, cao vai tr ca ngoi cnh. ng nu ra quan nim v s di
truyn tp nhim, cho rng cc bin i thu c trong i c th cng di truyn c.

Vo th k XIX, sinh vt hc pht trin mnh m, cc php lai ging c s dng rng
ri ng thc vt. Qua , cc nh sinh vt hc hiu c rng c cha v m u gp
phn vo cc tnh trng ca hu th. Tuy nhin quan nim ph bin thi ny l s di
truyn ha hp, tc l tnh trng ca cha m trn ln nhau to nn tnh trng trung
gian ca con ci.

70/132

Darwin chu nh hng ca thuyt di truyn gin tip. Trong tc phm S bin i
ca cc ng vt v thc vt trong nui trng (1868), ng pht trin thnh thuyt
pangen (Pangenesis). Theo , mi phn trong c th sn sinh ra nhng phn t nh gi
l gemmule (mm) theo mu tp trung v c quan sinh dc. Mi c th sinh ra do s ha
hp tnh di truyn ca c cha v m, v hn th cn bao gm c cc tnh tp nhim.
Nm 1871, F. Galton tin hnh thc nghim kim tra thuyt pangen. ng truyn
mu th en cho th trng, sau lai nhng con c truyn mu vi nhau. Lp li th
nghim qua 3 th h vn khng tm thy nh hng g n th trng. Nh vy, trong
mu th khng cha gemmule.
n cui th k XIX, gii khoa hc vn cha c c quan nim ng n v tnh di
truyn. Darwin nhiu ln nhn mnh: v cc quy lut di truyn v bin d, chng ta hy
cn bit qu t. ng tic l, nm 1866 Mendel cng b tc phm Cc th nghim
lai thc vt nhng Darwin khng c bit n.

Giai on di truyn Mendel

Gregor Mendel (1822 1884) s dng cy u H Lan Pisum sativum lm i tng

chnh trong nghin cu di truyn. T nm 1856 n 1863 ng trng khong 37.000
cy v quan st c bit khong 300.000 ht. Nh c phng php th nghim c o,
ng pht hin c cc quy lut di truyn, t nn mng cho di truyn hc hin i.
Kt qu nghin cu ca ng c trnh by trong tc phm Cc th nghim lai thc
vt di 44 trang, bo co trc Hi cc nh T nhin hc thnh ph Brno vo ngy 8
thng 2 v 8 thng 3 nm 1865 v c cng b trong K yu ca Hi vo nm 1866.
Mendel chng minh s di truyn cc tnh trng c tnh gin on c chi phi bi
cc nhn t di truyn v dng cc k hiu s hc n gin biu hin cc quy lut
truyn th tnh di truyn.

71/132

Do hn ch v tri thc ca Sinh hc ng thi, cng trnh ca Mendel khng c cng

nhn trong sut 35 nm. Mi n nm 1900, khi H. de Vries (H Lan), E. K. Correns
(c) v E. V. Tschermak (o) c lp pht hin li cc quy lut Mendel th pht minh
ca ng mi c tip nhn. Nm 1900 c coi l nm khai sinh ca Di truyn hc v
th k XX l th k pht trin ca Di truyn hc.

Nm 1901, H. de Vries nu ra thuyt t bin. Nm 1902, W. Bateson v L. Cunot

chng minh cc quy lut Mendel ng vt. Trong khong thi gian ny, cc hin tng
tng tc gen cng c pht hin. Cc quan im u tin v s di truyn ca nhim
sc th c nu ra. Nm 1903, T. Boveri chng minh vai tr ca nhn v W. Sutton
gn cc nhn t Mendel vi nhim sc th. c bit, A. Weismann da trn s suy lun
xut thuyt di truyn nhim sc th. ng tin on c c ch nguyn phn,
gim phn, vai tr ca nhim sc th trong nhn t bo; ng thi ra gi thuyt th
quyt nh (determinant) mang tnh di truyn gin on, l c s cho khi nim gen sau
ny.

72/132

Tn gi mn Di truyn hc (Genetics) do nh di truyn hc Anh W. Bateson nu ra

nm 1906. Nm 1909, nh khoa hc an Mch W. Johannsen nu ra cc thut ng: gen
(gene), kiu gen (genotype), kiu hnh (phenotype).

S pht trin ca thuyt di truyn nhim sc th

S pht trin ca t bo hc na cui th k XIX to c s cho s hiu bit cng trnh
ca Mendel. S kt hp gia t bo hc v di truyn hc to ra bc pht trin mi
cho di truyn hc.

Nm 1911, T. H. Morgan cng cc cng s xy dng hc thuyt di truyn nhim sc

th v bn di truyn nhim sc th da trn cc nghin cu trn i tng rui gim
Drosophila melanogaster. Kt qu thu c chng minh cc gen nm trn nhim sc
th xp dc to thnh nhm lin kt gen. Hc thuyt di truyn nhim sc th xc nhn
s ng n ca cc quy lut Mendel, cho thy cc gen c c s vt cht, gn cht vi
cu trc t bo. Di truyn hc c in c lc c gi l Di truyn Mendel Morgan. T.
H. Morgan c nhn gii Nobel vo nm 1934.
Nm 1920, N. I. Vavilov nu ra quy lut v cc dy tng ng trong bin d v sau
ny nu ra thuyt v cc trung tm ging cy trng trn th gii.

73/132

Nm 1925 1927, Muller chng minh tc ng gy t bin ca tia X, t c s cho

cc nghin cu v t bin nhn to.
Nm 1933, T. Painter pht hin nhim sc th khng l cn trng hai cnh (Diptere),
t c s cho vic nghin cu t bin nhim sc th v lp bn di truyn t bo.
Thp nin 1940, thuyt mt gen- mt enzyme a v cho George Beadle v Edward
Tatum gii Nobel vi cng trnh nghin cu trn nm mc Neurospora crassa chng
minh gen kim tra cc phn ng sinh ha. Di truyn hc c c bc pht trin mi:
i vo chi tit hot ng ca gen.
Cng trong nhng nm 1930 - 1940, Barbara McClintock pht hin cc gen di chuyn
dc trn nhim sc th m sau ny c gi l cc yu t di ng (transposable
elements) khi nghin cu trn ng Zea mais. B c nhn gii Nobel vo nm 1983
khi tui 80.
Cho n cui nhng nm 40 ca th k XX, di truyn hc c coi l giai on kinh
in v nhng nguyn l c bn c tm ra, khi nim gen c pht trin v c th
ha cng vi s biu hin ca chng.

S pht trin ca di truyn hc phn t

Sau th chin th hai, s pht trin mnh m ca khoa hc trn nhiu lnh vc dn
n nhiu pht minh ln cho Di truyn hc cp phn t.
Tip ni nghin cu ca Frederick Griffith (1928), nm 1944, Oswald Avery, C. M.
MacLeod v M. McCarty xc nh c bn cht ca hin tng bin np v chng
minh DNA l vt cht di truyn. Nhng n nm 1952, vai tr di truyn ca DNA mi
c cng nhn sau th nghim tm ra bn cht ca hin tng ti np trn phage T2 do
Alfred Hershey v Martha Chase tin hnh.
S pht hin cu trc chui xon kp ca DNA vo nm 1953 bi James Watson v
Francis Crick l mt trong nhng khm ph quan trng nht trong lch s di truyn hc.
N m ra mt cch hiu mi v hot ng ca gen v s di truyn cp phn t.
Bn cht ha hc ca gen- n v di truyn c bn trong h thng sng- c lm
sng t.

74/132

Nm 1956, hc thuyt trung tm (central dogma) ca Sinh hc phn t c Francis

Crick xut.
Nm 1961, M. Nirenberg v J. Matthei gii c nhng m di truyn u tin v n
nm 1966, ton b 64 codon m ha c nhm ca M. Nirenberg v nhm ca H.G.
Khorana xc nh.
Nm 1961, J. Monod v F. Jacob pht hin c ch iu ha sinh tng hp protein
prokaryote theo m hnh operon.

T khi k thut di truyn ra i cho n nay

T u thp nin 70 th k XX, k thut di truyn ra i to nn mt cuc cch mng
ln trong Di truyn hc v Sinh hc.
S hiu bit v gen mc tng nucleotide dn n k thut mi: gy t bin
im nh hng. S bin i nh hng trn gen dn n s thay i trnh t amino
acid trong phn t protein mt cch c ch ; t thc y s pht trin ca cng ngh
protein.
u nhng nm 1990, nghin cu in silico (trn my in ton) to thun li ln cho
cc nghin Sinh hc trong c Di truyn hc.
K thut di truyn ko theo s bng n ca Cng ngh Sinh hc, mang li nhng ng
dng to ln nhng cng t ra nhiu vn ng lo ngi i vi con ngi.

