Sinh Hoc Dai Cuong

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B Y T
SINH HC I CNG
DNG CHO O TO DC S I HC
M s: .20.X.06
NH XUT BN GIO DC
H NI 2008
Ch o bin son:
file://C:\Windows\Temp\cacyqucjtt\Introduction.htm
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Ch o bin son:
V KHOA HC V O TO - B Y T
Ch bin:
PGS.TS. CAO VN THU
Nhng ngi bin son:
ThS. TRN TRNH CNG
ThS. NGC QUANG
PGS.TS. CAO VN THU
Th k bin son:
PGS.TS. PHAN TH HOAN
Tham gia t chc bn tho:
ThS. PH VN THM
TS. NGUYN MNH PHA
Bn quyn thuc B Y t (V Khoa hc v o to)

M s: 7K787Y8 - DAI
922 2008/CXB/7 1873/GD
Li gii thiu
Thc hin mt s iu ca Lut Gio dc, B Gio dc & o to v B Y t ban hnh ch
trnh khung o to Dc s i hc. B Y t t chc bin son ti liu dy hc cc mn c s
chuyn mn theo chng trnh trn nhm tng bc xy dng b sch t chun chuyn mn trong cng
tc o to nhn lc y t.
Sch Sinh hc i cng c bin son da vo chng trnh gio dc ca Trng i hc D
H Ni trn c s chng trnh khung c ph duyt. Sch c tp th cc nh gio giu kinh
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H Ni trn c s chng trnh khung c ph duyt. Sch c tp th cc nh gio giu kinh
nghim ca Trng i hc Dc H Ni bin son theo phng chm: kin thc c bn, h thng; n
dung chnh xc, khoa hc; cp nht cc tin b khoa hc, k thut hin i v thc tin Vit Nam.
Sch Sinh hc i cng c Hi ng chuyn mn thm nh sch v ti liu dy hc chuyn
ngnh dc s i hc ca B Y t thm nh nm 2007. B Y t quyt nh ban hnh ti liu dy
t chun chuyn mn ca ngnh trong giai on hin nay. Trong thi gian t 3 n 5 nm, sch phi
chnh l, b sung v cp nht.
B Y t chn thnh cm n cc tc gi dnh nhiu tm huyt hon thnh cun sch; cm n GS.
TS. Nguyn Vn Thanh, PGS. TS. L Hng Hinh c v phn bin cun sch sm hon thnh, k
thi phc v cho cng tc o to nhn lc y t.
Ln u xut bn, chng ti mong nhn c kin ng gp ca ng nghip, cc bn sinh vin v
cc c gi ln xut bn sau sch c hon thin hn.
V KHOA HC V O TO - B Y T
Li ni u
Ti liu Sinh hc i cng v c bn bao gm ni dung chnh ca nhng bi ging cho cc lp sinh vin
dc c tp hp trong mt s nm ging dy. Cng ngh sinh hc ngy nay (thi k hu gii m genom
ngi) thay i rt mnh m, nn bn cnh nhng kin thc c bn cn thit cho sinh vin ngnh khoa h
Dc cc tc gi c gng cp nht nhng kin thc khoa hc mi nht lin quan ti Sinh hc phn t, Cng
ngh sinh hc Dc. Ti liu ny c bin son cho sinh vin, hc vin nm th nht i hc Dc. N
dung bao gm:
Cu trc ca t bo tin nhn v t bo nhn chun, s vn chuyn vt cht qua mng.
Qu trnh trao i cht trong t bo, enzym, qu trnh h hp, qu trnh quang hp.
Vt cht di truyn, qu trnh sao chp ADN, di truyn nhim sc th v ngoi nhim sc th, bin d.
H thng phn loi 5 gii ca Whittaker.
Vi thi lng 2 n v hc trnh, c nhng vn c bn ch nu ln khi nim. V cu trc, ti liu ny
chia lm 4 chng chnh:
Chng 1: T bo.
Chng 2: S trao i cht v nng lng.
Chng 3: Di truyn v bin d.
Chng 4: Ngun gc s sng v a dng sinh hc.
Ti liu ny c th tip thu c mt cch d dng khi ngi hc nm vng c cc thut ng, gii h
c vn , bit so snh cc khi nim v qu trnh cng nh tham kho trc v tch cc tham gia vo bi
ging.
Vi tinh thn cu th v nht qun, hy vng rng nhng kin thc c trnh by trong ti liu Sinh h
i cng ny gp phn to nn nn tng kin thc tng i vng chc v lnh vc sinh hc cho cc dc s
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i cng ny gp phn to nn nn tng kin thc tng i vng chc v lnh vc sinh hc cho cc dc s
tng lai. Mc d c gng, nhng do thi gian c gii hn nn khng trnh khi nhng khim khuy
thiu st. Cc tc gi rt mong nhn c nhng kin ng gp ti liu ny hon thin hn trong cc l
ti bn sau. Mi kin xin gi v Cng ty C phn sch i hc Dy ngh, Nh xut bn Gio dc, 25 Hn
Thuyn, H Ni.
CC TC GI
DANH MC CH VIT TT
Adenin
ADN, DNA
Acid 2-deoxyribonucleic
ARN, RNA
Acid ribonucleic
Cytosin
dATP
2-deoxyadenosintriphosphat
dCTP
2-deoxycytidintriphosphat
dGTP
2-deoxyguanosintriphosphat
dTTP
2-deoxythymidintriphosphat
EST
u trnh t biu hin (expressed sequence tags)
Guanin
G1
Gap 1
G2
Gap 2
Nguyn phn (mitosis)
NST
Nhim sc th
PCR
Phn ng chui trng hp (polymerase chain reaction)
Tng hp (synthesis)
SSB
Protein lin kt si n (single strand binding protein)
Thymin
TMV
Virus khm thuc l (tobaco mosaic virus)
TTH
Trung tm hot ng
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Chng 1
T BO
MC TIU
1. Trnh by c c im, cu to v chc nng ca cc thnh phn c trong t bo Prokaryota: thnh
t bo, mng sinh cht, ribosom, th nhn, lng, roi, bao nhy.
2. Trnh by c c im, cu to v chc nng ca cc thnh phn c trong t bo Eukaryota: mng
sinh cht, li ni cht, th Golgi, lysosom, peroxysom, nhn, ribosom, ty th, lc lp, khung t bo,
trung t, lng, roi.
3. Trnh by c m hnh phn t phospholipid, qua gii thch c s hnh thnh ca lp mng kp.
4. Trnh by c cu to ca mng sinh cht theo m hnh khm ng. Gii thch c ti sao mng sinh
cht c tnh linh hot.
5. Trnh by c tnh cht v c ch vn chuyn vt cht qua mng theo phng thc c tiu ph nng
lng v khng tiu ph nng lng.
1. HC THUYT T BO
T thi nguyn thy, con ngi bit quan st v tm hiu th gii sinh vt bao quanh mnh. Du tch
ca nhng quan st vn cn lu li n ngy nay qua cc bc v c xa ca ngi tin s. Cho n th k
XVII, mt s kin quan trng gip qu trnh tm ti bc sang mt trang mi. l vo nm 1665, ln
u tin Rober Hook quan st th gii sinh vt bng knh hin vi t to c phng i 30 ln. ng
quan st m bn thc vt v thy rng cu trc ca chng c dng cc xoang rng c thnh bao quanh v
t tn l Cella (theo ting Latin, Cella c ngha l xoang rng hoc t bo). Nhng quan st ca Rober Hook
t nn mng cho mt mn khoa hc mi, l T bo hc.
Tip , n nm 1674, Antoni Van Leeuwenhoek vi knh hin vi c phng i 270 ln tin hnh
quan st v m t cc loi t bo ng vt (t bo mu, tinh trng v.v..) v xc nh rng t bo khng n
gin l cc xoang rng nh Rober Hook quan st trc y m c cu trc phc tp.
Cho n th k XIX, nh s hon thin ca k thut hin vi, cng vi tng kt t cng trnh nghin cu
trn cc lnh vc thc vt, ng vt, vi khun ca nhiu nh khoa hc khc nhau, hc thuyt t bo ra i.
Ni dung c bn ca hc thuyt bao gm hai sau:
Sinh vt c tnh a dng cao song u c cu to t t bo.
Mi t bo sng u c cu trc v chc nng tng t nhau.
Theo F.Engel (1870), hc thuyt t bo l mt trong ba pht kin v i ca khoa hc t nhin th k
XIX (cng vi hc thuyt tin ho v hc thuyt chuyn ho nng lng). T bo hc tr thnh mt khoa
hc tht s c lp v pht trin nhanh chng c v nghin cu cu trc v chc nng.
Theo thi gian, cu trc ca t bo ngy cng c nghin cu chi tit. T quan nim u tin l mt
xoang rng, v sau t bo c m t gm 3 phn l khi t bo cht (Purkinje, 1838 v Pholmon,
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1844), c gii hn bi mng t bo v bn trong c cha nhn (R.Brawn, 1831). Hng lot bo quan
trong t bo cht c pht hin v sau nh trung t (Van Beneden v Boverie pht hin vo nm 1876),
ty th (Altman v Benda, 1894), phc h Golgi (Golgi, 1898),...
Vic nghin cu cu trc cng cho thy tnh a dng ca t bo. Cng ging nh sinh vt, cc nhm t
bo c s phn ha v bin i thch nghi vi tng vai tr nht nh. V d, ngi c ti hn 200 loi t
bo khc nhau. Da vo c im nhn, ngi ta chia t bo ra thnh hai nhm ln l t bo Prokaryota v t
bo Eukaryota. Hai nhm t bo ny l n v t chc c bn ca tt c cc c th sng v phng din cu
trc v chc nng.
2. CU TRC T BO PROKARYOTA
T bo Prokaryota (hay cn gi l t bo nhn nguyn thu) c kch thc nh, ng knh khong 0,2 2,0m, chiu di khong 2,0 - 8,0m. y l dng t bo n gin, bn trong t bo cht hu nh khng c
cc bo quan (hnh 1.1). Vi khun l nhm sinh vt duy nht c cu trc t loi t bo ny.
Hnh 1.1. Cu trc t bo Prokaryota
2.1. Thnh t bo
Thnh t bo (hay cn gi l vch t bo) l lp ngoi cng, c rn chc nht nh bo v v duy
tr hnh dng t bo. Thng thng, nng cc cht tan trong t bo cao hn mi trng bn ngoi, bi vy
t bo hp thu kh nhiu nc. S thm thu ca nc s khin cho t bo b trng ln. Lc ny, thnh t
bo c vai tr quan trng trong vic nng , gip cho t bo khng b v di tc ng ca p sut thu
tnh. Tu theo cu to m thnh t bo c th gip chng chu mt p sut thm thu t 5atm ln n 20atm.
Khng ch c vai tr bo v v mt c hc, thnh t bo cn gip bo v t bo khi nhng tc ng ca
cc hp cht ho hc. V d, thnh t bo mt s vi khun c th gy cn tr s xm nhp ca cht khng
sinh vo bn trong t bo. Vi chc nng bo v nh vy, thnh t bo chnh l mt trong s cc ch tc
dng ca thuc i vi vi khun.
V mt cu to, thnh t bo vi khun c hai kiu cu to chnh tng ng vi hai nhm vi khun c tn
l Gram (+) v Gram (-). Ngi u tin xng phng php nhum phn bit hai nhm vi khun ny
l H.C.Gram, mt nh vi sinh vt hc an Mch. Theo phng php nhum Gram, vi khun Gram (+) s bt
mu xanh tm, cn vi khun Gram (-) s bt mu ta. Cu to khi qut ca hai loi thnh vi khun c th
tm lc nh sau:
Vi khun Gram (+): thnh t bo dy, gm mt lp, thnh phn tng i ng nht.
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Vi khun Gram (-): thnh t bo mng, gm nhiu lp, thnh phn v cu trc kh phc tp.
Thnh t bo c th bt gp c hai nhm t bo Prokaryota v Eukaryota vi nhng chc nng gn
ging nhau nhng thnh phn, cu to th khc nhau. Thnh t bo Prokaryota c c trng bi s c mt
ca mt thnh phn c tn l peptidoglycan (PG). y l mt loi polyme xp, khng tan, kh cng v bn
vng, bao quanh t bo nh mt mng li. Cu trc c bn ca peptidolgycan gm c 3 thnh phn: Nacetylglucosamin; N-acetylmuramic acid v chui acid amin. vi khun Gram (+), PG chim ti 50% trng
lng kh ca thnh t bo. Cn vi khun Gram (-), PG ch chim 5-10%, cn li l cc lipid, protein v.v..
Khng phi tt c cc t bo Prokaryota u c thnh t bo, v d nh mt s loi vi khun thuc h
Mycoplasma. h vi khun ny, lp ngoi cng s l mng t bo cht.
2.2. Mng t bo cht