75/132

i tng v phng php nghin cu di

truyn hc
i tng nghin cu
Di truyn hc (Genetics t ch Latin Genetikos, ngha l ngun gc, sinh sn) l
mn khoa hc nghin cu v tnh di truyn v bin d ca sinh vt. Hay ni cch khc,
di truyn hc nghin cu cc qui lut truyn t thng tin t th h t tin cho con chu
v nhng qui lut bin i ca qu trnh truyn t .
C th coi tnh di truyn l c tnh ca b m truyn li cho con ci nhng tnh cht
v c im pht trin ca minh, m nh cc loi sinh vt bo tn c nhng c
im ring ca mnh qua hng lot th h. Tnh di truyn c m bo qua qu trnh
sinh sn. Trong qu trnh sinh sn hu tnh, nh s kt hp gia t bo sinh dc c v
t bo sinh dc ci m thc cht l s kt hp vt cht di truyn ca b v m, m
m bo s k tc vt cht di truyn gia cc th h. Vi hnh thc sinh sn v tnh, tnh
di truyn c m bo nh s phn chia ca cc t bo soma. Tnh di truyn va m
bo cho s k tc cc c tnh ca sinh vt qua cc th h, va m bo cho cc c th
sinh vt mt hnh thc pht trin c th, hnh thnh nn nhng tnh trng v c tnh
nht nh.
Trong bt k hnh thc sinh sn no, sinh vt cha phi c sn tt c cc tnh trng
v c tnh ca c th trng thnh m cc tnh trng v c tnh s c hnh thnh
dn trong qu trnh pht trin c th trong nhng iu kin mi trng nht nh.
C s vt cht ca tnh di truyn l tt c nhng yu t cu trc t bo c kh nng
ti sinh, phn ly, t hp v cc t bo con trong qu trnh phn chia ca t bo c th.
Vt cht di truyn xt cp t bo l nhim sc th, cp phn t l cc gen, trn
cc phn t nucleic acid. S nhn i ca nhim sc th cng nh qu trnh phn ly ca
chng trong nguyn phn cng nh gim phn c vai tr c bit quan trng m bo
k tc vt cht di truyn n nh qua cc th h.
Tm li, di truyn l c tnh c bn ca c th sinh vt m bo cho s k tc vt cht
di truyn v chc nng qua cc th h. Nh vy i tng nghin cu ca di truyn hc,
ngoi vic nghin cu tnh di truyn cn nghin cu qu trnh bin d tnh di truyn ca
sinh vt.
Bin d biu hin s sai khc gia cc c th con ci vi cha m hay vi cc c th
khc cng n. Mt mt s bin i ca b my thng tin di truyn dn n cc bin d,
mt khc cng chnh cc c ch di truyn to s a dng trong sinh gii. Tuy bin d c

76/132

s lng rt ln v ht sc a dng nhng xy ra trong mt khun kh nht nh nn c

th xp cc sinh vt vo nhng n v phn loi nh loi, chi, h, b, lp, ngnh
Bin d, di truyn, chn lc v cch ly l nhng nhn t tin ha c bn. C hai loi bin
d: bin d khng di truyn v bin d di truyn. Bin d khng di truyn l cc thng
bin c gi tr thch nghi trong i c th. Bin d di truyn to ra s a dng, cung cp
nguyn liu cho tin ha; di truyn gip duy tr cc c tnh; cn chn lc t nhin l
nhn t nh hng, dn n s a dng ca s sng nh ngy nay.

Phng php nghin cu

Cc phng php c s dng ph bin trong nghin cu Sinh hc nh phng php
sinh ha, phng php sinh l cng c s dng trong nghin cu Di truyn hc.
Cc phng php vt l hc, ha hc cng h tr rt nhiu. Ngoi ra, Di truyn hc cn
c nhng phng php nghin cu c th v thng dng sau:
Phng php lai
Ngi ta tin hnh tp giao gia cc c th v theo di s di truyn cc tnh trng qua
cc th h ni tip, phn tch kt qu v rt ra quy lut di truyn. y l phng php
c bn v c th ca Di truyn hc.
Phng php lai thng kt hp vi vic thu nhn cc bin d v v sau cn c s
dng vi khun. Phng php lai xa cho php xc nh mi quan h gia cc loi v
cc ging. Sau ny, phng php lai t bo sinh dng cng c s dng ph bin
ng vt, thc vt. Mt bin th ca phng php lai c s dng trong nghin cu di
truyn ngi l phng php nghin cu ph h.
Phng php ton hc
Trong phn tch di truyn, cc phng php ton hc gip i chiu kt qu thc nghim
vi d on l thuyt. y l phng php khng th thiu trong nghin cu di truyn
s lng v thng bin.
Phng php t bo hc
S kt hp gia T bo hc v Di truyn hc nng Di truyn hc ln tm cao mi.
Cc k thut mi trong nhum tiu bn, cc loi knh hin vi hin i gip cho s
phn tch cu trc, hot ng ca nhim sc th cng nh bn cht ca cc qu trnh xy
ra trong t bo t c nhiu thnh tu mi.

77/132

K thut di truyn
K thut ti t hp DNA va l thnh tu ca Di truyn hc, va l cng c quan trong
gip i su vo cc c ch di truyn v c cc qu trnh Sinh hc rt kh nghin cu
trc y.
Ngy nay, theo s pht trin chung ca Khoa hc, ngy cng c nhiu phng php v
k thut mi c ng dng trong nghin cu Di truyn hc.

78/132

Quan h gia di truyn hc vi cc khoa

hc khc v vi thc tin
ngha l lun ca Di truyn hc l xy dng nhng l lun sinh vt hc v tnh di
truyn, gp phn nghin cu cc quy lut ca s sng, cc quy lut tin ha ca sinh
vt. ngha thc tin ca Di truyn hc l vn dng cc nguyn l di truyn nhm xy
dng nhng phng php iu khin tnh di truyn v tnh bin d, gii quyt nhng vn
thc tin trong chn ging, trong y hc, cc Khoa hc x hi v cc vn lin quan
n o c, x hi v tng lai loi ngi.

Di truyn hc v Khoa hc chn ging

Di truyn hc l c s khoa hc cho chn ging. Cc thnh tu ca Di truyn hc c
ng dng sm nht v nhiu nht trong Khoa hc chn ging. Kin thc Di truyn hc
l c s xy dng cc phng php to ngun bin d cho chn ging (lai to, gy
t bin nhn to, k thut di truyn), ci tin cc phng php chn lc, thm ch to
ra c nhng ging mi vt gii hn ca tin ha.

Di truyn hc v Y hc
Di truyn y hc gip pht hin c nguyn nhn nhiu bnh tt di truyn v ra
phng hng ngn nga, iu tr mt s bnh ny. Phng php iu tr cc bnh di
truyn bng liu php gen (genotherapy) - a gen bnh thng vo thay th gen bnh m ra mt hy vng mi. Y hc tng lai vn nng v d phng, a b sp sinh c
chn on phn t, on bit c cc bnh c kh nng mc phi c bin php d
phng hoc iu tr sm.
Cui nm 1989, Hoa K bt u u t 3 t USD cho vic gii trnh t b gen ngi v
vo thng 10/2004, kt qu D n b gen ngi c cng b. Hng chc ngn gen
hot ng b no ngi c bit n to nn bc ngot ln cho vic nghin cu
cc cht iu khin tr thng minh. Gi y, ngi ta bt u ni n nhng a tr
theo n t hng ng vic ci bin di truyn ngi theo mun. Vn ny lin
quan n o l con ngi nn cn nhiu tranh ci.

Di truyn hc v tin hc
Nhng nm cui th k XX xy ra cuc rp ni ca th k gia hai lnh vc ang
pht trin nh cao l Sinh hc v Tin hc. ng dng Sinh hc vo Tin hc th hin
trong tng ch to cc my in ton sinh hc (biocomputer). Ngc li, vi kh
nng lu tr v x l mt khi lng thng tin s ca mnh, Tin hc tr thnh
79/132

cng c h tr c lc cho Sinh hc t nhng bc tin di. C th k ra 3 lnh vc

ng dng sau y:
Gii trnh t nucleotide ca nhiu b gen
Xc nh trnh t nucleotide ca b gen l mt cng vic s. Hn 80 phng th
nghim trn th gii trong c Hoa K thc hin D n B gen ngi. Nht Bn u t
2 t yn cho vic gii trnh t b gen ca nm men Saccharomyces cerevisiae, vi khun
Escherichia coli, Bacillus subtilis. Chu u chi chi khong 70 triu USD cho chng
trnh trn i tng thc vt Arabidopsis thaliana.
Tin hc ng vai tr thu thp, lu tr v khai thc s dng s liu. ng thi, c ch
lu tr v biu hin thng tin di truyn ca sinh vt s cung cp nhiu tng cho Tin
hc m phng.
Th nghim trn my in ton (in silico)
T cc phng php th nghim trong c th sinh vt (in vivo) tin n cc th nghim
trong ng nghim (in vitro) v gi y l s kt ni gia Tin hc v Sinh hc to ra
phng nghin cu mi: th nghim trn my in ton (in silico). Ngi ta xy dng
cc i tng nghin cu nhn to chng hn nh gen, protein nh in ton, thc hin
cc th nghim trn my tnh, sau ra thc tin kim nghim kt qu v chnh l.
Th nghim in silico c th h thp chi ph, rt ngn thi gian v an ton. Tuy nhin t
m phng n thc tin c th l mt khong cch kh ln.
ng dng ca bioinformatics trong bin i cht lng protein
Trnh t nucleotide ca gen quy nh trnh t chui polypeptide, t quy nh cc bc
cu trc khng gian tng ng ca phn t protein. Da vo tnh cht v quy lut v
cu trc phn t protein, phn mm in ton s xy dng nn cc m hnh cu trc
khng gian khc nhau ca mt phn t protein nht nh, chn m hnh ti u ri kim
chng bng cc th nghim in vitro, in vivo. Bng cch ny, ngi ta to ra nhng loi
protein vi nhng tnh cht c ci thin so vi t nhin hoc thm ch to ra nhng
loi protein cha h c trong t nhin.

Di truyn hc v kha cnh o c, x hi

Di truyn hc t lu l c s khoa hc cho mt s o lut, chng hn Lut hn nhn
v gia nh cm kt hn gia nhng ngi c quan h h hng trc h ba i. Ngy
nay, s pht trin vt bc ca Cng ngh Sinh hc ni chung v Cng ngh gen ni
ring t con ngi trc nhng vn mi v gio dc hc, lut hc, trit hc, x
hi hc.