Mng t bo cht (cytoplasmic membrane hay plasmamembrane) nm ngay di thnh t bo. Mng t
bo cht dy khong 5 - 10nm, c hnh thnh bi lp kp phospholipid. Chc nng ca lp mng ny c
th c tm lc nh sau:
Ngn cch t bo vi mi trng, gip t bo tr thnh mt h thng bit lp.
Thc hin qu trnh trao i cht, thng tin gia t bo v mi trng
L gi th gn cc enzym ca qu trnh trao i cht trong t bo.
2.3. T bo cht
T bo cht (cyloplasm) l vng dch th c gii hn bi mng t bo. T bo cht c cu to dng
keo, cha 80% l nc. c im quan trng to nn s khc bit vi t bo Eukaryota l t bo cht ca t
bo Prokaryota hu nh khng cha cc bo quan. Ton b t bo cht l mt khi thng nht, khng phn
ho thnh vng chc nng. Cc hot ng sng ca t bo u din ra chung trong t bo cht m khng c
s phn nh ranh gii r rt. nhiu loi vi khun, trong t bo cht cng khng cha h thng si nng
gip duy tr hnh dng t bo mt cch n nh. Lc ny hnh dng c c l do thnh t bo.
Nm ri rc trong t bo cht l cc ht ribosom, bo quan c vai tr tng hp protein. S lng ribosom
trong t bo cht tng i ln, chim ti 70% trng lng kh ca t bo vi khun. Ribosom ca t bo
Prokaryota c hnh thnh t hai tiu n v 50S v 30S. S l n v Svedberg, i lng o tc lng ca
cc ht trong mt huyn dch khi ly tm cao tc. Ribosom ca vi khun chu tc ng ca nhiu khng sinh
nh streptomycin, neomycin, tetracyclin,...
2.4. Th nhn
T bo Prokaryota cha c nhn hon chnh m ch tn ti th nhn (nuclear body). y l mt dng
nhn nguyn thu, cha c mng bao bc nn th nhn v t bo cht khng c tch bit r rng. V thnh
phn, th nhn cha nhim sc th (NST) c cu to t mt si ADN xon kp, dng trn khng lin kt
vi protein. ADN ca t bo vi khun c chiu di nm trong khong 0,25 - 3,0m, tng ng vi khong
6,6 - 13,0.106 cp nucleotid. Do ch cha 1 si NST duy nht nn i a s vi khun l t bo dng n
bi. Vai tr ca th nhn l ni cha ng thng tin di truyn v l trung tm iu khin mi hot ng sng
ca t bo.
nhiu loi vi khun, ngoi th nhn, ADN cn nm trong t bo cht di dng mt vng ADN nh
c gi l plasmid. Plasmid c kh nng sao chp mt cch c lp i vi ADN trong th nhn. Cc gen
nm trn plasmid thng m ho cho cc protein khng ng vai tr quan trng trong qu trnh sinh trng
ca t bo. Trong mt s trng hp, gen nm trn plasmid to nn c tnh khng khng sinh hoc quyt
nh gii tnh ca vi khun.
2.5. Bao nhy
Nm bn ngoi ca thnh t bo mt s loi vi khun cn c thm mt lp bao nhy (capsule). y l
mt lp vt cht dng keo, c dy khng c nh. Thnh phn ch yu ca bao nhy l cc polysaccarid.
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Vai tr ca bao nhy l gip bo v t bo khi cc tc ng bn ngoi (nh s kh hn hoc s tn

cng ca bch cu). Do c cu to t polysaccarid nn bao nhy cn ging nh mt ngun d tr dinh dng
cho t bo, phng khi ngun dinh dng trong mi trng b cn kit. Cc vi khun c bao nhy thng
c kh nng bm dnh tt trn cc gi th hoc kt dnh vi nhau thnh mng. V d, cc vi khun gy su
rng nh Streptococcus salivarius v Streptococcus mutans, bao nhy gip cc loi vi khun ny bm dnh
vo b mt rng. Trong qu trnh sng, vi khun s ln men ng (c trong thc phm cn st li trong k
rng) sinh ra acid lactic lm hng dn men rng. Nhiu loi vi khun sng thu sinh cng c bao nhy.
Bao nhy s gip chng bm c trn cc gi th v khng b nc ra tri. y l nhng vi khun c vai
tr quan trng trong vic lm sch nc.
2.6. Lng (roi) v khun mao

Khi quan st nhiu loi vi khun, ngi ta thy chng c th chuyn ng c trong mi trng lng.
S chuyn ng c khi l hon ton ngu nhin nhng cng khng t trng hp l do chng tm n
hoc trnh xa mt yu t no nh: tm n ngun thc n, ti ch c nh sng, m p hoc trnh xa nhng
ni c ho cht c hi. S chuyn ng c thc hin nh h thng roi trn b mt vi khun.
Lng (cilia, roi-flagella) l nhng si lng di, un khc, mc mt ngoi ca t bo. Lng c bit
c gi l roi. Thnh phn cu to ca lng (roi) l cc protein flagellin. Roi khng nm ngu nhin m
phn b c quy lut trn b mt t bo. c im phn b c tnh c th tu theo loi vi khun. Roi c th
nm mt u, c hai u, nm gia hoc khp xung quanh t bo. Roi hot ng theo cch quay nh
kiu vn nt chai. Nh c s vn ng ca roi m vi khun c th chuyn ng trong dch lng vi tc
khong 20 - 80m/s.
Ngoi lng (roi), trn b mt ca vi khun cn c h thng khun mao (pilus hay fimbria) bao ph. Khc
vi roi, khun mao c kch thc nh v c s lng ln hn rt nhiu. Khun mao khng c vai tr vn
ng m gip cho t bo bm dnh vo gi th. y l c im khin nhiu loi vi khun gy bnh c th
sng bm trn vt ch (v d ng tiu ho, ng h hp, ng tit niu,...).
3. CU TRC T BO EUKARYOTA
Theo h thng phn loi ca R. H. Whittaker, cc sinh vt trn Tri t c phn thnh 5 gii l
Monera, Protista, Nm (Fungi), ng vt (Animalia) v Thc vt (Plantae). Trong , cc sinh vt thuc gii
Monera c cu to c th t t bo Prokaryota, bn gii cn li c cu to t t bo Eukaryota. T bo
Eukaryota c kch thc thay i tu loi nhng ni chung ln hn t bo Prokaryota (10 - 100m). Cu trc
t bo Eukaryota kh phc tp. T bo cht khng cn l mt khi ng nht nh t bo Prokaryota m
thay vo cha rt nhiu cc bo quan vi nhng chc nng chuyn ho khc nhau (hnh 1.2).
Tnh t ngoi vo trong, t bo Eukaryota cng gm 3 thnh phn chnh l mng sinh cht bao bc bn
ngoi, bn trong l mt khi t bo cht, gia t bo l nhn. d dng cho vic nghin cu, c th tm
chia cc bo quan trong t bo ra lm 3 nhm ln c lin h vi nhau v mt chc nng nh sau:
Cc bo quan thuc h mng trong: li ni cht (c ht, khng ht), phc h Golgi, lysosom,
peroxysom.
Cc bo quan tham gia sn sinh nng lng: ty th, lc lp.
Cc bo quan tham gia biu hin gen: nhn, ribosom.
Ngoi nhng bo quan k trn, trong t bo cn mt s cc cu trc khc nh b khung xng, trung t,
cc ht d tr v.v... Mi loi t bo u c nhng cu trc c th ring, ph hp vi chc nng m n m
nhim. Ni dung chng ny ch cp n mt s bo quan chnh.
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Hnh 1.2. Cu trc t bo Eukaryota
3.1. Mng sinh cht

Mng sinh cht l mt dng ca mng sinh hc, c coi l tin thn ca tt c cc h thng mng trong
t bo. V cu to, mng sinh hc l mt mng lipoprotein dy ch khong 7-10nm. Ngi ta gi thit rng,
t bo s khai c hnh thnh di dng mt ti mng bao bc cc phn t hu c (tng t t bo
Prokaryota). V sau, lp mng ny c phn ho vo trong, to nn mt h thng mng ni bo phn chia
t bo cht thnh nhng khu vc vi chc nng ring bit (t bo Eukaryota). Nhng khu vc l cc bo
quan. Nh c s phn ho ny, cc hot ng ca t bo c thc hin mt cch c trt t v hiu qu hn
rt nhiu. Mng sinh hc khng ch n thun c vai tr ngn cch m cn l ni din ra rt nhiu hot ng
trao i cng nh chuyn ho ca t bo.
Mng sinh cht ca t bo hu ht cc sinh vt u c nhng chc nng chung, l:
Ngn cch t bo vi mi trng, gip t bo tr thnh mt h thng bit lp.
Thc hin qu trnh trao i cht, thng tin gia t bo v mi trng.
L gi th gn cc enzym ca qu trnh trao i cht trong t bo.
Vi c im bao bc ton b t bo, mng sinh cht to cho t bo tr thnh mt h thng bit lp. Qua
mng sinh cht, t bo thc hin s trao i cht vi mi trng mt cch c chn lc, ph hp vi nhu cu
sinh trng v pht trin ca t bo cng nh c th.
3.1.1. Thnh phn v cu trc ca mng sinh cht
Mng sinh cht l mt lp mng bao ph t bo cht c c t bo Prokaryota v Eukaryota. Thm ch
mt s loi virus cng c bao bc bi mng sinh cht (nh Togavirus, Rabdovirus, Micovirus,...). Mng
sinh cht cc dng t bo thng khc nhau v t l cng nh s phn b ca mt s cu trc, th nhng
chng u c bn cht l lp lipoprotein.
Cu trc ca mng sinh cht c nghin cu t rt sm, c th k ra mt s cng trnh ni bt nh
sau. Nm 1899, sau nhng thc nghim cho thy tc thm vo mng ca cc cht t l thun vi tc
ho tan trong lipid, Overton i n gi thuyt mng t bo l mt lp lipid. n nm 1925, Gortner v
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Grendel tin hnh xc nh din tch lipid ca mng hng cu sau khi tri trn mt nc v thy rng
din tch rng gp i din tch tng s mng hng cu c tch. T , hai tc gi xut mng t bo
l mt h thng 2 lp mng lipid. Vo nm 1952, Davson v Danielli a ra mt trong nhng m hnh
hon thin u tin v cu to ca mng sinh cht. Theo , nn tng ca mng sinh cht l lp lipid kp,
nm mt trong v mt ngoi mng cn c thm hai lp protein lin tc. M hnh ny ph hp vi nhng
phn tch v thnh phn ho hc cng nh nhng quan st trn knh hin vi thi by gi. Tuy nhin, cng
nghin cu su, ngi ta cng thy m hnh ca Davson v Danielli cn nhiu khim khuyt. Protein khng
nm c nh mt cch cng nhc hai mt ca mng m c s phn b linh hot v c kh nng vn ng
c. chng minh, ngi ta tin hnh lai hai loi t bo ca ngi v ca chut vi nhau. Kt qu l
ch sau mt vi pht, protein c hiu ca hai loi t bo ny hon ton trn u vi nhau (hnh 1.3).
n nm 1972, Singer v Nicolson xut mt m hnh mng sinh cht mi, gi l m hnh khm
lng. y l m hnh c tha nhn rng ri hin nay. Theo Singer v Nicolson, mng t bo cng c nn
tng l lp phospholipid kp. Trn mng cng c cc protein nhng protein khng ch nm hai b mt m
c nh khu v phn tn linh hot theo kiu khm vo lp kp phospholipid. V thnh phn ho hc,
mng sinh cht cha 3 nhm hp cht l: lipid, protein v hydratcarbon.
Hnh 1.3. S th nghim lai t bo chut v t bo ngi
3.1.1.1. Lipid
Lipid chim khong 50% (dao ng t 25-75%) trng lng mng. Cc lipid ch yu ca mng gm:
Phospholipid (chim khong 55-57%).
Cholesterol.
Glycolipid.
Nn tng c bn ca mng sinh cht l mt lp kp cc phn t phospholipid. Khung ca phospholipid
l mt phn t glycerol c 3 carbon. Gn vi C1 v C2 l cc acid bo c trng lng phn t ln (acid
palmitic v oleic), cc acid ny c th dng no hoc khng no. Hai acid bo ny cng quay v mt pha so
vi khung glycerol. Do tnh khng phn cc nn hai acid bo ny s to nn u k nc ca phn t
phospholipid. Gn vo v tr C3 l phn t ru c phosphoryl ho v c tnh phn cc (ru
phosphocholin). Phn t ny quay v pha i din vi hai acid bo v to nn u a nc ca phn t
phospholipid. Vi kiu cu to ny phospholipid l mt phn t lng cc: mt u phn cc (a nc) v
mt u khng phn cc (k nc) (hnh 1.4).
Khi a mt nhm cc phn t phospholipid vo nc, cc phn t nc s ht u phn cc ca phn
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t phospholipid v pha mnh v y u khng phn cc ra xa. Bi hin tng ny nn cc phn t