80/132

C ba xu hng vt gii hn tin ha t nhin ang din ra: (1) a gen ngi vo cc
sinh vt; (2) Nhn gen hoc c quan t cc sinh vt; (3) Nng cao tui th con ngi,
tin n con ngi bt t. Cc th nghim to dng phi ngi (human embryon cloning)
v sinh sn v tnh ng vt thnh cng c cng b. Tng lai, s c s can thip
trc tip vo b my di truyn ci thin con ngi sinh hc, thm ch to ra chng
ngi mi u vit hn con ngi t nhin.
Vy u l gii hn? V mt o l, nhng th nghim tng t nh trn khng c
php tin hnh trn ngi. l o l sinh hc (Bioethics). Nhng nm u ca thp
nin 1990, Ngh vin Chu u thng qua 3 lut cm cc th nghim lin quan n
o l. y ban Quc t v o l sinh hc IBC (International Commitee of Bioethics)
ca UNESCO cng c thnh lp. Cu tr li ty thuc vo quan im ca cc Nh
cm quyn v c Nh khoa hc.

81/132

Acid Nucleic l vt cht di truyn cp

phn t
Tiu chun ca vt cht di truyn
Mi sinh vt trong qu trnh pht sinh v pht trin u mang nhng c im ring
bit ca loi v ca c th. Cc c im ny phi c m ha di dng thng tin
di truyn. ng thi cc c im ca loi nh s pht trin phi, s bit ha t bo
cng phi c chng trnh ha v mt di truyn.
Theo George J. Brewer, 1983, vt cht di truyn phi hi 3 tnh cht sau y:
Mang thng tin di truyn c trng cho loi
Vt cht di truyn phi mang thng tin c trng cho loi. y khng ch n gin l
nhng thng tin v thnh phn cu trc c th m cn l nhng thng tin v c im
pht trin c th qua cc giai on khc nhau trong chu trnh sng ca c th. Khi nim
ny bao hm hai :
Vt cht di truyn phi c kh nng m ha mi thng tin di truyn ca th h trc
chuyn giao cho th h sau. Phng thc m ha da trn nguyn tc: s lng, thnh
phn, trnh t sp xp ca cc nucleotide trn gen cu trc s quy nh s lng, thnh
phn, trnh t sp xp ca cc acid amin trn chui polypeptide tng ng.
S bit ha t bo v qu trnh pht trin c th cho thy mi loi t bo, ty vo giai
on pht trin m c nhng yu cu v biu hin khc nhau. iu ny chng t cc gen
trong h gen ca chng khng hot ng ng lot v lin tc: c gen hot ng giai
on ny nhng li b km hm giai on khc. chnh l c ch iu ha biu hin
gen. C ch ny do mt h thng cc gen chuyn trch ngoi gen cu trc m nhim.
C kh nng ti bn
Vt cht di truyn phi c kh nng hnh thnh cc bn sao, trong cha y cc thng
tin di truyn ca loi v ca c th truyn li cho th h sau.
prokaryote, thng qua hnh thc phn bo trc tip (trc phn), mi t bo con s
nhn c mt bn sao vt cht nhn ging ht t bo m.
eukaryote, hot ng phn bo gin tip (gin phn) c hai hnh thc: phn chia
nguyn nhim (nguyn phn) v phn chia gim nhim (gim phn). Trong , qua
nguyn phn, mi t bo con nhn c mt bn sao cha ton b thng tin di truyn
82/132

trong nhn. Qua gim phn, mi t bo n bi ch nhn c bn sao ca mt na vt

cht di truyn trong nhn.
i vi akaryote, phng thc sinh sn l nhn ln hng lot trong t bo k ch. Trong
5 giai on ca mt chu trnh gy c (hp ph, xm nhp, tng hp, lp rp, phng
thch), giai on th 3 bao hm s ti bn vt cht di truyn cho th h sau.
C kh nng bin i
C ch ti bn ca vt cht di truyn d rt chnh xc vn khng th trnh khi nhng
sai st vi tn sut thp. Nhng sai st ny to nn cc t bin, lm thnh ngun
nguyn liu ch yu cho qu trnh tin ha ca sinh gii.
Ngoi t bin, hin tng ti t hp v cc yu t di truyn vn ng (transposable
genetic element) cng gp phn lm bin i vt cht di truyn ca cc sinh vt.

Chng minh acid nucleic l vt cht mang thng tin di truyn

S khm ph ra hin tng Bin np (transformation)
Nm 1928, Frederick Griffith quan st thy mt hin tng b n khi tin hnh cc th
nghim trn Streptococcus pneumoniae. Vi khun ny vn gy bnh vim phi ngi
nhng li c kh nng gy cht chut. Tuy vy, nhng chng khc nhau ca loi ny
c c lc khc nhau.

83/132

Trong th nghim, Griffith s dng hai chng vi khun khc bit nhau bi hnh dng
khun lc v c tnh. Chng c gy cht chut, k hiu l S (smooth) cho khun lc
nhn, lng v t bo c bc bi mt v nhy bng polysaccharide. Chng khng c
c k hiu l R (rough), khng c v nhy, cho khun lc sn.
Nu dng chng R sng hoc chng S b un si, tim mt cch ring r cho chut
th chut u khng cht. Nhng nu b tim ng thi chng R sng v chng S un
si th chut s cht v vim phi. Hn th na, t xc chut cht c th phn lp c
nhng vi khun sng. Nhng vi khun ny s hnh thnh khun lc trn lng v biu
hin c tnh nu cho ly nhim ln sau. Nh vy, bng mt cch no , nhng mnh
v t bo t chng S b un si bin i cc vi khun R sng tr thnh cc vi khun
S sng. Hin tng ny c gi l Bin np.
Tip ni nghin cu trn, nm 1944, Oswald Avery, C. M. MacLeod v M. McCarty
xc nh c bn cht ca hin tng bin np. Cc tc gi phn tch cc phn t
thu c t cc mnh v t bo S thnh tng nhm cht v kim tra kh nng bin np
ca chng.

84/132

u tin, h thy rng bn thn cc polysaccharide khng gy bin np t bo R. V

vy, d chc chn c lin quan n tnh gy bnh, v nhy polysaccharide ch l mt
biu hin kiu hnh ca c tnh. Quan st cc nhm cht khc, cc tc gi nhn thy
duy nht ch c nhm phn t DNA gy ra s bin np t bo R. H suy lun rng DNA
chnh l yu t xc nh c tnh v polysaccharide v t xc nh c tnh gy bnh.
Vic cung cp DNA ca t bo S cho cc t bo R sng cng ngang bng vi vic cung
cp gen S (gen quy nh v nhy) cho chng.
Th nghim ca Hershey Chase
Mc d cc th nghim ca Avery v cng s c cu tr li cui cng, nhng cc nh
khoa hc thi by gi vn min cng chp nhn DNA (hn l protein) l vt cht di
truyn. Nm 1952, th nghim ca Alfred Hershey v Martha Chase trn phage T2
chm dt tranh lun . H suy on rng s nhim phage phi bao hm vic a vo
vi khun mt thng tin chuyn bit gip ti sn xut virus.

Phosphor khng tm thy trong protein nhng li c trong cu trc DNA; ngc li, lu
hunh ch hin din trong protein m khng c mt trong DNA. Hershey v Chase
dng 32P nh du DNA ca mt nhm phage T2 v dng 35S nh du protein
ca mt nhm phage T2 khc. K , dng hai nhm phage ny cho nhim ring r vo
E.coli vi s lng ln virus. Sau thi gian gy nhim thch hp, h dng lc khuy
85/132

tch v virus cn bm bn ngoi ra khi t bo vi khun. S dng phng php ly tm

tch ring v virus vi t bo vi khun ri phn tch phng x. Vi nhm phage nh
du bng 32P, trong t bo vi khun c cha cht phng x chng t DNA ca phage
vo trong vi khun. Vi nhm phage nh du bng 35S, cht phng x nm trong phn
v virus b li.
Kt qu th nghim trn cho thy protein v ca phage khng xm nhp t bo vi khun
m ch c DNA ca phage c np vo. Phn t DNA ny gip sn sinh ra th h phage
mi. Nh vy, DNA chnh l vt liu di truyn ca phage.
RNA cng l vt cht di truyn

Phn ln virus k sinh thc vt c cu to chnh gm phn v bng protein v phn li

l RNA, in hnh nh virus gy bnh khm thuc l TMV (Tobacco Mosaic Virus) v
HRV (Holmes ribgrass virus). C hai loi virus ny khi xm nhp vo t bo l s lm
bin i s trao i cht ca t bo khin cho dip lc t b phn hy, gy ra nhng m

86/132

mu trn nn l xanh. Tuy nhin, chng khc bit nhau bi cch gy bnh, mu sc v
kch thc cc m khm.
Ngi ta c th phn lp ring r protein v v RNA di dng tinh, sau tng hp tr
li thnh ht virus. Cng c th dng protein v RNA ca hai loi hoc hai chng virus
khc nhau tng hp nn mt dng virus ghp.
Nm 1957, Fraenkel Conrat v cng s tng hp nn mt loi virus ghp t RNA
ca HRV v protein v ca TMV. Cho nhim loi virus ghp ny vo cy thuc l lnh
th thy cy b nhim bnh vi triu chng bnh ca HRV. Ngi ta cng phn lp c
HRV t l cy b bnh. Nh vy trong trng hp ny, RNA chnh l c s vt cht ca
s di truyn cn protein ch ng vai tr v bc h tr.