phospholipid s ngu nhin c sp xp theo kiu quay u khng phn cc vo nhau, cn u phn cc
hng ra ngoi. Cu trc ny c th c dng hnh cu (cn gi l mixen) hoc dng lp kp. Trong , dng
lp kp chnh l nn tng ca mng sinh cht. S sp xp cc phn t phospholipid to nn lp mng ny
c tnh t pht v khng tiu tn nng lng.
Thnh phn lipid th hai c mt trong mng sinh cht l cc cholesterol. Phn t cholesterol c cu to
gm mt nhm phn cc lin kt vi nhn steroid. Cc phn t cholesterol xp xen k vo lp mng kp
phospholipid theo phng thc nhn steroid th nm cnh cc ui acid bo, cn u phn cc th xp gn
vi ru phosphocholin, ngha l cc thnh phn c cng tnh cht phn cc th nm gn nhau. Khi nm xen
k vi cc phn t phospholipid, cholesterol s c vai tr c nh c hc cho mng bi chng lm cho cc
ui acid bo tr nn bt ng. Cc phn t phospholipid v th kh c th chuyn dch c.
Dng lipid th ba l glycolipid. l cc lipid lin kt vi oligosaccarid.
Hnh 1.4. Cng thc cu to phn t phospholipid v s hnh thnh lp mng kp trong mi trng nc
3.1.1.2. Protein
Protein trong mng sinh cht chim khong 25-75% trng lng mng. Thnh phn cng nh chc nng
ca protein mng rt a dng. N c th l protein cu trc, enzym xc tc, protein vn chuyn cht qua
mng, th th mng, thu nhn thng tin, v.v.. C th v protein nh nhng cng c c gn trn lp mng
kp phospholipid.
Tu theo cch sp xp m ngi ta phn ra hai loi protein:
Protein xuyn mng.
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Protein ra mng.
a) Protein xuyn mng
Nhng protein ny nm xuyn qua chiu dy ca mng v lin kt rt cht ch vi lp kp lipid qua
chui acid bo. C loi protein xuyn mng 1 ln, v d nh glycophorin mng t bo hng cu. C loi
protein xuyn mng nhiu ln, v d nh protein band3 cng t bo hng cu ging glycophorin nhng
xuyn mng ti 12 ln hoc bacteriorodopsin l mt loi protein xuyn mng ti 7 ln c vi khun. Vi
nhng protein xuyn mng, phn nm trong mng thng c tnh khng phn cc v lin kt vi ui k
nc ca lp kp phospholipid. Cn u th ra ngoi ra mng t bo (c th ra ngoi hoc ra trong) ca
cc protein ny c tnh phn cc v thng l nhm amin hoc carboxyl. Bn cnh , nhng phn protein
nm ra ngoi mng hay lin kt vi hydratcarbon to nn cc glycoprotein.
Nhiu nghin cu cho thy cc protein xuyn mng mt ln phn nhiu c vai tr l cc th th
(receptor). Cn cc protein xuyn mng nhiu ln thng l cc knh dn truyn phn t.
Mt s v d v protein xuyn mng:
Glycophorin l mt loi protein xuyn mng mt ln hng cu, c cu to gm 131 acid amin. Nm
pha ra ngoi mng l chui peptid cha nhm amin u, chui ny lin kt vi mt loi hydratcarbon
cha ti 16 mch oligosaccarid. Cc oligosaccarid ny to phn ln cc hydratcarbon ca b mt mng t
bo. Nm pha ra trong l chui peptid cha nhm carboxyl u. Phn ny thng lin kt vi cc
protein nm bn trong t bo cht.
Protein band3 c nghin cu u tin hng cu. y l mt protein xuyn mng nhiu ln, c cu
to t mt chui peptid rt di, gm ti 930 acid amin. Ging nh glycophorin, phn th ra ra ngoi mng
ca band3 cng lin kt vi mt s oligosaccarid. Vai tr ca protein band3 l mt knh dn truyn anion.
Hot ng ca protein band3 c lin quan mt thit n chc nng chuyn ch O2 v CO2 ca hng cu.
b) Protein ra mng
Protein ra mng chim khong 30% thnh phn protein mng. Chng c th nm ra ngoi hoc ra
trong ca mng. Nhng protein ny lin kt vi lp kp phospholipid thng qua lin kt ho tr. Ngoi ra,
chng cng c th lin kt vi cc protein xuyn mng thng qua lc hp ph.
Nhiu protein ra mng tham gia vo chc nng lin kt ca t bo. V d protein fibronectin l mt loi
protein ra ngoi mng, c hu ht cc loi ng vt. Fibronectin c dng mng li p st b mt ngoi
ca mng t bo. Nh c protein ny m t bo ng vt c th bm dnh vi nhau hoc bm vo gi th.
Fibronectin c gi trn b mt t bo nh cc protein xuyn mng. T bo ung th c kh nng sn sinh ra
fibronectin nhng li khng gi c n trn b mt mng, lm t bo mt kh nng bm dnh vi cc t bo
khc. cng l mt trong nhng nguyn nhn khin cho t bo ung th c th di c i nhiu ni trong c
th.
Mt v d khc v protein ra mng l cc protein actin, spectrin, ankyrin v band4.1 c hng cu. Bn
loi protein trn an vi nhau to nn mt cu trc li nm ra trong ca mng hng cu. Mt li l hnh
6 cnh. Trong , cnh li l protein spectrin dng si xon. Cn cc gc c cu to t protein actin,
ankyrin v band4.1. Li c gn vi mng thng qua lin kt gia ankyrin v protein band3. Nh c cu
trc li ny neo gi m t bo hng cu c dng a lm hai mt, gip tng cng kh nng trao i kh vi
mi trng xung quanh.
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3.1.1.3. Hydratcarbon
Hnh 1.5. M hnh mng sinh cht theo Singer v Nicolson
Khong 2 - 10% hydratcarbon trong mng sinh cht l nhng mch olygosaccarid hoc polysaccarid.
Chng thng lin kt vi lipid hoc protein to nn cc glycolipid v glycoprotein. Ngi ta thy rng
khong mt phn mi s phn t lipid mng c gn vi hydratcarbon u a nc. c im phn b ca
hydratcarbon l lun c nh khu b mt ngoi ca mng. Phn hydratcarbon ny to nn mt lp cu
trc si vi chc nng kh a dng. V d, nh to in tch m cho mng (hu ht mt ngoi t bo ng vt
u tch in m), to khng nguyn b mt, tham gia lin kt t bo.
3.1.2. Tnh linh hot ca mng sinh cht
Mng sinh cht khng phi l mt cu trc cng nhc m l mt h thng c tnh linh hot cao. S linh
hot c quy nh bi cc thnh phn cu to nn mng.
3.1.2.1. Tnh linh hot ca lp kp phospholipid
Nh trnh by trn, phn t phospholipid cha hai acid bo c mch carbon rt di to nn u k
nc nm bn trong mng. Hai acid bo ny c th dng no hoc khng no. Khi trng thi no, hai mch
acid bo c dng dui thng. Cc phn t phospholipid nm st nhau nn mng trng thi nht. Khi trng
thi cha no, mch carbon ca acid bo c dng gp khc. iu ny khin cho cc phn t phospholipid y
nhau ra xa. Cu trc mng tr nn lng lo v c dng lng. S lng phn t phospholipid trng thi no
hoc khng no v th c nh hng n tnh linh hot ca mng.
S lng phn t cholesterol cng lm thay i tnh n nh ca mng. Cc phn t ny c vai tr c
nh cc phn t phospholipid v mt c hc nn mng cha nhiu cholesterol th tnh n nh cng cao.
Cc phn t phospholipid trong lp kp khng ng yn ti ch m c s dch chuyn v tr. S chuyn
ng ny ph thuc vo tnh trng ca mng sinh cht ang dng lng hay nht. S chuyn ng s din ra
mnh hn khi mng sinh cht dng lng. Khi chuyn ng dch ch, cc phn t lipid s chuyn ch theo
chiu ngang (sang bn cnh), y l loi chuyn ng nhanh. Khng ch chuyn ng ngang, phn t
phospholipid cn c kh nng chuyn ch t lp lipid ny sang lp lipid khc (cn gi l chuyn dch bp
bnh hay chuyn dch Flip-Flop), y l kiu chuyn dch chm.
3.1.2.2. Tnh linh hot ca cc protein mng
Nhiu loi protein c kh nng chuyn ng quay v dch ch trong mng. Bnh thng cc phn t
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protein phn b tng i ng u, nhng khi c mt tc ng no y ca mi trng (nhit , pH,

khng th,... ) chng c th di chuyn v tp hp thnh m. Nhiu thc nghim cho thy, nu khng c
neo gi, phn t protein c th chuyn dch trn mng vi tc chng 10nm/pht.
Mt trong nhng v d v s chuyn dch ca protein mng do s tc ng bn ngoi l cc glycoprotein
t bo lympho. Cc glycoprotein ny bnh thng phn b ng u trn mng sinh cht. Khi x l t bo
nhit lnh hoc khi t bo ngng trao i cht, chng s tp hp li thnh nhm, to nn m khng
nguyn t bo lympho.
3.2. T bo cht
T bo cht ca t bo Eukaryota cng l mt khi dch lng song khng ng nht m c phn chia
thnh nhiu khu vc khc nhau, gi l cc bo quan. Mi bo quan c mt cu trc v chc nng chuyn
bit.
3.2.1. Li ni cht
Mng li ni cht (endoplasmic reticulum) c pht hin vo nhng nm 50 ca th k XX nh
nhng tin b ca k thut knh hin vi in t. y l mt h thng phc tp bao gm cc knh, cc ti lin
thng nhau v phn b khp trong t bo cht. Chnh v phn b rng nh vy nn li ni cht l thnh
phn ch yu ca h mng trong. Hu ht cc t bo Eukaryota u c li ni cht (tr hng cu trng
thnh). Cu trc cng nh mc pht trin ca li ni cht thay i tu loi t bo, trong pht trin
nht l nhng t bo c chc nng bi tit protein. Ngay c i vi mt t bo, nhng giai on khc
nhau, mc pht trin ca li ni cht cng khc nhau. Nhng t bo ang trong trng thi phn chia
thng c h thng li ni cht pht trin km.
V cu to, li ni cht cng c cu to t lp phospholipid ging nh mng sinh cht tuy c mc
linh hot cao hn. Li ni cht c chia ra lm hai loi l li ni cht c ht v li ni cht trn.
3.2.1.1. Li ni cht c ht
Loi li ni cht ny pht trin nhng loi t bo c mc tng hp protein mnh. V d, t bo
bch cu, t bo tuyn tu... Li ni cht c ht c cu to gm cc knh, cc xoang dt thng nhau. Trn
b mt c gn rt nhiu ht ribosom vi chc nng tng hp protein. Li ni cht c ht thng nm st v
ni lin vi mng nhn, t lan rng vo t bo cht. mt s loi t bo, li ni cht cn thng vi c
mng sinh cht (hnh 1.6).
Khi li ni cht hot ng, cc protein sau khi c ribosom tng hp s tp trung trong lng ti. Tip
, cc protein ny s i theo cc knh ca li ni cht v chuyn n phc h Golgi tip tc c hon
thin thnh sn phm hon chnh.
Hnh 1.6. M hnh li ni cht c ht (theo I. Livingstone

Biodidac)
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3.2.1.2. Li ni cht trn