87/132

Ti bn DNA
Cc nguyn tc v c im chung ca ti bn DNA
S ti bn DNA theo c ch bn bo tn (semi-conservative mechanism)

u im ni bt ca m hnh Watson Crick l cho php d on ngay c c ch ti

bn ca DNA. Ngay sau khi m hnh cu trc DNA c nu ra, nhiu th nghim c
tin hnh xc nhn cc d on.
Trong chui xon kp DNA, hai mch n lin kt vi nhau bng mt quan h b sung.
Nh vy, trong qu trnh ti bn, nu hai mch n tch ri nhau v mi mch n c
dng lm khun tng hp nn mt mch mi theo nguyn tc b sung th kt qu l
t mt phn t DNA ban u to ra c hai phn t mi ging ht nhau. V trong
mi phn t DNA con u mang mt mch c v mt mch mi nn kiu ti bn ny
c gi l bn bo tn.
C ch bn bo tn c Meselson v Stahl chng minh bng thc nghim vo nm
1958. Ngi ta nui E.coli trong mi trng c ngun N15, t bo s s dng N15
88/132

tng hp DNA cho n khi tt c DNA ca vi khun u mang ng v nng N15. Sau
cc t bo c chuyn sang mi trng c cha N14. Cch khong thi gian u
n tng ng vi mi t phn bo, ngi ta ly cc t bo em chit tch DNA. Bng
phng php ly tm trong gradient t trng CsCl, cc loi DNA nng, nh v lai c
tch ra, kt qu phn tch ph hp vi kiu ti bn bn bo tn.

C ch phn t ca qu trnh ti bn DNA

Nhng nghin cu tip theo tm ra c ch phn t ca qu trnh ti bn. l mt
qu trnh phc tp c s khc bit prokaryote v eukaryote nhng phi tri qua c ch
chung sau:
Cc lin kt hydrogen n nh cu trc xon v lin kt hai mch n vi nhau
phi c ph v tch ri hai mch.
Phi c on mi (primer), tc l on DNA/RNA ngn, bt cp b sung vi
mch khun to u 3OH t do.
Nguyn liu tng hp DNA l cc desoxynucleosid 5-triphosphate (dNTPs):
dATP, dGTP, dCTP v TTP.
Mch khun lun c c theo chiu 3-5 trong khi mch mi c tng hp
theo chiu 5-3. Mi nucleotide mi c gn vo u 3OH ca mch ang
ko di bng lin kt cng ho tr. Nng lng cho s polymer ho n t vic
thy phn cc dNTPs, loi ra cc pyrophosphate.
Mi bc c thc hin mt cch nhanh chng, chnh xc di s iu khin
ca enzyme c hiu.

89/132

n v ti bn, im khi u v im kt thc ti bn

Mi on DNA c ti bn nh l mt n v ring l c gi l mt n v ti bn
(replicon). Cc DNA dng vng, kch thc nh (chng hn nh nhim sc th ca vi
khun v DNA virus) ch gm mt n v ti bn duy nht. S ti bn khi u t mt
im gi l im khi u sao chp (origin) v lan ra theo hai hng, hnh thnh hai
chc ba ti bn (replication fork) cho n khi gp nhau ti im kt thc (terminus).
im khi u ti bn c khuynh hng giu A-T d dng khi u tch ri hai mch
n.
eukaryote, mi phn t DNA bao gm nhiu n v ti bn. Mi n v ti bn c
im khi u ring v s ti bn cng lan ra theo hai hng. Khi cc chc ba ti bn
gp nhau th s ti bn DNA c hon thnh. V d nh t bo hu nh in hnh c t
50.000 100.000 n v ti bn, mi n v ti bn c kch thc khong 40 200 kb.
1.4 S tng hp mch mi ca DNA xy ra lin tc trn mt mch v gin ontrn
mch kia(Semi-discontinuous replication - Ti bn bn gin on)

90/132

Trong ti bn bn bo tn, c hai mch mi ca DNA c tng hp ng thi mi

chc ba ti bn. Tuy nhin, c ch ti bn DNA ch cho php s tng hp theo hng
5 3, v th trn hai mch khun c hng ngc nhau, s tng hp mch mi khng
din tin ging nhau.
T im khi u ti bn, trn mch khun 3- 5 s tng hp mch mi din
ra mt cch lin tc theo chiu 5- 3 cng chiu vi hng tho xon. Mch
ny c gi l mch ti (leading strand).
Trong khi trn mch khun 5 3, s tng hp mch mi cng din ra theo
hng 5 3 nhng ngc vi hng tho xon. Chnh v th, s tng hp
mch mi khng xy ra lin tc m di dng nhng on ngn c gi l
on Okazaki (kch thc 1000 2000 bp prokaryote v 100 200 bp
eukaryote). Mch mi c tng hp theo kiu gin on ny c gi l mch
chm (lagging strand).
Trn thc t, s tng hp c hai mch theo cng hng v mch khun chm c un
vng quay 180o ti chc ba ti bn, tr nn cng hng vi mch khun ti.

91/132

Mi RNA
Mch ti v tt c nhng on Okazaki ca mch chm ch c th c tng hp bng
cch ko di mt mi bt cp sn trn mch khun. Mi l mt on RNA ngn, c
tng hp bi mt phc hp protein gi l primosome bao gm nhiu protein v mt
enzyme c tn l primase. V d, E. coli, phc hp primosome bao gm cc Dna B
helicase (tch ri hai mch n) v DNA primase (tng hp mi RNA t khun DNA).
Cc mi RNA ny sau s b phn hy bi RNase v c thay bng trnh t DNA
nh vo hot ng ca DNA polymerase. Enzyme ligase s ni lin cc on DNA li
vi nhau.
C ch dng mi l RNA thay v DNA dng nh lin quan n tnh chnh xc cao
trong ti bn DNA v hot tnh sa sai ca cc DNA polymerase.

Ti bn DNA prokaryote
Cc nghin cu v ti bn DNA c tin hnh da trn cc h thng th nghim in
vitro. theo di ti bn DNA, ngi ta dng thymidine H3 phng x. Qu trnh ti bn
xut pht t im khi u v lan ra hai pha. Khi quan st DNA vng trn ang ti bn,
ta thy dng DNA hnh con mt (eye replication). Hai chc ba ti bn lan dn, cui
cng to thnh hai phn t DNA lai, trong c mt mch mang du phng x T-H3.
C trng hp, s ti bn ch xy ra v mt pha.

Din bin ti bn nhim sc th vng trn c th chia lm ba giai on: khi u

(initiation), ko di (elongation), kt thc v phn chia (termination & segregation).

92/132

Giai on khi u
Tho xon phn t DNA

M hnh phn t ca DNA vng vi khun hay virus cho ta mt khi nim
chung v cc cu hnh c th c ca mt phn t DNA. Dng th nht l siu
xon. Dng th hai c cu trc li hn, thng l do b t mt ch trn mt
trong hai mch ca phn t DNA. Dng th ba tng ng vi cu trc thng
do s ct t trn c hai mch trong phn t DNA. Trong , dng siu xon
l dng c bn v cu trc v chc nng.

93/132

Nh vy, bt u ti bn, phn t DNA phi c tho xon nh vo

hot ng ca cc enzyme c tn l topoisomerase. C hai loi topoisomerase.
Loi I tho dng siu xon, chng gn vo phn t DNA v ct mt trong hai
mch. Sau khi gii phng mt phn t DNA c tho xon, cc enzyme
ny s ni li ch t. Enzyme loi I c bit r nht l protein ca E.
coli. Cc topoisomerase loi II c kh nng tho cc nt ny sinh do cc bin
i cu trc ca chui xon kp bng cch ct t c hai mch DNA. Enzyme
c bit r nht l gyrase ca E. coli. Gyrase s dng nng lng t s thy
phn ATP tho xon DNA.

94/132

Tch ri hai mch n ti im khi u ti bn

E.coli, OriC cha bn v tr gn vi protein khi u c tn l Dna A. Mi

v tr c kch thc 9bp. S tng hp cc protein ny gn lin vi tc tng
trng t bo v th vic khi u ti bn DNA cng gn lin vi tc tng
trng. tc tng trng cao, cc nhim sc th ca vi khun c th bt
u ln ti bn th hai trc khi ln ti bn th nht kt thc ti hai im
khi u mi. V vy, mi t bo con s nhn c mt nhim sc th ang
c ti bn mt phn.
Qu trnh ti bn bt u khi DNA OriC qun quanh mt phc hp protein
Dna A gm 30-40 phn t, mi phn t gn vi mt ATP. iu ny thc
y s tch ri hai mch ti ba trnh t lp 13bp giu A T, cho php
protein Dna B gn vo. Dna B l mt DNA hehicase, nm trong phc hp
primosome. Cc helicase ph v lin kt hydro gia cc base nh nng lng
thy phn ATP. C nhiu loi helicase cng hot ng ng thi: Mt s gn
trn mch 3 5 nh cc Rep, s khc gn trn mch 5 3 nh helicase II
v III.
Cc mch tch ri s c n nh di dng mch n nh cc protein
SSB (Single Strand Binding lin kt vi mch n). Cc protein ny gn
95/132

ln khp phn mch n lm cho hai mch khng kt hp tr li c. Mch

khun c s dng n u th cc protein SSB c gii phng khi khun
n .
Giai on ko di
Tng hp mi RNA

Cc DNA polymerase ch c th tng hp DNA bng cch ko di mt mi

RNA bt cp sn trn khun. Nh bit, mi ny c tng hp nh
hot tnh ca enzyme primase c trong phc hp primosome.
S tng hp mch mi din ra theo kiu bn gin on, ko di mi RNA

96/132

DNA polymerase III l mt phc hp dimer. Mi phn gm nhiu n v gn

vi nhau, chu trch nhim tng hp mt mch n DNA nhm m bo cho
tc tng hp ca c hai mch bng nhau. n v c hot tnh polymerase
tht s. n v c chc nng c sa nh hot tnh exonuclease 3- 5, t
lm tng tnh chnh xc trong ti bn. n v gip gn polymerase vo
DNA. y l nhn t duy tr, khc nhau hai mch khun, quy nh di
ca on DNA c tng hp mch ti (di) khc vi mch chm (ngn).
DNA polymerase III bt u ti bn trn mi mch khun bng cch gn
vo mch (nh n v ) v lp cc nucleotide b sung vo v tr tng ng,
ko di on mi RNA bt cp sn trn khun t u 3OH t do ca
mi (nh n v ). Cc DNA polymerase c tnh c hiu cao, ch thm
nucleotide vo u 3OH ca mch ang tng hp.
Ngoi chc nng polymer ha theo hng 5 - 3, DNA polymerase III cn c
kh nng sa sai nh hot tnh exonuclease theo hng 3- 5. Exonuclease
l hot tnh enzyme ct DNA t u mt mt mch. Trn ng di chuyn
tng hp mch mi, nu gp ch nucleotide va lp sai v tr, DNA
polymerase III s dng hot tnh exonuclease 3- 5 ct li li b
nucleotide sai v lp ci ng vo ri tip tc ti bn (n v ).
Qu trnh ti bn DNA E.coli din ra vi tc nhanh, c th n 50.000
nucleotide/pht.