Li ni cht trn c cu to t cc ng ln, nh phn nhnh. B mt ca li khng c ribosom m
thay vo l mt h thng cc enzym phc v cho qu trnh chuyn ho lipid v hydratcarbon. Li ni
cht trn khng nm tch bit m ni lin vi li ni cht c ht. iu ny khin cho hai loi li ni cht
i khi rt kh phn bit vi nhau.
Chc nng ca li ni cht trn kh a dng. Vi h thng enzym trn b mt, li ni cht thc hin
qu trnh tng hp v chuyn ha cc lipid, hydratcarbon. t bo gan, li ni cht tham gia vo vic tng
hp cc enzym gip kh mt s c t xm nhp vo t bo. Cc loi thuc nh amphetamin, morphin,
codein, phenobarbitan,... u l nhng thnh phn b chuyn ho bi t bo gan. T bo no cng l mt
trong nhng loi t bo c li ni cht trn pht trin mnh vi chc nng tng hp mt s hormone gii
tnh. Ngoi ra, li ni cht trn cn pht trin mnh cc t bo m m, t bo tuyn nhn da, v tuyn
thng thn,...
c im chung ca li ni cht c ht v li ni cht trn l cc sn phm sau khi to ra c vn
chuyn trong lng li n cc vng khc nhau ca t bo. Vi c im ny, h thng li ni cht c vai
tr nh l mt h thng giao thng ni bo.
3.2.2. Ribosom
Mc d ribosom l mt bo quan c kch thc nh v cu to khng phc tp song n c vai tr v
cng quan trng i vi t bo. Chng l ni thc hin qu trnh sinh tng hp protein. Phn ln ribosom
c nh khu trn b mt ngoi ca li ni cht c ht. Ti y, chng c sp xp theo tng chui di,
theo hnh hoa th hoc hnh xon. Cc ribosom ny c gn vi li ni cht thng qua protein riboforin.
Ribosom cn nm t do trong t bo cht. nhng t bo c mng li ni cht c ht pht trin km th
y l trng thi ch yu ca ribosom. Ngoi ra, ribosom cn c tm thy trong c ty th v lc lp l cc
bo quan c kch thc ln trong t bo.
Hnh 1.7. S lp rp hai tiu n v to nn ribosom
V mt cu to, ribosom c kch thc khong 20 - 35nm. c cu to t hai tiu n v ln (60S) v

nh (40S). C hai tiu n v u c thnh phn gm ARN ribosom v cc protein (hnh 1.7). Mc d c c
hai loi t bo nhng ribosom ca t bo Eukaryota khc vi t bo Prokaryota v kch thc. Hng s lng
ribosom ca t bo Eukaryota l 80S, cn ca t bo Prokaryota l 70S.
3.2.3. Phc h Golgi
Phc h Golgi (Golgi complex) l mt h thng tng i phc tp, thng nm gn vi li ni cht.
ng vt, s lng phc h Golgi c th thay i t 3 - 20 phc h/t bo. t bo thc vt, phc h
Golgi cn c mt tn gi khc l dictiosom vi s lng khong mt vi trm/ t bo.
Phc h Golgi c cu to gm cc ti dt ln nh, xp chng ln nhau. Hai mp ti un cong. Chiu dy
mi ti khong 20 - 40nm. Mng ti cng ging nh mng li ni cht, c thnh phn l lp lipoprotein.
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Da vo kch thc, ngi ta chia cc ti to nn phc h Golgi thnh 3 loi:

H thng cc b cha: thng xp thnh b hoc chng 5-8 b k st nhau.
Khng bo ln: thng nm cnh hoc xen k vo cc b b cha.
Khng bo b: c kch thc nh nht, thng nm mp b cha.
Cc loi ti ny c mi lin h v ngun gc lin quan n nhau. Khi cc b cha phnh ln, chng c
th tr thnh cc khng bo ln. Cc khng bo ny khi xp xung li tr thnh b cha. Cn cc khng bo
b c th c hnh thnh do s tch ra mp b cha.
Phc h Golgi c ba min cis, trans v min trung gian. Min cis hng vo li ni cht, min trans
hng ra ngoi mng t bo, cn min trung gian nm gia (hnh 1.8).
Mc pht trin ca cc thnh t cu to nn phc h Golgi khc nhau tu loi t bo. V d, trong t
bo thn, t bo nron, t bo gan th h thng b cha pht trin. Trong khi tinh t, tinh trng v non
bo th h thng b cha pht trin rt yu hoc thiu hn, phc h Golgi ch gm cc khng bo ln nh.
Trong cng mt c th nhng cc giai on pht trin khc nhau, cu to ca phc h Golgi cng khc
nhau. chut cng, trong giai on u pht trin ca phi, phc h Golgi ca t bo tuyn tu ch l h
thng b cha pht trin yu, khng c khng bo ln nh. V sau, t h thng b cha ny mi tch ra cc
khng bo b vi s lng ngy cng nhiu. nhng giai on pht trin mun hn ca phi, b cha bt
u phnh to ln tr thnh cc khng bo ln.
Hnh 1.8. Phc h Golgi (theo David H. Cormack)
Vai tr ca th Golgi l thu gp, ch bin, ng gi v phn pht cc sn phm ca li ni cht. Sn

phm ca li ni cht, bao gm cc hydratcarbon, cc lipid, cc protein thng ch dng thnh t c bn.
tr thnh cc sn phm c chc nng hon chnh, chng cn c ch bin phc h Golgi. V d, qu
trnh hnh thnh cc enzym thu phn cha trong lysosom: cc protein sau khi c chuyn n phc h
Golgi t li ni cht, ti y chng s c gn cc phn t ng to nn glycoprotein. Tip , cc
glycoprotein ny li c phosphoryl ho tr thnh enzym c chc nng hon chnh, c bao gi trong
cc ti nh gi l lysosom. Cc sn phm ca phc h Golgi gm nhiu chng loi vi cc vai tr khc nhau.
V d, cc enzym thu phn, cc ht non hong, cc hormone steroid, cc hormone insulin v.v... Chng c
th c s dng ngay trong t bo hoc bi xut ra ngoi mi trng.
Qu trnh ch bin ti phc h Golgi bt u t mt cis v hon thnh mt trans. Cc sn phm ca
li ni cht nh protein, lipid, hydratcarbon c vn chuyn n phc h Golgi di dng cc ti nh. Cc
ti ny nhp vo h thng b cha ca phc h Golgi mt cis (hnh 1.9). Tip , cc sn phm s c
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chuyn dn ln mt trans. Trong qu trnh vn chuyn , sn phm s c lp rp vi nhau, c bin

i khi nm mt trans th chng tr thnh sn phm hon chnh. Theo mt s nghin cu, qu trnh
vn chuyn bn trong phc h Golgi c thc hin thng qua phng thc ny chi. Sau khi kt thc mi
giai on ch bin, sn phm s tch ra mp b cha di dng cc ti nh v nhp vo b cha pha trn.
Ti y, qu trnh hon thin sn phm li tip tc din ra. Khi nm mt trans, cc sn phm khng bi
xut ra ngoi t bo cht mt cch t do m c bao gi trong cc ti mng tch ra t mt trans ca phc h
Golgi.
C th thy rng phc h Golgi cng vi li ni cht l hai thnh phn to nn dy chuyn sn xut ca
t bo.
Hnh 1.9. S vn chuyn sn phm t li ni cht c ht n th Golgi

1. Ny chi t li ni cht ht; 2. Vn chuyn di dng bng ti; 3. Nhp vo phc h Golgi.
3.2.4. Lysosom
Lysosom l mt trong rt nhiu sn phm ca phc h Golgi vi vai tr l tham gia vo qu trnh tiu
ho ni bo. Cc khi cht hu c sau khi a vo t bo s c phn gii nh lysosom.
Lc mi hnh thnh, lysosom ch l cc ti c kch thc nh (0,3 - 0,5m) c dng hnh cu hoc hnh
trng c gii hn bi mng lipoprotein, bn trong cha cc enzym thu phn. Loi lysosom ny c gi
l lysosom cp 1 v trng thi cha hot ng. Chng c to ra do s ny chi mp cc b cha ca
phc h Golgi. S lng lysosom cp 1 c sn sinh tu thuc loi t bo. Cc t bo bch cu mono, cc
bch cu trung tnh l nhng loi t bo c s lng lysosom trong t bo cht tng i ln.
Lysosom cp 1 chuyn sang trng thi hot ng khi chng c dung hp vi cc ti nhp bo. Trong
qu trnh nhp bo, cc khi cht hu c c a vo t bo di dng cc ti mng. Cc ti ny c a
su vo trong t bo cht v dung hp vi cc lysosom cp 1. Qua , enzym thu phn t lysosom cp 1 s
c gii phng vo ti v tin hnh phn ct cc phn t hu c. Dng dung hp gia ti thc bo v
lysosom cp 1 c gi l lysosom cp 2, dng lysosom ang hot ng. Cc ng n, acid amin,
nucleotid,... c to ra sau khi enzym phn ct khi cht hu c s c gii phng vo trong t bo cht.
Cc thnh phn khng s dng c nm li trong ti v c thi ra ngoi theo phng thc xut bo (hnh
1.10).
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Hnh 1.10. S to thnh v hot ng ca lysosom
Cc enzym thu phn cha trong lysosom c th quy v bn nhm chnh l protease, lipase, glucosidase,
nuclease. Ngoi ra, cn c mt s nhm khc nh phosphatase, phospholipase, sunfatase v.v... Cc enzym
ny c c im chung l hot ng trong iu kin pH = 5, trong khi pH ca t bo cht bng 7. Chnh
iu kin pH c trng nh vy gii hn cc enzym trn ch hot ng c trong lysosom m khng th
hot ng trong t bo cht.
Khng ch tham gia vo qu trnh thc bo, lysosom cn thc hin tiu hu cc mnh v t bo, cc bo
quan b gi ci, h hng, gip t bo ti s dng li cc n v cu trc. sinh vt a bo, lysosom ca
mt s t bo nht nh khi n mt thi im no s ng lot b v v gii phng cc enzym thu phn
vo t bo cht, pH ca t bo cht b gim xung. Lc ny, ton b cc cu trc ca t bo s b enzym phn
gii. l qu trnh t tiu. Mt v d in hnh ca hin tng ny l qu trnh bin i nng nc thnh ch.
Khi pht trin n mt giai on nht nh, ui ca nng nc tr nn khng cn thit na v s b tiu bin
nng nc tr thnh ch. C th thy rng qu trnh t tiu ng gp vo qu trnh bin thi c th ca
mt s loi t dng n gin tr thnh dng c t chc phc tp hn.
ngi, nhng ri lon sai hng v cu trc ca lysosom l nguyn nhn gy nn nhiu cn bnh m
i khi l kh trm trng. V d, bnh Tay-Sachs pht sinh do lysosom thiu mt enzym c tn l -Nhexominase A. Nhng tr b bnh ny c biu hin ri lon h thn kinh trung ng, chm pht trin tr tu
v cht tui th 5. Mt cn bnh khc hay gp cc th m l bnh vim phi. Cc loi bi nh bi silic,
amian, cc kim loi nng khi xm nhp nhiu vo phi s lm hng cc lysosom ca t bo ph nang. Enzym
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ca lysosom b gii phng tc ng ln cc ph nang v gy bnh vim phi.

3.2.5. Peroxysom
y l mt loi bo quan gn vi lysosom. Chng c cu to t cc ti mng lipoprotein c kch thc
t 0,15 1,7m. Bn trong mng l cht nn (matrix) ng nht, hoc c cha cc ht nh hay si phn
nhnh. Trong peroxysom cha cc enzym oxy ho nh catalase, D-aminoacid oxydase, urat oxydase. Ring
enzym catalase c mt trong mi loi peroxysom v chim ti 15% thnh phn protein ca bo quan ny.
Chc nng c nhc n nhiu nht ca peroxysom l tham gia phn gii hydro peroxyd (H2O2) thnh
nc nh s c mt ca enzym catalase. Bn cnh , peroxysom cn tham gia oxy ho acid uric, tham gia
chuyn ho cc acid amin. ng vt c xng sng, peroxysom c nhiu trong t bo gan, thn.
thc vt c mt loi peroxysom c bit c tn l glioxysom. Bo quan ny tham gia vo qu trnh
chuyn ho cc lipid thnh glucid, mt qu trnh quan trng c trng cho thc vt.
3.2.6. Ty th
Ty th l bo quan c mt trong tt c cc t bo h hp hiu kh, thc hin chc nng v cng quan
trng l ni chuyn ho nng lng t cc phn t dinh dng thnh dng nng lng tch tr trong phn t
ATP, dng nng lng cn thit cho tt c cc hot ng sng ca t bo.
Ty th thng c dng ht hoc dng si vi ng knh t 0,5 1m, chiu di t 1 - 7m. Hnh dng
v kch thc ca ty th thay i tu loi v tu theo trng thi hot ng ca t bo. nhng t bo ang
hot ng mnh, ty th thng c kch thc ln hn bnh thng. Cc nhn t nh p sut thm thu, pH,
tnh trng bnh l ca t bo u gy nn nhng bin i v hnh dng ca ty th.
Thng thng, ty th phn b ng u khp t bo cht. Tuy nhin, c nhng lc chng tp trung nhiu
phn ny hoc phn khc ca t bo, ni cn nhiu nng lng hot ng. V d, t bo ng thn, ty
th thng tp trung phn y, ni mng t bo cn nhiu ATP vn chuyn nc v cc cht. Khi tin
hnh quay phim, ngi ta thy ty th khng nm yn mt ch m lun chuyn ng. Ty th c th co ngn
li, dui di ra, lin kt li vi nhau thnh si di hoc t thnh nhiu khc nh. Ty th cng chuyn ng
t v tr ny n v tr khc ca t bo. S chuyn ng dch ch ca ty th c ngun gc do s vn ng ca
t bo cht to nn.
Cng ging nh hnh dng v kch thc, s lng ty th thay i t vi chc n vi nghn tu loi v
trng thi hot ng ca t bo. T bo gan l mt trong nhng t bo c rt nhiu ty th. V d, chut c
th ln n 1.000 - 2.000 ty th/ t bo.
V cu to, ty th bao gm hai lp mng lipoprotein, gia hai lp mng l khe gian mng hay cn gi l
xoang ngoi. Mng ngoi ca ty th bng phng, c thnh phn protein chim khong 60%, 40% cn li l
lipid. Thnh phn protein ca mng ngoi gm nhiu loi knh dn truyn v cc enzym nh transferase,
cytocrom B, phospholipase,... Mng trong c thnh phn v cu trc khc vi mng ngoi. B mt mng
trong khng phng m gp hoc lm vo trong to nn cc cu trc dng ng hoc ti c gi l mo
(cristae). Hm lng protein ca
mng trong rt cao, c th ln n 80%, c b tr xuyn mng hoc ra mng. Cc protein ny bao gm
cc phc h dn truyn in t, cc enzym, knh dn truyn phn t v cc ht c bn. Cc ht c bn c
dng hnh cu c cung m xuyn qua mng trong ty th. ng knh ca ht c bn khong 11nm. Vai tr
ca ht c bn l thc hin cng on cui cng to ra sn phm ATP ca ty th. Vic gp np ca mng
trong lm tng din tch b mt, qua lm tng hiu qu hot ng ca ty th.
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Hnh 1.11. Ty th
Mng trong gii hn mt khong khng gian c gi l xoang trong (cn gi l cht nn) ty th. y l
mt phn dch lng cha h enzym ca chu trnh Kreb (mt giai on ca qu trnh h hp) cha cc enzym
tng hp acid bo, cc ion v c bit l acid nucleic (ADN, ARN) v ribosom. Nh c acid nucleic v
ribosom ring nn ty th c kh nng t tng hp mt s thnh phn protein cu to cho chnh n. Tuy nhin,
s protein ny khng nhiu, ch khong 10%, phn ln protein cn li vn c m ho trn ADN trong
nhn.
ADN ca ty th c dng vng, khng lin kt vi protein histon, nm t do hoc bm vo mng trong
ca ty th. So vi ADN trong nhn, ADN ty th c mt s im khc bit v m b ba. Trong khi mt s m
b ba trn ADN nhn c vai tr l b ba kt thc th ty th li m ho cho acid amin v ngc li. ngi,
b gen ca ty th c tnh n nh cao, t b bin i qua cc th h.
3.2.7. Lc lp
Lc lp thuc nhm bo quan c tn l lp th, c Schimper m t vo nm 1885 nh l mt thnh
phn c trng ch c t bo thc vt. Ngy nay, ngi ta chia lp th ra lm hai nhm ln l bch lp (loi
lp th khng c mu) v sc lp (loi lp th c cha sc t). Trong , lc lp l mt loi sc lp c vai tr
thc hin qu trnh quang hp thc vt.
S lng lc lp trong mt t bo thc vt thay i tu theo loi v chng loi m. thc vt bc cao,
mi t bo l c khong 20 40 lc lp. Ch mt s t t bo l c mt hoc hai lc lp. Lc lp thng phn
b ng u trong t bo cht. Tuy nhin, chng khng nm c nh m c s vn ng do iu kin ngoi
cnh hoc nh hng ca dng chy t bo cht.
Lc lp thng c dng hnh cu, hnh trng hoc a. Trong mt s trng hp, lc lp c th dng
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li, di xon hoc hnh sao nh t bo to. thc vt bc cao, lc lp thng c ng knh trung
bnh 4 6m. Kch thc ca lc lp cng c s a dng tu theo loi. Ngi ta thy rng, nhng thc vt
mc trong bng rm, lc lp thng ln v cha nhiu chlorophyll hn nhng thc vt c chiu sng
thng xuyn.
Ging nh ty th, lc lp c cu to gm hai lp mng lipoprotein, nhng khc ch c hai lp mng
ny u phng ch khng gp np nh ty th. Gia hai lp mng l khong khng gian c gi l khe
gian mng. Phn dch c gii hn bi mng trong gi l cht nn (stroma). Trong cht nn cha thnh
phn quan trng nht ca lc lp. l h thng cc ct c ni vi nhau thnh mng li. Cc ct c tn
l grana cn cc di mng ni cc ct c gi l lamella. S lng grana c th thay i t vi ct n 50
ct tu loi lc lp. Chng c cu to gm rt nhiu ti nh, c gi l ti thylacoid, xp chng ln nhau.
Ti thylacoid c dng dt, ng knh 0,6m, dy 20nm v cng c cu to t mng lipoprotein. Trn mng
ti thylacoid c cha h thng sc t v chui truyn in t thc hin pha sng ca quang hp. Ngoi h
thng grana, trong cht nn lc lp cn cha cc enzym tng hp hydratcarbon, cc acid nucleic v ribosom.
ADN ca lc lp cng c dng vng ging nh ty th vi chiu di khong 150m v khng lin kt vi
protein histon. Mc d l thnh phn ca t bo Eukaryota nhng ribosom ca c ty th v lc lp u c
hng s lng l 70S ch khng phi l 80S nh trong t bo cht.
Vi c im chung l u cha acid nucleic, ribosom v c th t tng hp mt s thnh phn protein
cho ring mnh, ty th v lc lp c gi l bo quan bn t tr. So snh c im cu to ADN v ribosom
ca cc bo quan ny vi ca t bo Prokaryota, ngi ta a ra gi thuyt rng ty th v lc lp vn l
nhng t bo Prokaryota xm nhp v sng cng
sinh trong t bo Eukaryota. Dn dn, b gen ca ty th, lc lp tch ra v st nhp vo b gen ca t bo,
cc thnh phn khng cn thit b tiu bin v chng tr thnh nhng bo quan ca t bo Eukaryota
Hnh 1.12. Lc lp
3.3. Nhn t bo
Nhn l thnh phn bt buc ca tt c cc loi t bo Eukaryota. t bo Prokaryota tuy khng c
nhn nhng cng c mt vng vi chc nng tng ng c gi l th nhn. Nhn thc hin hai chc nng
quan trng l:
Lu tr v truyn thng tin di truyn t th h t bo ny sang th h t bo khc.
iu khin mi hot ng sng ca t bo.
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Phn ln t bo ch c mt nhn. Th nhng cng c nhng trng hp m mt t bo c th c 2, 3

hoc nhiu nhn. V d, trng giy Paramecium l mt sinh vt n bo c hai nhn vi mt nhn ln v
mt nhn b, khong 15% t bo gan ca ngi cng c hai nhn, cn t bo megakaryocyte trong tu xng
th c ti hng chc nhn.
Hnh dng ca nhn thng ph thuc vo hnh dng t bo. Vi nhng t bo hnh khi, a gic, hnh
cu,... nhn thng c dng hnh cu. nhng t bo hnh tr th nhn c dng ko di, v d, t bo c.
Nhn cng c th c dng c bit nh dng chia thu bch cu ht. Ngoi ra, hnh dng nhn c th thay
i tu theo tui v hot ng ca t bo. Nhng t bo hot ng mnh thng c nhn ln hoc c th chia
thu.
Kch thc ca nhn thay i tu loi t bo. Ngay c trong cng mt loi, kch thc nhn cng thay
i tu theo trng thi hot ng ca t bo. Mi mt loi t bo thng c t s gia th tch nhn v th
tch t bo cht dao ng trong mt khong xc nh. Nu ln hn khong , ng ngha vi vic nhn ln
hn bnh thng th l lc t bo chuyn sang trng thi phn chia.
Bn cnh hnh dng v kch thc, v tr ca nhn cng c tnh c th cho loi t bo v cc trng thi
khc nhau ca t bo. Trong cc t bo phi, nhn thng nm trung tm. Trong t bo trng giu non
hong, nhn li nm ngoi vi. t bo tit, nhn hay nm lch xung phn y ca t bo, cn phn ngn
thng cha nhiu ht tit. V cu trc, c th chia nhn thnh bn phn: mng nhn, dch nhn, nhim sc
th v hch nhn (hnh 1.13).
Hnh 1.13. Nhn t bo
3.3.1. Mng nhn