97/132

Hon chnh si mi tng hp

Trn mch chm, sau khi mi on DNA c ko di, mi RNA b di

i nh hot tnh exonuclease 53 ca DNA polymerase I v s b enzyme
RNase H phn hy. Cc ch trng m mi li s c thay bng trnh t
DNA mt cch chnh xc nh kt hp hot tnh polymerase 5 3 (ko di
u 3 ca on Okazaki trc) vi hot tnh exonuclease 3 5 (c v sa
sai) ca DNA polymerase I. Cui cng, ligase s ni tt c cc on DNA trn
mch mi tng hp li vi nhau.
Giai on kt thc ti bn v phn chia t bo

Hai chc ba ti bn s gp nhau khong 180o i din vi OricC. Quanh

vng kt thc ny c vi im lm dng li s ti bn bng cch gn vi mt
sn phm ca gen tus, l mt nhn t km hm hot ng helicase ca
Dna B.
Khi s ti bn hon tt, hai phn t DNA vng vn cn dnh vi nhau. Mt
topoisomerase loi II c tn l topoisomerase IV s tch ri chng sau
hai nhim sc th con s c phn phi vo hai t bo con.

98/132

Ti bn DNA eukaryote
S ti bn eukaryote vn cn nhng iu cha tng tn nhng cc d liu thu c
cho thy h thng ny kh gn vi h thng ti bn prokaryote. Khc bit ch yu l
cc loi DNA polymerase tham gia vo qu trnh.
im khi u ti bn v giai on khi u:
V s phc tp trong cu trc ca cht nhim sc, tc di chuyn ca cc chc ba ti
bn eukaryote ch t khong 50 bp/giy. Vi tc ny, nu ch s dng hai chc ba
ti bn, phi mt 30 ngy mi ti bn xong mt phn t DNA nhim sc th in hnh
ca ng vt hu nh vi kch thc 105 kb. Do cn phi c nhiu n v ti bn
trong mt t bo, in hnh nh ng vt hu nh l 50.000 100.000 n v ti bn.
eukaryote, c khong 20 50 n v ti bn khi u cng lc ti mi thi im,
xuyn sut phase S. Phn ti bn sm nht ch yu bao gm nguyn nhim sc cht
(euchromatin), tc l phn DNA c hot ng phin m. Trong khi , d nhim sc cht
(heterochromatin) c hot ha mun hn v phn DNA ca tm ng (centromere)
v on cui nhim sc th (telomere) c ti bn sau cng. iu ny phn nh kh
nng p ng khc nhau ca nhng cu trc nhim sc cht vi nhn t khi u.
Cc trnh t khi u ng vt hu nh cha c phn lp nhng ngi ta tin rng
s khi u ca mi n v ti bn c l din ra ty bn trong mt vng khi u c
th di n vi kb v l mt phn ca trnh t DNA lp li trung bnh.
Khc vi prokaryote, cc n v ti bn ca eukaryote ch khi u mt ln trong mi
chu trnh t bo. Mt protein gi l nhn t cho php (licensing factor) cn thit cho s
khi u v ti hot ha ch c kh nng tip cn vo nhn khi mng nhn tan i, v th
ngn chn c vic ti khi u trc hn nh.

99/132

Chc ba ti bn v h thng cc enzim ti bn

Trc khi ti bn, DNA phi c tho khi nucleosome ti mi chc ba ti bn. iu
ny lm chm s di chuyn ca chc ba, ch cn 50 bp/giy. Sau khi chc ba ti bn
i qua, cc nucleosome c ti lp rp t cc histone c v histone mi c tng hp.
Kch thc nh ca cc on Okazaki (v d 135 bp SV40) phn nh lng DNA
c tho khi mi nucleosome khi chc ba ti bn i qua. Ngoi s nhn i DNA, cc
histone cng phi c nhn ln gp i trong sut phase S.

100/132

H thng cc DNA polymerase eukaryote phc tp hn so vi prokaryote, bao gm:

Polymerase /primase: C chc nng tng hp mi cho mch ti v cho c
nhng on Okazaki ca mch chm. Polymerase ny tip tc ko di DNA
nhng nhanh chng b thay bi polymerase trn mch ti v polymerase
trn mch chm. Polymerase khng c hot tnh exenuclease.
Polymerase : C chc nng ging DNA polymerase I prokaryote, ngha l
tng hp i km vi sa sai v hon chnh mch mi sau khi mi RNA c
loi b.
101/132

 Polymerase v polymerase : C chc nng ko di DNA. Trong kh

nng tng hp on DNA di nht thuc v polymerase vi s tr gip ca
PCNA. C hai enzyme ny u c kh nng c v sa sai.
Polymerase ?: c tm thy trong ti th, chc nng cha r.
Ngoi cc polymerase k trn, h thng ti bn eukaryote cn c s tham gia ca nhiu
protein chuyn bit nh PCNA (Proliferating Cell Nuclear Antigen - khng nguyn
trong t bo ang phn chia) c chc nng hot ha cc polymerase v , cc nhn
t ti bn A v C (Replication Factor, RF -A, RF - C) cn cho hot ng ca cc
polymerase v ...
Ti bn on cui nhim sc th (telomere replication):
Hai on cui ca nhim sc th dng thng khng th c ti bn y bi v khng
th ko di DNA t u 5 ca mch mi tng hp sau khi loi b mi RNA. Do
thng tin di truyn s mt dn khi DNA sau mi t ti bn.
gii quyt vn ny, on cui mi nhim sc th eukaryote bao gm hng trm
bn sao mt trnh t n gin, khng m ha (v d nh TTAGGG ngi) vi u 3
nh ra so vi u 5.
Telomerase l mt enzyme duy tr di ca telomere. Enzyme ny c cha mt phn
t RNA ngn. Mt phn trnh t ca RNA ny b sung vi trnh t lp ca telomere, do
n c th hot ng nh mt mch khun ko di cc trnh t lp t u 3 t do
ca telomere bng cch tng hp ri dch chuyn lin tip nhiu ln.
Cc eukaryote n bo, chng hn nh nm men, nh vo hot ng ca telomerase m
duy tr telomere, t bo v t bo khi s mt thng tin di truyn. Th nhng cc
sinh vt eukaryote a bo, cc t bo sinh dng, gen m ha telomerase b km hm.
V th, qua mi t phn bo, cc nhim sc th ngn dn i cho n khi chm ti cc
trnh t m ha ca DNA, t bo s gi i v cht. Trong nui cy t bo sinh dng,
hu ht cc t bo ch phn chia 30-40 hoc 50 ln ri cht.
cc t bo ung th, s ti hot ha hot ng ca telomerase khin cho cc t bo ny
c kh nng bt t.

102/132

103/132

Nhim sc th
Nhim sc th prokaryote
Nhim sc th prokaryote gm chui xon kp DNA dng vng, cun li mt cch
tinh vi hay siu xon, tng tc vi cc phn t protein. Cc protein ny c chc nng
n nh cu trc ca nhim sc th
VD: E.coli, cc protein HU (positive charged & dimeric protein), H-MS ging nh
histon Eukaryote c vai tr trong nn DNA. Cc protein khc nh IHF (integration
host factor, RNA polymerase v RNA c vai tr trong t chc DNA
prokaryote, nhim sc th nn cht hnh thnh vng nhn v nh trn mng nguyn
sinh.

104/132

DNA ca vi khun th xon (a), vng trn (b) v siu xon (c)

Nhim sc th Eukaryote
Nhim sc th ca Eukaryote c cu to t si nhim sc (chromatin) l phc hp
gia DNA v protein. Protein ca nhim sc th: Histon v nonhiston chim khong
80% trong lng nhim sc th.
Histon protein c thnh phn amino acid tch in dng (base) chng hn nh
Arginine, Lysine v Histidine cao. Histon tham gia vo cu trc chui nucleosome v
solenoid. C nm loi protein histon: H2A, H2B H3, H4 v H1.
Nonhiston protein c thnh phn amino acid tch im m (acid) cao. Protein ny tham
gia vo ng xon DNA hnh thnh cc bc cu trc cao hn, sao chp DNA (DNA
polymerase), phn ly nhim sc th (protein ng c trong trung tm 0 ca tm ng)
v tham gia vo qu trnh sao m, iu ho biu hin gene.
n v cu trc c bn ca nhim sc th l cc nucleosme c ng knh 100
Anstrong. Nucleosome c hnh thnh do mt chui DNA khong 146 bp qun quanh
mt li protein gm 8 phn t histon (2H2A, 2H2B, 2H3 v 2H4). Hai nucleosome c
ni vi nhau bi mt on DNA khong 55 bp.
Chui nucleosome tip tc xon hnh thnh cu trc phc tp hn l solenoid c
ng knh 300 Anstrong. Mi solenoid c khong 60 nucleosome c n nh bi
Histon H1. Khi H1 c gii phng solenoid tr v dng nucleosome.
Solenoid tip tc xon hnh thnh cc si c ng knh 300nm, 700nm (chromatin) v
1400nm (chromosome) di s h tr ca cc protein nonhiston.