Ging nh mng sinh cht, mng nhn thc hin hai chc nng chnh l:
Tch bit nhn vi phn t bo cht bn ngoi.
iu chnh s trao i cht gia nhn v phn cn li ca t bo.
Mng nhn c cu to t mng lipoprotein. Nh vy l v thnh phn, mng nhn cng tng t nh
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mng ca cc bo quan khc v tng t nh mng sinh cht. Cn v tnh cht, mng nhn c nhng
c im ring. Mng nhn khng c kh nng t hn gn li nh mng sinh cht. Nu b chc thng, nhn
s b ph hu. Tnh thm thu ca mng nhn cng khc vi mng sinh cht. Nhiu thnh phn thm c
qua mng t bo nhng khng th thm qua mng nhn.
Quan st trn knh hin vi in t cho thy, mng nhn l mt cu trc gm hai lp mng: mng ngoi
v mng trong. Xoang c gii hn bi hai mng ny c gi l xoang quanh nhn. a s mng nhn c
dy chng 40nm, dy ca mng trong v mng ngoi gn 10nm, cn ca xoang quanh nhn l 10 20nm. B mt mng ngoi gn nhiu ht ribosom. Mng ngoi c thnh phn gn ging v ni lin vi li
ni cht. nhiu loi t bo, v d nh t bo ng thn, t bo bch cu, mt s t bo thc vt,... xoang
quanh nhn v h thng cc b cha ca li ni cht cng c ni vi nhau bi cc khe hp, to nn mt
h thng xoang thng nhau. Trong mt s trng hp, h thng xoang ny c th thng ra ngoi t bo. p
st b mt ca mng trong l mt h thng si dy c gi l lamina. Lamina c to nn do s an kt li
ca ba loi protein laminin A, B v C. H thng si lamina c tc dng nh gi cho mng nhn v ng
thi cng l ni nh bm ca cc si nhim sc vng ngoi vi bn trong nhn.
Mng nhn c mt c im quan trng, l tnh khng lin tc. Trn mng nhn c rt nhiu cc l
nh gip cho phn bn trong ca nhn thng vi t bo cht. Cc l phn b trn b mt mng nhn tng
i ng u vi khong cch t 50 - 100nm. L mng nhn khng phi l l n gin m c to nn t
mt phc h cc protein. Mi phc h ny gm tm n v xp vng quanh to thnh l nhn vi ng knh
l khong 9nm, nm gia l nhn l mt ht protein trung tm. Phc h protein ny s m nhn vai tr
iu tit cc cht i qua mng nhn. Ngoi ra, l nhn cn ging nh mt h thng ct nng gip n nh
cu trc ca mng nhn.
3.3.2. Dch nhn
Dch nhn l mt khi dch lng cha cht nhim sc, ion, enzym ca qu trnh nh nhn i ADN, qu
trnh phin m to ARN v.v... Trong dch nhn ngi ta cng thy c h thng cc si ging nh trong t bo
cht vi vai tr to nn b khung nng ca nhn. Bn cnh , b khung ny cng tham gia vo qu trnh
vn ng ca acid nucleic v cc protein bn trong nhn.
3.3.3. Nhim sc th
Trong t bo Prokaryota cng nh Eukaryota, thng tin di truyn u c m ho trn ADN (acid
deoxyribonucleic). t bo Eukaryota, nhim sc th c cu to do s kt hp ca phn t ADN v cc
protein (ch yu l histon). y l im khc bit ln so vi nhim sc th ca t bo Prokaryota vn ch l
si ADN trn. Mt im khc bit na l s lng nhim sc th gia hai loi t bo. T bo Prokaryota ch
c mt si nhim sc th duy nht trong khi t bo Eukaryota c rt nhiu si tng ng vi b nhim sc
th ca loi, v d ngi l 46 si, mui l 6 si, rui gim l 8 si v.v...
Qua mi giai on ca chu trnh t bo, nhim sc th li c nhng hnh thi khc nhau. k trung gian,
nhim sc th dng si mnh. Khi t bo bc sang trng thi phn chia, cc nhim sc th c nhn i
v bt u co xon li thnh dng ch X in hnh. iu ny khng c t bo Prokaryota.
T nhng thng tin m ho trn ADN trong nhn, t bo tin hnh chuyn i chng thnh trnh t acid
amin ca cc protein. Cc protein ny c th l protein cu trc, cng c th l protein vi chc nng iu
khin cc hot ng sng ca t bo.
3.3.4. Hch nhn
Khi quan st trn knh hin vi, ngi ta thy trong nhn c mt hoc mt s vng bt mu sm v gi l
hch nhn. Hch nhn khng c cu trc tch bit vi cc phn khc ca nhn. ch l mt ni c mt
cc acid nucleic v protein tp trung rt cao. ngi, hch nhn do 10 nhim sc th thuc cc cp 13, 14,
15, 21 v 22 chm u li to thnh. Trong hch nhn, ngi ta thy c s tng hp mnh m ca cc ARN.
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V thnh phn, hch nhn cha 10 20% ARN ca c t bo. Mt s loi protein sau khi c tng hp
t bo cht s c a vo nhn v chuyn ti hch nhn. Ti y s din ra qu trnh kt hp cc on
ARN vi protein to nn cc ht nh. y l cc tiu n v ca ribosom. Ni cch khc, hch nhn chnh
l ni sn xut cc tiu n v ribosom ca t bo. Cc tiu n v ny sau khi c to nn s i ra ngoi
nhn v ghp vi nhau thnh ribosom hon chnh. Phn nhim sc th nm trong hch nhn l nhng on
mang gen m ho cho ARN ribosom v c gi l vng NOR (Nuclear Organization Regions). Hch nhn
ch tn ti trong nhn k trung gian. Chng s bin mt k u v xut hin li khi kt thc k cui ca
qu trnh phn bo.
3.4. Mt s thnh phn khc trong t bo Eukaryota

3.4.1. B khung t bo
B khung t bo c cu to t mt h thng cc vi si v vi ng an xen vo nhau. Chng thc hin
chc nng nng , neo gi cc b phn trong t bo, gip chng khng b tri dt mt cch t do trong t
bo cht (hnh 1.14). Ngoi ra, h thng cc vi si, vi ng cn tham gia chc nng vn ng ca t bo.
Hnh 1.14. B khung t bo
3.4.1.1. Vi si
Trong t bo c 3 loi vi si ch yu l vi si actin, vi si myozin v vi si trung gian.
Vi si actin
L loi vi si c cu to t protein actin. Loi protein ny rt ph bin t bo Eukaryota vi hm
lng ln (chim 5% protein tng s ca t bo). Protein actin phn b hu khp t bo cht. a s t bo
ng vt, chng xp thnh b song song hoc mng li nm ngay st mng sinh cht, gn vi mng sinh
cht thng qua cc protein lin kt (spectrin, -actinin, clathrin) v c vai tr nng , c nh mng.
Protein actin c chia lm hai loi: actin G (dng cu) v actin F (dng si). Dng actin F c hnh
thnh t hai chui actin G xon quanh nhau, c ng knh 7nm v chiu di bc xon l 72nm. Qu trnh
trng hp ca cc actin G din ra khi c mt ion Mg2+ v ATP. y l mt qu trnh c tnh thun nghch,
si actin F c th c hnh thnh v phn gii nhanh chng nhng ni, nhng thi im cn thit tu theo
nhu cu ca t bo. Dng actin F chnh l cc vi si actin vi vai tr to nn b khung t bo (hnh 1.15). S
trng hp v gii trng hp ca cc protein actin gp phn lm thay i c tnh ca t bo cht. Khi cha
nhiu actin F, t bo cht c trng thi nht, cn khi cha nhiu actin G s c dng lng.
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Hnh 1.15. Vi si actin
Ngoi vai tr to nn b khung, cc vi si cn tham gia vo qu trnh vn ng t bo. Cc dng vn

ng t bo nh vn ng chn gi, vn ng dng t bo cht hay vn ng ca cc bo quan t phn ny
n phn khc ca t bo u lin quan n hot ng ca vi si actin kt hp vi vi si myozin. S vn
ng ca ton b h thng t bo c ca c th sinh vt cng c ngun gc t hot ng ca cc vi si to
nn. Ngoi ra, cc vi si cn ng vai tr to cc mi lin kt v cu ni gia cc t bo cnh nhau.
Vi si myozin
L loi vi si cu to t protein myozin. y l mt loi protein phc tp, c cu trc bt i xng vi
trng lng phn t khong 450.000 Dalton. Mt phn t myozin c hnh thnh t 6 mch polypeptid: 2
mch nng, 4 mch nh. t bo bnh thng vi si myozin c cu trc ngn, trong khi t bo c chng c
kch thc kh di v c th ln n 1,5m. Vi si myozin gp phn to nn b khung v kt hp cng vi si
actin thc hin chc nng vn ng ca t bo.
Vi si trung gian
y l loi vi si ph bin trong cc t bo Eukaryota. Chng c dy t 8 - 10nm v c hnh thnh
t nhiu loi protein khc nhau nh vimentin, desmin, cytokeratin v.v... Da vo thnh phn protein m
ngi ta chia vi si trung gian thnh nhiu nhm khc nhau. c im chung ca cc loi vi si ny l u
c cu trc phc tp, gm nhiu si nh bn xon vi nhau. Chc nng ca vi si trung gian l tham gia to
b khung t bo.
3.4.1.2. Vi ng
Vi ng l nhng cu trc hnh tr di, khng phn nhnh, c ng knh trung bnh 25nm, thnh bn
dy 5nm v rng gia. Chiu di ca vi ng thay i tu tnh trng t bo v c th ln n vi m. Vi ng
c cu to t protein tubulin. Mi phn t protein tubulin l mt dimer gm hai monomer l tubulin A v B
ni vi nhau. Cc protein tubulin xp vi nhau thnh vng xon v to nn vi ng. Nhn t trn xung, thnh
vi ng c cu to t 13 protein tubulin xp vi nhau. S trng hp to nn vi ng c tnh thun nghch v
xy ra vi s c mt ca ion Mg2+ v GTP.
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Hnh 1.16. Vi ng
Ngoi vic kt hp vi vi si to nn b khung t bo, vi ng cn tham gia vo nhiu vai tr khc

nh vn ti ni bo hay l thnh phn cu to ca trung t, lng v roi. Cc bo quan nh ty th, lc lp, cc
bng xut nhp bo c vn chuyn t phn ny n phn kia ca t bo cht l nh hot ng ca vi ng.
Trong qu trnh phn bo, thoi v sc ni vi cc nhim sc th cng c cu to t cc vi ng. Colchicin l
mt cht c tc dng ngn cn s trng hp ca cc protein tubulin. Khi c mt cht ny, thoi v sc khng
th hnh thnh dn n ri lon trong vic phn chia nhim sc th. Qu trnh phn bo c th b dng li
k gia.
3.4.2. Trung th
Trung th (centrosome) l bo quan c trong tt c cc t bo ng vt a bo cng nh n bo. Mi t
bo c mt trung th nm gn nhn. Cu to ca trung th bao gm trung t (centriole) v cht quanh trung
t (pericentriole). t bo thc vt, cu to ca trung th khng c trung t m ch c cht quanh trung t.
Trung t ca t bo ng vt thng c hai chic nm vung gc vi nhau trong trung th. Trung t c
cu to hnh tr vi ng knh 0,15 - 0,25m v chiu di 0,7m. Thnh tr c to bi 9 b 3 vi ng xp
ni nhau. Cc vi ng trong b ba c ng knh 20 - 26nm, c gi l vi ng A, B v C. Vi ng A nm
gn tm nht, ri n vi ng B v xa nht l vi ng C. Nhng quan st trn knh hin vi in t cho thy
trong ba vi ng, ch c vi ng A l hon chnh, cn vi ng B v C c dng ch U v ni vi vi ng pha
trc. Cc b ba vi ng c ni vi nhau bi protein nexin, protein ny s ni vi ng A ca b ba pha
trc vi vi ng C ca b ba nm pha sau. Vi cch t chc nh vy, ngi ta gi cu trc ca trung t l 9
+ 0.
Cht bao quanh trung t c cu to t hai phn. Phn th nht l cc th km c cu trc hnh cu
ng knh 40 - 70nm c ni vi vi ng ca trung t thng qua cc cung. Phn th hai l mt h thng
gm cc vi ng t do xp xung quanh trung t theo hnh phng x.
Trung th ng vai tr quan trng trong qu trnh phn bo. Khi t bo bc sang k gia, trung th
c nhn i v i v hai cc ca t bo. Vi s c mt ca ATP, trung th kch thch s trng hp to nn
cc vi ng. Vic hai trung th nm hai cc ca t bo s gip nh hng cho cc vi ng to thnh thoi v
sc. Nh c thoi v sc, cc nhim sc th c phn chia v vn chuyn ng v hai cc ca t bo. n k
cui ca phn bo, cc vi ng to nn thoi v sc c gii trng hp, thoi v sc bin mt. Do hot ng
ca trung th nh vy nn ngi ta cn gi chng vi mt ci tn khc l MTOC (Microtubule Organizing
Center - trung tm t chc vi ng).
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Hnh 1.17. Trung t
3.4.3. Lng v roi

Lng (cilia) v roi (flagella) l nhng phn li ra ca t bo cht c bao bc bi mng, bn trong c cha
h thng cc vi ng. Lng v roi c chc nng vn ng. Ngi ta phn bit lng v roi thng qua s lng
v kch thc ca chng. Lng c chiu di t 10 20m v c s lng rt nhiu, c khi ti 300 lng/t
bo. Roi c kch thc ln hn lng, chiu di c th ti 150m. Thng thng mi t bo ch c mt vi
roi. Lng v roi ch c mt s loi t bo nht nh. V d, cc ng vt nguyn sinh, lng v roi l
phng tin ch yu gip chng chuyn ng. c th a bo, b mt cc t bo biu m thnh ng cng c
lng nhng chng khng ng vai tr gip t bo chuyn ng m gip to nn dng chy trong lng ng.
Tinh trng cng l mt loi t bo c roi l phng tin chuyn ng.
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Hnh 1.18. Cu to ca lng v roi
V cu trc, lng v roi ging nhau. Chng c dng hnh tr, ng knh 0,2m. Cc vi ng bn trong
lng v roi c b tr theo kiu 9 + 1. gia l mt i vi ng trung tm ng knh 20nm, c mng bao
quanh. Xp xung quanh i vi ng trung tm l 9 i vi ng ngoi vi. Mi vi ng ngoi vi c ng knh t
18 - 22nm. Mi i vi ng bao gm hai vi ng k hiu l A v B xp st nhau. ng A c kch thc b v
nm gn trung tm hn. Thnh ca ng A c gn 2 mu protein dynein. Khi c mt ATP v Ca2+, cc mu
dynein s lin kt vi i ng bn cnh, gy ra s trt ca cc i vi ng ngoi vi v lm cho lng, roi
chuyn ng. Cc i vi ng ngoi vi c ni vi nhau bng cu ni nexin v ni vi i vi ng trung tm
thng qua cc di mnh nh hnh nan hoa.
4. S VN CHUYN CHT QUA MNG

4.1. Mt s khi nim
4.1.1. S khuch tn
Khi nh mt git mc vo trong cc nc, chng ta d dng thy ban u mc tp trung ti mt ch, v
sau git mc s lan dn ra ton b cc nc. y l mt th nghim n gin v s khuch tn. Khuch
tn c th hiu l s chuyn ng ca cc phn t t ni c nng cao n ni c nng thp hn.
Nguyn nhn ca s chuyn ng ny l do c tnh chuyn ng hn lon ca cc phn t. Khi mt cng
dy c, xc sut va chm gia cc phn t cng ln. iu khin chng c xu hng chuyn ng ra xa
nhau v dn u trong mi trng. Khuch tn l mt trong nhng cch thc t bo c th trao i vt
cht vi bn ngoi.
4.1.2. S thm thu
Mng sinh cht l mt mng bn thm. Khng phi phn t no cng c th t do i qua c mc d
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c s chnh lch nhiu v nng gia hai bn mng. Trong trng hp ny, cn bng nng , i
tng khuch tn qua mng s l dung mi. i vi t bo, dung mi ho tan ca phn ln cc cht l nc.
Chnh s chnh lch v nng cht tan to ng lc ko nc qua mng nhm thit lp s cn bng
nng . S vn chuyn nc nh vy gi l thm thu. Chnh lch v nng cht tan cng cao, p sut
thm thu sinh ra cng mnh.
Hng chuyn ng ca nc lun i t ni c nng cht ho tan thp n ni c nng cht ha
tan cao. Do t bo l mt h thng tch bit vi mi trng v nng cc cht ni bo lun c s n nh
mt cch tng i. Bi vy, cng ging nh s khuch tn cht tan, s thm thu nc ph thuc vo iu
kin mi trng. Da vo mi tng quan vi t bo, ngi ta chia mi trng ngoi bo ra lm ba kiu: u
trng, nhc trng v ng trng. S bin i ca t bo khi a vo cc mi trng trn cng khc
nhau (hnh 1.19).
Mi trng u trng: l mi trng c nng cht tan cao hn t bo. Nc v th s b rt ra ngoi
khin t bo b co rt li. Nu khng hi phc c, t bo c th cht.
Mi trng nhc trng: l mi trng c nng cht tan thp hn t bo. Nc s thm thu vo
trong v lm t bo trng ln. Mng t bo s phi chu mt p sut thu tnh do nc gy nn. Cng nhiu
nc, p sut thu tnh cng ln. Nu nh qu ngng chu ng, mng t bo s b v, phn t bo cht bn
trong tro ra ngoi. l hin tng tan bo.
Mi trng ng trng: l mi trng c nng cht tan cn bng vi t bo. Lng nc i vo
v i ra khi t bo l cn bng. Lc ny t bo trng thi n nh.
Hnh 1.19. S vn chuyn nc qua mng
C th thy rng tr trng hp ng trng, cn li cc mi trng nhc trng v u trng u c

nguy c gy hi cho t bo. Th nhng trong mi trng t nhin, rt t khi nng cht tan li c th cn
bng vi t bo. V th t bo hay c th phi c nhng phng thc nht nh gim thiu tc hi ca s
thm thu nc qu mc. Mt trong nhng cch thc ph bin l s dng vch t bo. T bo vi khun hay
thc vt hu nh lun c bao bc bi mt lp vch t bo dy. Chnh lp vch ny gip t bo c th chu
c p sut rt cao. Khi nc thm thu nhiu vo bn trong, t bo ch cng ln ch khng b v. Cch th
hai l iu chnh nng cht tan cn bng vi mi trng. Nhiu trng hp khi cho t bo vo mi trng
u trng, t bo b rt nc v co li. Th nhng nu c vy trong mt thi gian th t bo li phnh ln
v tr v trng thi bnh thng. S d c hin tng nh vy l v t bo vn chuyn cht tan vo trong t
bo cht v lm nng ni bo tng ln cn bng vi mi trng. Khi nc s khng b rt ra na. Mt
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cch thch nghi khc l s dng cc bo quan chuyn trch vn chuyn nc. iu ny thng gp
cc ng vt nguyn sinh. Trong mi t bo ng vt nguyn sinh thng c mt hoc vi khng bo co bp
nm st mng. Nhim v ca nhng khng bo co bp ny l ht nc t trong t bo cht vo n v y ra
ngoi mi trng.
4.2. Vn chuyn cc cht khng tiu ph nng lng

Hnh thc vn chuyn ny cn c mt tn gi khc l vn chuyn th ng. Kh nng vn chuyn th
ng cc cht ph thuc vo ba yu t:
Gradient nng : y l yu t quan trng nht v cng l ng lc ca s vn chuyn th ng. Khi
c s chnh lch v nng mt cht hai pha ca mng, cc phn t ca cht s dch chuyn theo
nguyn tc khuch tn t ni c nng cao ti ni c nng thp hn. Nh vy, c th thy rng chiu
hng ca s vn chuyn hon ton ph thuc vo hng ca gradient nng . chnh l tnh cht th
ng ca kiu vn chuyn ny.
Kch thc phn t: Thng thng cc cht c kch thc cng ln th vn chuyn cng chm. Tuy
nhin, chng cng phi p ng c yu cu l khng phn cc v khng tch in.
Tnh cht ca phn t: Cc cht c tch in thng kh i qua mng. V d, cc ion mc d c
kch thc nguyn t nhng rt kh i qua c lp kp phospholipid, trong khi , phn t O2 v CO2 th
li c th i qua d dng. Cc hp cht ho tan c trong lipid th cng i qua c mng nh alcol,
aldehyd, ceton, glycerol v.v... Cn cc hp cht ho tan trong nc hu nh u b lp k nc ca mng
y ra ngoi.
Hnh 1.20. nh hng ca Gradient nng n tc vn chuyn cht qua mng
A. Qua cc knh Protein, B. Thm thu trc tip

Trong nghin cu y dc hc, ngi ta li dng tnh cht d thm qua mng ca hp cht lipid to
ra cc liposome lm phng tin vn chuyn thuc vo t bo. l cc ti mng lipid kp c kch thc
khong 50nm, bn trong c cha cc hp cht cn a vo t bo.
S vn chuyn th ng cc cht din ra kh d dng v c th thc hin qua hai con ng: thm trc
tip qua mng kp phospholipid hoc i qua cc knh protein dn truyn. Hai con ng ny c mt s khc
bit v tc vn chuyn cc cht. Vi cch thm trc tip qua mng, tc vn chuyn s t l thun vi
gradient nng . Cn khi i qua knh protein, tc v gradient ch t l thun trong mt gii hn nht
nh. Khi qu gii hn , d tng gradient th tc cng khng tng na. S d c hin tng bo ho nh
vy l v s lng cng nh kh nng hot ng ca knh protein trn mng l c hn. S chnh lch nng
cng cao, cc phn t i qua knh cng nhiu. Nhng n khi cc knh hot ng ht cng sut th tc
khng th tng ln na.
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4.3. Vn chuyn cc cht c km theo tiu ph nng lng

y l hnh thc vn chuyn cc cht khng ph thuc vo gradient nng nh s dng nng lng
ho hc. Nh c tnh cht nh vy nn c th gi hnh thc vn chuyn ny l vn chuyn ch ng. Vn
chuyn ch ng bt buc phi thc hin thng qua cc knh dn truyn c bn cht l protein xuyn mng.
Vn chuyn ch ng c vai tr rt quan trng trong t bo. N gip t bo c th ch ng thit lp
nng ni bo ca cc cht theo nhu cu m c khi l chnh lch vi mi trng rt nhiu (v d Na+, K+,
Cl-, HPO42-,...). S duy tr ny to iu kin n nh cho t bo hot ng bnh thng. Vn chuyn ch
ng khng ch din ra gia t bo vi mi trng m cn gia cc phn ca t bo vi nhau. Bm H+ l
mt v d nh vy. Bm ny nm trn mng ca ty th v lc lp vi vai tr to s chnh lch v nng H+
duy tr hot ng chc nng ca cc bo quan ny.
Bng 1.1. Nng ca mt s ion trong v ngoi t bo
H+.
Ion
Nng ngoi bo
Nng ni bo
+
140mM
10mM
Na
4mM
140mM
K+
2+
2,5mM
0,1 M
Ca
100mM
4mM
Cl
Trong t bo c 3 loi knh (bm) vn chuyn rt quan trng l: knh Na+/ K+,
Chnh lch
14x
35x
25000x
25x
knh lin kt v knh
Knh Na+/K+: Cc t bo ng vt c kh nng duy tr nng ni bo ca Na+ rt thp, cn K+ rt
cao so vi mi trng ngoi bo. Kh nng l do trn mng sinh cht c rt nhiu knh Na+/ K+, chng
vn chuyn mt cch ch ng Na+ ra ngoi v a K+ vo trong t bo. Nng lng thc hin s vn
chuyn ly t phn t ATP c sn sinh trong qu trnh h hp t bo. Ngi ta tnh ra c ti 1/3 lng
ATP c sn sinh trong t bo ng vt l duy tr hot ng ny. Bi vy, khi s h hp b nh tr th
hot ng vn chuyn Na+, K+ cng b ngng li.
Mt trong nhng t bo thc hin tch cc vic vn chuyn Na+, K+ l t bo thn kinh. Thng qua vic
a Na+ ra ngoi v K+ vo trong, t bo to nn s khc bit v nng ion mt trong v mt ngoi
ca mng, qua to nn in th mng. Vic thay i t in th tnh sang in th hot ng l c s
dn truyn xung thn kinh. Ngoi ra, nng Na+/ K+ cn gin tip nh hng n cc hot ng khc ca
t bo. V d, t bo c tim, nu v mt l do no y m s vn chuyn Na+ khng thc hin c, lc
nng Na+ ni bo tng cao s gin tip lm nng Ca2+ tng theo v kt qu l s co bp ca c tim
tng ln.
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Hnh 1.21. C ch hot ng ca knh Na+/K+

Knh Na+/K+ cn c tn gi khc l Na+, K+ adenosintriphosphatase (Na+, K+ ATPase). y l mt
loi protein xuyn mng c khi lng phn t 270.000D. Knh c to nn t 4 tiu n v: 2 v 2.
Vic thu phn ATP ly nng lng c thc hin ti mt khu vc trn tiu n v . Hot ng ca knh
din ra theo chu k. u tin, 2 ion Na+ s bm vo mt trong ca knh. Vi s c mt ca Mg2+, knh s
tin hnh thu phn ATP ly gc phosphat cao nng v gii phng ra ADP. Sau khi c phosphoryl ho
bng cch tip nhn gc phosphat ca ATP, knh Na+/K+ s b bin i th hnh v nh 3 ion Na+ ra ngoi.
Ngay sau khi nh Na+, 2 ion K+ t mi trng s bm vo mt ngoi ca knh. S bin i th hnh ca
knh Na+/K+ ch din ra trong giy lt ri li gii phng nt gc phosphat v quay tr li hnh dng ban u.
Qu trnh ly li th hnh ban u cng l lc 2 ion K+ c vn chuyn v nh vo trong t bo. Nh vy,
sau mt chu k c 3 ion Na+ c chuyn ra ngoi v 2 ion K+ c a vo trong qua knh Na+/K+.
Knh lin kt: Knh ny gm nhiu loi v c vai tr vn chuyn ng cng nh cc acid amin. Vic
vn chuyn cng cn s tham gia ca ATP v cc ion. T lin kt y ch qu trnh dn truyn cc
cht khng din ra mt cch n l m i km vi s vn chuyn cc ion. V d, qu trnh vn chuyn
glucose c thc hin ng thi vi vn chuyn Na+ qua mng theo cng hng. Khi knh Na+/K+ hot
ng, n s bm Na+ ra ngoi t bo v to s chnh lch v nng rt ln gia hai bn mng. Lc ny,
Na+ c xu hng khuch tn tr li qua knh lin kt. S khuch tn ca Na+ tng i ln, v th n h tr
v gip cho phn t glucose i qua mng mt cch d dng, thm ch l c th i ngc vi gradient nng
.
Knh H+: Nh trnh by trn, knh H+ khng nm trn mng sinh cht m nm trn mng ca ty
th v lc lp. Nng lng knh H+ hot ng ly t cc in t cao nng c to ra thng qua qu trnh
oxy ho cc hp cht hu c hoc hp th nng lng photon nh sng din ra trong cc bo quan ny. Qu
trnh hot ng ca knh s gip to ra s chnh lch v nng H+ gia hai bn mng. Mc ch ca s
chnh lch v nng H+ l to ng lc cho qu trnh tng hp ATP theo phng thc ho thm (trnh by
nhng chng sau).
4.4. S nhp bo v xut bo
Trong hai qu trnh trnh by trn, s vn chuyn c thc hin mc cc phn t. Ngoi ra,
mt s t bo cn c mt phng thc khc cho php vn chuyn c khi vt cht i vo hoc ra khi t bo.
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l nhp bo v xut bo. y l mt hnh thc vn chuyn ch ng. Trong , vt cht s c vn