105/132

Cc mc ng xon ca chromatin
Cht nguyn nhim sc: l cht nhim sc trng thi xon, trng thi hot ng.
Cht d nhim sc: l cht nhim sc cun xon cao nht, trng thi khng hot ng.

Tnh c trng ca b nhim sc th

Mi loi sinh vt eukaryote u c b nhim sc th c trng v s lng, hnh thi v
cu trc. y l tnh c trng phn bit cc loi vi nhau, khng phn nh trnh
tin ho cao hay thp.
nhng loi giao phi, t bo sinh dng (soma) mang b nhim sc th lng bi (2n)
ca loi. Trong , nhim sc th tn ti thnh tng cp tng ng (homologous), mt
chic c ngun gc t b, mt chic c ngun gc t m. Trong giao t cha b nhim
sc th n bi (n), mi nhim sc th ch c mt chic.
106/132

Bng cc k thut t bo hc hin i, cn c cc mt chc nng, cu trc, hnh

thi v tnh c th trong hot ng, ngi ta phn bit cc loi nhim sc th khc
nhau:
- Nhim sc th thng (nhim sc th A - autosome): ging nhau c 2 gii
c, ci.
- Nhim sc th gii tnh (sex chromosome): khc nhau gia 2 gii c v ci.
VD: Ngi c 46 nhim sc th gm 22 cp nhim sc th thng v 1 cp nhim sc
th gii tnh. Nhim sc th gii tnh n l XX v nam l XY.
- Nhim sc th ph (nhim sc th B): c pht hin mt s ging thc vt nh ng,
la mch en cha qua chn lc, s lng nhiu hay t ty dng. Nhng cy c nhim
sc th B th yu hn v hu th km hn so vi cc cy ch cha nhim sc th A
bnh thng. V d, la mch en, nhng cy c ti 9 nhim sc th B thng khng
c kh nng sng. Nhim sc th B c kch thc nh v hiu qu di truyn thp. Nhim
sc th B cng c bt gp mt s ng vt nh su b, giun dp.
S lng nhim sc th trong b lng bi n nh i vi mi loi nhng khng mang
tnh c trng cao, chng hn nh g (Gallus gallus), ngan (Cairina moschata), vt nh
(Anas phatyryncha) u c 2n = 80. Tnh c trng ch th hin r trong s lng, thnh
phn, trnh t phn b cc gen trn mi nhim sc th v cc c im hot ng ca
nhim sc th trong ti bn, phn ly, t hp, trao i on, t bin.

Hnh thi nhim sc th

Trong nhn t bo, cht nhim sc tn ti thng xuyn di dng si nhim sc mnh,
kh quan st. Khi bc vo phn bo, si nhim sc bt u ng xon v t nn
cc i k gia. Lc ny, nhim sc th (chromosome) dy hn v dng kp gm
hai nhim sc t (chromatid) nh nhau tm ng (centromere); chng c hnh dng
v kch thc c trng nn c th quan st v m s lng thng qua mt knh hin
vi quang hc.

107/132

Nhim sc th vi vng tm ng
Mi nhim sc th c mt tm ng, l im tht eo chia nhim sc th thnh
2 vai vi chiu di khc nhau, vai ngn hn gi l vai p v vai di hn gi l vai q. Da
vo v tr ca tm ng c th phn bit hnh thi cc nhim sc th:
- Tm gia (metacentric): 2 vai bng nhau.
- Tm u (acrocentric): 2 vai khng bng nhau.
- Tm mt (telocentric): tm ng nm gn cui.

108/132

S cc kiu hnh thi nhim sc th k gia v k sau

mt s t chc, c quan ca mt s loi thng xut hin cc nhim sc th c hnh
thi c bit nh nhim sc th khng l (polytene chromosome), nhim sc th chi
n (lambrush chromosome):
Nm 1981, E. Balbiani pht hin nhim sc th khng l tuyn nc bt u
trng Chironomus. n nay, loi nhim sc th ny c tm thy trong t bo ca
tuyn nc bt, tuyn Manpighi, mng rut mt s cn trng b 2 cnh (Diptera) nh:
Drosophilidae, Chironomidae.
Nhim sc th khng l c s lng si nhim sc gp nhiu ln so vi nhim sc th
thng, c th cha ti 150 - 1600 si. Nguyn nhn ca hin tng ny l do c ch
ni nguyn phn (endomitosis). Nhim sc th t nhn i nhiu ln nhng khng phn
ly, to nhim sc th c dng chm nhiu si, b ngang ca nhim sc th tng ln. Do
khng ng xon nn chiu di ca nhim sc th khng l c th t ti 250-300 m,
gp 100-200 ln chiu di nhim sc th thng. Dc theo chiu di ca nhim sc th
khng l phn ha thnh nhng khoanh bt mu m, nht khng ng nht nh cc a
sng, ti xen nhau. Ngi ta cho rng cc a sm mu l ni tch ly nhiu DNA, c
to ra do xon nh khu dy c hoc do tp trung nhiu ht nhim sc.
rui gim, NST khng l tuyn nc bt c hnh thnh do DNA t nhn i 10
ln, to ra 210 = 1024 si dnh lin nhau sut dc theo chiu di.

(a) Nhim sc th khng l cua rui gim tao im nhim sc (chromocenter).

(b) B nhim sc th c ban trong t bao ang phn chia vi cac nhanh c biu hin
bng cac mau khac nhau.

109/132

(c) Anh chup nhim sc th khng l

Nhim sc th khng l cua rui gim

Nhim sc th chi n: Nhim sc th ny c th di n 800 m, c k u ca gim
phn trong t bo trng ca ng vt c xng sng nht l giai on Diplotene ca
trng c nhiu non hong (trng g, chim hoc b st). c im ca nhim sc th
kiu chi n l t trc ca nhim sc th c nhiu vng DNA, cnh cc vng DNA ny
l nhng loi ARN c tng hp t cc vng DNA m xon.

Nhim sc th chi en

110/132

Kiu nhn v nhim sc

Do s n nh v hnh thi ca mi nhim sc th v s c nh v s lng
nhim sc th ca mi loi nn mi loi c mt kiu nhn c trng. Kiu nhn
(karyotype) l s m t hnh thi ca b nhim sc th. Kiu nhn c th c biu th
dng nhim sc (Idiogram) khi cc nhim sc th c xp theo th t bt u t
di nht n ngn nht.
Sau ny k thut nhum mu (mu giemsa hay quinacrin) c hon chnh, lm
r hn cc vt c trng th hnh thi ca mi nhim sc th c xc nh chi tit hn.
Da vo nhim sc nhum mu, c th tm thy cc on tng ng trn cc nhim
sc th cng loi ca cc loi c h hng gn nhau. V d so snh nhim sc ca
ngi v vn cho thy c mi quan h h hng rt gn v nhim sc th th hai ca
ngi do s ni li ca 2 nhim sc th khc nhau vn ngi.

Cp nhim sc th tng ng v nhim sc ca ngi

111/132

S phn bo
Phn bo prokaryote
Phn bo prokaryote l trc phn (binary fission). Mt t bo prokaryote sau mt ln
phn bo trc phn to hai t bo con ging nhau.
Trong phn bo trc phn, nhim sc th ca prokaryote nhn i v nh trn mng t
bo ti mt cu trc gi l mesosome (cc np gp ca mng t bo). Thnh t bo xut
hin hnh thnh vch ngn, tch i hai nhim sc th v chia t bo m thnh hai t bo
con (daughter cell). Mi t bo con mang mt b gene hon chnh.

Phn bo ca t bo vi khun

112/132

Chu trnh t bo v s phn bo eukaryote

Chu trnh t bo
Khi nim
Chu k t bo, hay chu k phn bo, l mt vng tun hon cc s kin xy ra trong
mt t bo eukaryote t ln phn bo ny cho n ln k tip. Chu k t bo c
iu khin bi nhiu loi cyclin v cdk (mt loi kinase ph thuc cyclin). Leland H.
Hartwell, R. Timothy Hunt v Paul M. Nurse t gii Nobel trong lnh vc Sinh l
v Y hc nm 2001 v nhng pht hin ca h v vai tr trung tm ca nhng phn t
ny trong chu k t bo.
Chu trnh t bo gm hi gia on: k trung gian (interphase) v k phn chia (mitosis).

Chu trnh t bo
Mt chu trnh t bo ca eukaryote gm 4 phase:

G1 l phase di nht. Trong , t bo chun b cho ti bn DNA.

S l phase duy nht trong chu trnh t bo c s ti bn DNA.
G2 l mt phase ngn trc khi nguyn phn.
M (Mitosis) l phase nguyn phn, gm s phn chia nhim sc th v phn
chia t bo

G1, S v G2 l ba phase ca k trung gian (interphase)

Phase M gm 4 k: k trc (prophase), k gia (metaphase), k sau (anaphase), k
cui (telophase).

113/132

Sau phase M, t bo con bc vo phase G1 ca chu trnh k tip. Cc t bo cng c

th thot khi chu trnh k tip v ri vo tnh trng khng sinh sn gi l G0.
iu ha chu trnh t bo
Thi gian v t l ca s phn bo trong nhng phn khc nhau ca thc vt v ng vt
l rt quan trng cho s sinh trng, pht trin v tn ti bnh thng. Mc thng
xuyn ca s phn bo khc nhau ty loi t bo, chng hn t bo da ngi phn bo
thng xuyn, t bo gan ch phn chia khi cn thit cn t bo thn kinh ngi trng
thnh th khng phn chia na.
S khc bit trn l kt qu ca qu trnh iu ha chu trnh t bo mc phn t. H
thng iu ha chu trnh t bo gm cc checkpoint. Mt checkpoint trong chu trnh t
bo l ni m tn hiu cho php tin trnh phn bo tip tc hay dng. C ba checkpoint
quan trng trong chu tnh t bo: checkpoint G1, G2 v M phase.

Nhng phn t tham gia iu ha chu trnh t bo l cc protein hay enzyme hot ha
hay c ch cc protein khc bi s phosphoryl ha. hiu r c ch iu ha ny ta
xem xt c ch iu ha checkpoint G2 ca Cdk (cyclin-dependent kinase) mt kinase
ph thuc vo cyclin. Hot ng ca Cdk lm thay i nng ca cyclin trong t bo.
Cdk tn ti trong t bo trng thi bt hot. Khi cyclin tch ly trong t bo trong
pha G2, n s kt hp vi v hot ha Cdk hnh thnh phc hp Cyclin-Cdk. CyclinCdk c pht hin u tin l MPF (maturation promation factor). MPF phosphoryl
114/132

nhiu protein khc, chng hn nh phosphoryl ha mng nhn, kch thch cc kinase
khc phosphoryl ha cc protein khc ca mng nhn t gip t bo vt qua
checkpoint G2 v tin vo pha phn bo (M phase).
Cui pha M, enzym phn gii cyclin, nh vy lm bt hot Cdk. Cdk tn ti trong t bo
cho n khi kt hp vi cyclin mi. Nhng enzyme ny cng lin quan n vic gip
chu trnh t bo vt qua im checkpoint M. C t nht ba protein Cdk v nhiu cyclin
lin quan n vic gip t bo vt qua im checkpoint G1. Nh vy, hot ng tng
gim ca cc phc hp cyclin v Cdk c th kim sot tt c cc giai on ca chu trnh
t bo.
Phn bo nguyn nhim (Nguyn phn - Mitosis)
K trung gian
Chim 90% thi gian ca chu trnh t bo. Trong sut k trung gian t bo sinh trng,
nhn i nhim sc th chun b cho s phn chia t bo. K trung gian gm 3 giai on
(phase): G1 (first gap), S (synthesis), G2 (second gap).
Trong sut k trung gian, trung th (centrosome) nhn i. Mi trung th mang hai trung
t (centriols). Sao thoi v sc hnh thnh. Nhim sc th tho xon, nhn i thnh
nhim sc th kp gm hai nhim sc t ch em (sister chromatid) dnh nhau tm ng.
Phn bo nguyn nhim
Gm giai on phn chia nhim sc th v phn chia t bo cht (cytokinesis).
Phn chia nhim sc th:
K trc (prophase)
Nhim sc th bt u xon li, c th quan st bng knh hin vi quang hc. Nhim sc
th vn dng kp.
Hai trung th ( t bo ng vt) di chuyn v hai cc t bo. Thoi v sc (mitotic
spindle) hnh thnh. Hch nhn bin mt.
K gia (metaphase)
u k gia mng nhn tan bin. Si v sc di ra, xuyn qua nhn tng tc vi nhim
sc th. Mt vi si v sc gn vi tm ng ca nhim sc th ti kinetochore. Hai
kinetochore ca cp chromatid ch em gn vi hai si v sc n t hai cc i din.
Cc si v sc khng gn vi kinetochore tng tc vi nhau.

115/132

Cui k gia, nhim sc th kp ng xon cc i tp trung mt phng xch o ca

thoi v sc.
K sau (anaphase)
Hai chromatid ch em tch nhau ra tm ng v mi si chromatid by gi gi l
nhim sc th n di chuyn v hai cc ca t bo. Si v sc thu ngn li.
K cui (telophase)
hai cc t bo, mng nhn xut hin v hnh thnh hai nhn ging nhau. Cui k
nhim sc th dui xon.

116/132

Phn bo (cytokinesis):
Cui k cui xy ra s phn chia t bo cht.
t bo ng vt, mt rnh phn chia xut hin trn b mt t bo gn mt phng xch
o, sau n su vo do tc ng co rt ca vng vi si actin bn trong t bo cht, ct
t bo m thnh hai t bo con (daughter cells) ging nhau. Trong khi t bo thc
vt khng xut hin rnh phn ct. Nhng ti (vesicle) cha nguyn liu xy dng vch
t bo t Golgi di chuyn ti gia t bo v t chc thnh a t bo (cell plate) chia t
bo m thnh hai t bo con.

117/132

Phn chia t bo cht t bo ng vt v thc vt .

ngha ca nguyn phn
Nguyn phn l c s ca s tng trng sinh vt a bo v s sinh sn v tnh.
Qua nguyn phn, cc th h t bo trong mt c th a bo cng nh cc th h c th
ca loi sinh sn v tnh c duy tr mt b nhim sc th (2n) c trng.
Phn bo gim nhim (Gim phn - Meiosis)
L qu trnh phn bo chuyn bit xy ra t bo sinh dc, trong s lng nhim sc
th gim i mt na. Gim phn gm hai ln phn chia: ln phn chia th nht l phn
chia gim nhim, ln phn chia th hai l phn chia nguyn nhim. Mt t bo sinh dc
qua gim phn to ra 4 t bo con vi b nhim sc th gim i mt na.

118/132

So snh gia gim phn v nguyn phn

Gim phn I
Gm 4 k
K trc I
Cc s kin xy ra trong gim phn I tng t trong nguyn phn. Tuy nhin, hin tng
tip hp gia cc cp nhim sc th tng ng v s bt cho trao i on gia cc
chromatid ch xy ra gim phn.
K gia I
Cc nhim sc th kp tp trung theo cp tng ng trn mt phng xch o ca thoi
v sc. Mi nhim sc th kp trong cp tng ng ch gn vi mt si v sc ti mt
kinetochore ca n.
K sau I

119/132

Si v sc thu ngn li. Cp nhim sc th kp phn ly, mi chic kp i v mt cc t

bo.
K cui I v phn chia t bo cht
Hai t bo con hnh thnh, mi nhn t bo mang mt b nhim sc th n bi. Nhim
sc th vn trng thi kp.

120/132

Gim phn II
Tip theo k cui ca gim phn I l giai on chuyn tip, tng t nh k trung gian
nhng khng c s nhn i nhim sc th.
K trc II
B nhim sc th kp (n bi) tin ti mt phng xch o ca t bo.
K gia II
Nhim sc th kp ng xon cc i tp trung mt phng xch o ca thoi v sc.
Hai kinetochore ca cp chromatid ch em gn vi hai si v sc n t hai cc i din.
K sau II
Hai chromatid ca nhim sc th kp tch nhau ra tm ng, mi chromatid phn ly
v mt cc ca t bo.
K cui II v phn chia t bo
Mi nhim sc th v n cc t bo, mng nhn hnh thnh. Phn chia t bo cht xy
ra.
Kt thc gim phn, mt t bo sinh dc (2n) hnh thnh 4 t bo con. Mi t bo con
c b nhim sc th gim i mt na (n).
ngha ca gim phn
Gim phn l c s ca s sinh sn hu tnh v lm tng s a dng di truyn.
Thng qua gim phn, mt t bo sinh dc chn vi 2n nhim sc th c th to 2n loi
giao t. VD: ngi t bo sinh dc c 2n = 46 nhim sc th, qua gim phn c th
hnh thnh 223 loi giao t.
S trao i cho dn n hon v gene k trc gim phn I cng lm tng s loi giao
t.
S kt hp ngu nhin ca cc loi giao t trong th tinh lm tng s loi hp t. VD:
S lng loi hp t c th hnh thnh ngi l 223 x 223 .

121/132

Tm li: S phn ly c lp ca nhim th, s trao i cho ca cc chromatid v s th

tinh ngu nhin gia cc loi giao t l ba ngun gc ca s a dng di truyn.

Phn li c lp ca nhim sc th

122/132

123/132

Gene v m di truyn
Khi nim v gene
Mendel (Mitchell, 1999) nhn t di truyn l mt n v di truyn tch bit nh hng
mt tnh trng.
Khi nim gene c Wilhelm Johannsen ra ln u tin vo nm 1909 nhng ng
ch mi quan nim mt cch s lc rng y l nhng mm mng c bit, di truyn
tch bit nhiu tnh trng ca c th.
Morgan (Mitchell, 1999): gene (locus) l n v di truyn khng th chia nh nm
trn nhim sc th. Khi nim ny bao hm:
Gene l n v chc nng: Cc phn ca gene nu nh tn ti th khng th hot ng
chc nng. t bin ca mt gen s nh hng n mt chc nng di truyn.
Gene l n v ti t hp: S trao i cho ch c th din ra gia cc gene, khng th
xy ra trong gene.
Gene l n v t bin: Gene s b bin i nh mt tng th hon chnh trc cc tc
nhn t bin liu lng.
Benzer (Hong, 1995) nghin cu phage T4 k sinh trn E.coli thy rng trao i cho
v t bin c th xy ra ngay trong mt gen v trn gen c th c hn mt v tr trao
i cho v t bin xy ra. Nh vy mi gen c nhiu n v ti t hp (recon-t
recombination) v t bin (muton- c gc t mutant). Mt on phn t DNA xc nh
cu trc ca mt chui polypeptide c gi l gen cu trc hay citron (gc t cistrans), trung bnh c t 500-1000 cp nucleotide, trong cha cc muton v recon.
Thp nin 1940, thuyt mt gen - mt enzyme ca George Beadle v Edward Tatum
chng minh: gen kim tra cc phn ng sinh ha. Hc thuyt trn chng t rng gen
chi phi cu trc protein. Sau , ngi ta bit rng c nhng protein gm nhiu chui
polypeptide do nhiu gen quy nh (v d nh hemoglobin) nn thuyt ny c chnh
li l 1 gen 1 polypeptide.
Thp nin 1950, m hnh cu trc ADN do James Watson v Francis Crick xut v
Hc thuyt trung tm ra i. Gen c hiu l mt on ADN trn nhim sc th m
ha cho mt polypeptide hay mt ARN.
Cui nhng nm 1970, ngi ta pht hin thy eukaryote c nhng on ADN khng
m ha cho cc acid amin trn phn t protein. Trong khi ton b DNA prokaryote u
124/132

mang thng tin m ha cho cc protein, th eukaryote, phn t DNA bao gm cc trnh
t m ha gi l cc exon xen k vi cc trnh t khng m ha gi l intron. Mt phn
t mRNA va mi c phin m t mt gen cu trc th cha c hot tnh; phn t
ny phi tri qua mt qu trnh ghp ni, loi b cc intron tr thnh phn t mRNA
trng thnh c kh nng tham gia dch m. Khi nim gen c chnh l nh sau: Gen
l mt on ADN m bo cho vic to ra mt polypeptide, n bao gm c vng trc,
vng sau v vng m ha cho protein, c nhng on m ha (exon) ln nhng on
khng m ha (intron) ca eukaryote.

(a)

(b)
Cu trc ca mt gene m ha cho mt polypeptide
Nh vy hin nay, c th nh ngha tng qut v gen nh sau:

125/132

Gen l n v chc nng c s ca b my di truyn, chim mt locus nht nh trn

nhim sc th. Gen l nhng on vt cht di truyn m ha cho mt i phn t sinh
hc nh ARN hoc polypeptide.

M di truyn
Nh bit, c mi lin h gia DNA, RNA v protein. DNA v RNA cu to t 4 loi
nucleotide trong khi protein c hnh thnh t 20 loi amino acid. Nh vy, s t hp
ca 4 loi nucleotide s m ha cho 20 loi amino acid. Nu t bo s dng 2 nucleotide
m ha cho 1 amino acid th 4 loi nucleotide ch to c 42 = 16 amino acid khng
cho nhu cu thc t. Nh vy, n v m ha cho mt amino acid hay cn gi l codon
phi t 3 nucleotide tr ln.
Nhng nghin cu trong thp 60 chng minh c rng t hp 3 nucleotide c th
m ha cho mt amino acid v xc nh cc codon m ha cho tng loi amino acid.
Nh vy 4 loi nucleotide hnh thnh 43 = 64 codon trong c 3 codon khng m ha
cho bt k amino acid no, l cc codon kt thc UAA, UGA, UAG v mt codon
m u AUG m ha cho methionin.

126/132

Bng m di truyn (mRNA)

M di truyn l m b ba (triplet), khng gi ln nhau (non-overlapping) v c c
mt cch lin tc, khng ngt qung (comma-less).
M di truyn c tnh thoi ha (degenerate) ngha l nhiu codon cng xc nh mt acid
amin, tr hai ngoi l: AUG m ha cho Met v UGG m ha cho Trp.
M di truyn c tnh ph bin (universal), ngha l thng nht cho hu nh ton b sinh
gii. Tuy nhin, t nm 1980 ngi ta pht hin thy ty th ca mt s loi eukaryote
v mt s sinh vt n bo c s bin i ngha ca mt s m di truyn.

127/132

Tham gia ng gp
Ti liu: Gio trnh sinh hc i cng
Bin tp bi: Nguyn Hi
URL: http://voer.edu.vn/c/3207fce7
Giy php: http://creativecommons.org/licenses/by/3.0/
Module: S hnh thnh tri t v kh quyn
Cc tc gi: Nguyn Hi
URL: http://www.voer.edu.vn/m/29045681
Giy php: http://creativecommons.org/licenses/by/3.0/
Module: Ngun gc ca s sng
Cc tc gi: Nguyn Hi
URL: http://www.voer.edu.vn/m/726733d6
Giy php: http://creativecommons.org/licenses/by/3.0/
Module: S tin ha ca t bo
Cc tc gi: Nguyn Hi
URL: http://www.voer.edu.vn/m/fd8e75a4
Giy php: http://creativecommons.org/licenses/by/3.0/
Module: Hc thuyt t bo v cc phng php nghin cu t bo hc
Cc tc gi: Nguyn Hi
URL: http://www.voer.edu.vn/m/0446df5e
Giy php: http://creativecommons.org/licenses/by/3.0/
Module: Thnh phn ha hc ca t bo
Cc tc gi: Nguyn Hi
URL: http://www.voer.edu.vn/m/9f76953b
Giy php: http://creativecommons.org/licenses/by/3.0/
Module: T bo Eukaryote
Cc tc gi: Nguyn Hi
URL: http://www.voer.edu.vn/m/0dfad704

128/132

Giy php: http://creativecommons.org/licenses/by/3.0/

Module: Cu to ca t bo Prokaryote
Cc tc gi: Nguyn Hi
URL: http://www.voer.edu.vn/m/df1b2a70
Giy php: http://creativecommons.org/licenses/by/3.0/
Module: Cc qu trnh sinh hc trong t bo
Cc tc gi: Nguyn Hi
URL: http://www.voer.edu.vn/m/1156e2d7
Giy php: http://creativecommons.org/licenses/by/3.0/
Module: S a dng ca t bo
Cc tc gi: Nguyn Hi
URL: http://www.voer.edu.vn/m/a8cdeffd
Giy php: http://creativecommons.org/licenses/by/3.0/
Module: Khi nim v cu trc Enzyme
Cc tc gi: Nguyn Hi
URL: http://www.voer.edu.vn/m/a8acd084
Giy php: http://creativecommons.org/licenses/by/3.0/
Module: C ch hot ng,phn loi v cc yu t nh hng n hot tnh Enzyme
Cc tc gi: Nguyn Hi
URL: http://www.voer.edu.vn/m/943c33c2
Giy php: http://creativecommons.org/licenses/by/3.0/
Module: H hp t bo
Cc tc gi: Nguyn Hi
URL: http://www.voer.edu.vn/m/2a11404c
Giy php: http://creativecommons.org/licenses/by/3.0/
Module: Quang hp
Cc tc gi: Nguyn Hi
URL: http://www.voer.edu.vn/m/f383ae59
Giy php: http://creativecommons.org/licenses/by/3.0/

129/132

Module: Lch s pht trin ca di truyn hc

Cc tc gi: Nguyn Hi
URL: http://www.voer.edu.vn/m/df62a83b
Giy php: http://creativecommons.org/licenses/by/3.0/
Module: i tng v phng php nghin cu di truyn hc
Cc tc gi: Nguyn Hi
URL: http://www.voer.edu.vn/m/846e1528
Giy php: http://creativecommons.org/licenses/by/3.0/
Module: Quan h gia di truyn hc vi cc khoa hc khc v vi thc tin
Cc tc gi: Nguyn Hi
URL: http://www.voer.edu.vn/m/c68a3e01
Giy php: http://creativecommons.org/licenses/by/3.0/
Module: Acid Nucleic l vt cht di truyn cp phn t
Cc tc gi: Nguyn Hi
URL: http://www.voer.edu.vn/m/b2d973b8
Giy php: http://creativecommons.org/licenses/by/3.0/
Module: Ti bn DNA
Cc tc gi: Nguyn Hi
URL: http://www.voer.edu.vn/m/e4c96037
Giy php: http://creativecommons.org/licenses/by/3.0/
Module: Nhim sc th
Cc tc gi: Nguyn Hi
URL: http://www.voer.edu.vn/m/476649ef
Giy php: http://creativecommons.org/licenses/by/3.0/
Module: S phn bo
Cc tc gi: Nguyn Hi
URL: http://www.voer.edu.vn/m/9f0d8098
Giy php: http://creativecommons.org/licenses/by/3.0/
Module: Gene v m di truyn

130/132

Cc tc gi: Nguyn Hi
URL: http://www.voer.edu.vn/m/1657d6c5
Giy php: http://creativecommons.org/licenses/by/3.0/

131/132

Chng trnh Th vin Hc liu M Vit Nam

Chng trnh Th vin Hc liu M Vit Nam (Vietnam Open Educational Resources
VOER) c h tr bi Qu Vit Nam. Mc tiu ca chng trnh l xy dng kho
Ti nguyn gio dc M min ph ca ngi Vit v cho ngi Vit, c ni dung phong
ph. Cc ni dung u tun th Giy php Creative Commons Attribution (CC-by) 4.0
do cc ni dung u c th c s dng, ti s dng v truy nhp min ph trc
ht trong trong mi trng ging dy, hc tp v nghin cu sau cho ton x hi.
Vi s h tr ca Qu Vit Nam, Th vin Hc liu M Vit Nam (VOER) tr thnh
mt cng thng tin chnh cho cc sinh vin v ging vin trong v ngoi Vit Nam. Mi
ngy c hng chc nghn lt truy cp VOER (www.voer.edu.vn) nghin cu, hc
tp v ti ti liu ging dy v. Vi hng chc nghn module kin thc t hng nghn
tc gi khc nhau ng gp, Th Vin Hc liu M Vit Nam l mt kho tng ti liu
khng l, ni dung phong ph phc v cho tt c cc nhu cu hc tp, nghin cu ca
c gi.
Ngun ti liu m phong ph c trn VOER c c l do s chia s t nguyn ca cc
tc gi trong v ngoi nc. Qu trnh chia s ti liu trn VOER tr ln d dng nh
m 1, 2, 3 nh vo sc mnh ca nn tng Hanoi Spring.
Hanoi Spring l mt nn tng cng ngh tin tin c thit k cho php cng chng d
dng chia s ti liu ging dy, hc tp cng nh ch ng pht trin chng trnh ging
dy da trn khi nim v hc liu m (OCW) v ti nguyn gio dc m (OER) . Khi
nim chia s tri thc c tnh cch mng c khi xng v pht trin tin phong
bi i hc MIT v i hc Rice Hoa K trong vng mt thp k qua. K t , phong
tro Ti nguyn Gio dc M pht trin nhanh chng, c UNESCO h tr v c
chp nhn nh mt chng trnh chnh thc nhiu nc trn th gii.

132/132