chuyn thng qua cc ti mng.
4.4.1. S nhp bo (endocytosis)
Hin tng nhp bo l s hnh thnh cc bng ni bo do s lm vo v tch ra ca mt phn mng c

cha mt khi rn hoc dch lng bn trong. Nhp bo c chia thnh 3 dng: m bo (pinocytosis), thc
bo (phagocytosis) v nhp bo-th th (receptor-mediated endocytosis).
m bo: l hin tng bt gi v a vo t bo cc git cht lng ngoi bo. Trong dch lng c cha
cc cht ho tan m t bo s dng. Din bin ca qu trnh m bo nh sau: mt phn mng sinh cht lm
vo thnh ti ng knh khong 0,1m, cht lng ngoi bo s trn vo lng ti, ming ti khp li, ti tch
ra khi mng tr thnh mt bng m bo v c a su vo t bo cht.
Thc bo: l hin tng hnh thnh v a vo t bo cc bng c kch thc ln (1 - 2m) c mng
bao bc v cha cc phn t rn, vi khun hoc mnh v t bo,... Hin tng ny c mt s bch cu, c
bit l bch cu mono v mt s ng vt nguyn sinh (v d amp). thc bo, mng sinh cht khng lm
vo nh m bo m thay vo c s hnh thnh chn gi vy bt cc phn t ngoi bo hnh thnh
cc ti c ng knh ln c gi l phagosome. V cu to, ti thc bo cng c cu to mng ging ti
m bo nhng khc ch c thm cc vi si actin bao bc bn ngoi.
Nhp bo - th th:
Hnh 1.22. Qu trnh nhp bo - th th

A. nh chp bng knh hin vi in t; B. M hnh.
Nhp bo th th l dng nhp bo, trong to thnh cc bng nhp bo do s lm vo v tch ra ca

mt phn mng c bit c cha nhm th th (receptor). Cc bng ny c bao bc bi mt h thng si
clathrin. Clathrin l mt loi protein gm 6 mch polypeptid (3 mch di v 3 mch ngn) xp vi nhau to
thnh dng king 3 chn. Khi cc receptor c gn vi cc phn t mang tn hiu c trng n t mi
trng (ligand), h thng si clathrin s c hnh thnh ko lm phn mng cha receptor xung, khp
ming li to nn cc bng nhp bo-th th. Bng cch to nn cc bng nhp bo nh vy, t bo thu nhn
c cc cht ho hc mang thng tin t iu chnh hot ng sng cho ph hp (hnh 1.22).
4.4.2. S xut bo (exocytosis)
Xut bo l mt hnh thc vn chuyn cht ra khi t bo qua mng sinh cht. Qu trnh xut bo din ra
ngc vi nhp bo. Cc bng cha cht bi tit c hnh thnh trong t bo cht sau chuyn dn ra
ngoi mng. Mng ca bng xut bo tip s ho nhp vi mng sinh cht v gii phng cc thnh phn
cha trong n ra ngoi t bo. Cng ging nh nhp bo, xut bo l mt qu trnh i hi tiu tn nng
lng. S vn ng ca bng xut bo din ra nh tc ng ca h thng vi si, vi ng c trong t bo cht.
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Thnh phn c a ra ngoi nh bng xut bo cng kh a dng. c th l cc cht cn b, cc

sn phm tha ca qu trnh tiu ho ni bo hoc cng c th l nhng sn phm c chc nng m t bo
chu trch nhim sn xut (v d cc enzym tiu ho, cc hormone,...).
4.5. S trao i thng tin qua mng t bo
Bn cnh s trao i cht, t bo cn trao i thng tin vi mi trng bn ngoi. Nhng thng tin c
th l nhng tn hiu ho hc, nhit , nh sng, B phn tip nhn thng tin chnh l cc th th
(receptor) gn trn mng sinh cht. Thng qua vic tip nhn nhng thng tin t mi trng m t bo hoc
c th c th iu chnh cc hot ng sng mt cc ph hp. Mi trng bn ngoi y khng ch hiu
n thun l mi trng t nhin m cn l mi trng bn trong c th sinh vt. Ngay c nhng t bo
su bn trong c th sinh vt a bo cng c s tip nhn thng tin.
sinh vt a bo, s phi hp hot ng ca cc t bo trong cng mt c th l v cng cn thit. C
th sinh vt thc hin iu thng qua mt h thng cc tn hiu, ch yu di dng cc cht ho hc
(hormone). Khi tip nhn tn hiu hormone, cc t bo s lp tc iu chnh hot ng trao i cht mt cch
ph hp. Khong cch gia t bo pht v t bo nhn tn hiu c th cch xa m cng c th ngay cnh
nhau. Mi trng hp nh vy u c phng thc gi v nhn thng tin tng ng.
i vi cc t bo xa nhau, s pht v nhn tn hiu c thc hin thng qua cc hormone v th th
mng. Cc tn hiu hormone ny xut pht t cc tuyn ni tit v i n cc ni trong c th thng qua h
thng mch mu. V d, hormone glucagon do tuyn o tit ra c tc ng hot ho cc t bo gan, lm tng
cng gii phng glucose vo mu... Mt tn hiu c th mang ngha kch thch m cng c th l c ch
cc hot ng ca t bo. Glucagon lm tng glucose trong mu, cn isulin li c vai tr ngc li. Cc th
th mng, b phn tip nhn thng tin t bo, c bn cht l cc protein xuyn mng. Mt u ca th th
th ra ngoi mng v l ni tip nhn cc phn t mang tn hiu t bn ngoi. S hnh thnh phc h phn t
mang tn hiu - th th (ligand-receptor) s to nn mt lot cc bin i dy chuyn, cui cng lan truyn
n u kia ca th th v lm thay i hot ng ca t bo. Sau khi thc hin chc nng, hormone s ri ra
ngoi v th th li tr v trng thi bnh thng. Trong nhiu trng hp, sau khi chuyn vo trong t bo,
thng tin c th c khuch i ln rt nhiu ln. V d, i vi hormone glucagon l t 105 n 109 ln. V
th nn ch cn mt lng rt nh hormone cng lm bin i trn din rng hot ng ca cc t bo
trong c th.
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Hnh 1.23. M hnh cu to v hot ng ca xynap
Khi hai t bo cnh nhau mun trao i thng tin th iu s c thc hin thng qua cc cu trc
chuyn bit nm mng. i vi hai t bo thn kinh, l cc xynap ho hc hoc xynap in. Cn hai
t bo bnh thng th l cc cu ni t bo (gap).
CU HI LNG GI
1. Hy trnh by c im cu to ca t bo Prokaryota.
2. Hy trnh by c im cu to v chc nng ca cc thnh phn c trong t bo Eukaryota.
3. Trnh by cu trc ca mng sinh cht theo m hnh khm ng. Tnh linh hot ca mng c th
hin ra sao ?
4. C ch vn chuyn vt cht quang mng theo phng thc th ng l g ? Tnh cht ra sao ?
5. Th no l vn chuyn ch ng ? Trnh by c im ca ba knh vn chuyn ch ng quan trng
trong t bo: knh Na+/K+, knh lin kt v knh H+.
6. Qu trnh xut/ nhp bo l g ? Din bin ra sao ?
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Chng 2
S TRAO I CHT V NNG LNG
MC TIU
1. Trnh by c khi nim v G, vai tr ca nng lng hot ha.
2. Trnh by c m hnh phn t v chc nng ca ATP, phng thc tng hp ATP trong t bo
3. Trnh by c c im cu trc v chc nng ca enzym, c ch hot ng ca enzym, gii thch
c tnh c hiu ca enzym.
4.
Trnh by c nh hng ca cc nhn t nhit , pH, nng enzym/c cht, cc cht c ch ln

tc phn ng do enzym xc tc
5.
Trnh by c khi nim v cc giai on chnh ca qu trnh h hp hiu kh. So snh h hp hiu
kh vi h hp k kh.
6.
Trnh by c khi nim v cc giai on chnh ca qu trnh quang hp.
1. NNG LNG SINH HC

Nghin cu v nng lng sinh hc l lnh vc nghin cu v s bin i nng lng din ra km vi
cc phn ng sinh ho trong c th. Ging nh cc phn ng ho hc, phn ng sinh ho l qu trnh mt
hoc mt vi c cht ny c chuyn ho thnh mt hoc mt vi c cht khc. Trong c th sinh vt, i
a s cc phn ng sinh ho ny u din ra vi s c mt ca cc cht xc tc. V d, phn ng bin i
dihydroxyaceton phosphat thnh glyceraldehyd 3 - phosphat cn mt xc tc c tn l triose phosphat
isomerase.
1.1. S bin i nng lng t do

Thng s nhit ng hc quan trng nht trong nghin cu nng lng sinh hc l s bin i nng
lng t do, c k hiu l G (ch G xut pht t tn J. Willard Gibbs , ngi c nhiu ng gp quan
trng trong lnh vc nghin cu nhit ng hc). y l s bin i nng lng din ra trong mt phn ng
iu kin nhit v p sut khng i (iu kin thng thng ca cc phn ng sinh ho). i vi phn
ng bin i dihydroxyaceton phosphat thnh glyceraldehyd 3- phosphat nu trn, ta c G c tnh
nh sau:
Trong : G:
G0:
s bin i nng lng t do

s bin i nng lng t do tiu chun
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R:
hng s kh
T:
nhit tuyt i
[G3P]:
nng glyceraldehyd 3 -phosphat
[DHAP]: nng dihydroxyaceton phosphat

Vi phn ng
ta c:
G0 tng ng vi s bin i nng lng din ra khi mt mol cht tham gia phn ng c chuyn
ho thnh mt mol sn phm to thnh v cc cht trong phn ng u c nng 1M. Trong c th, cc
phn ng din ra trong mi trng nc m lng nc l khng i. V vy, ngi ta coi tc ng ca nng
nc bng 1.
G c th mang gi tr m, dng hoc bng khng. Phn ng c G dng c gi l thu nhit
(endergonic). Trong phn ng ny, mc nng lng t do ca sn phm cao hn mc nng lng t do ca
cc cht tham gia phn ng. Bi vy, phn ng din ra c cn c s cung cp nng lng t mi trng
bn ngoi. Phn ng c G m c gi l phn ng pht nhit (exegonic). Lc ny, mc nng lng t do
ca sn phm thp hn cc cht tham gia phn ng. Mt phn ng c G m thng c xu th din ra mt
cch t pht.
1.2. Nng lng hot ho

Mc d phn ng pht nhit c xu th din ra t pht nhng c th thy rng chng t khi xy trong iu
kin bnh thng. V d nh cc phn ng chy, xng hay cn u khng th t bc chy m cn c mt
nhit lng ban u khi mo phn ng. Nng lng dng khi mo phn ng c gi l nng
lng hot ho. Vn tc ca mt phn ng khng ph thuc vo vic chng gii phng ra bao nhiu nng
lng m ph thuc vo mc nng lng hot ho. Cc phn ng c mc nng lng hot ho ln hn
thng c khuynh hng din ra chm hn.
Vai tr ca nng lng hot ho l gip ph v cc lin kt vn c ca phn t cht tham gia phn ng
t hnh thnh nn cc lin kt c mc nng lng thp hn. Nng lng hot ho khng phi l mt
hng s c nh. Chng ph thuc vo trng thi ca cc lin kt trong phn t. Nu c mt tc ng no
khin cho cc lin kt ny b yu i th nng lng hot ho cn thit cng s thp hn. Tc ng ny c
gi l s xc tc. Trong c th sinh vt, cht xc tc c gi l cc enzym.
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Hnh 2.1. Mc nng lng hot ha ca phn ng trong iu kin bnh thng v khi c enzym xc tc
1.3. ATP tin t nng lng ca c th

Ngun nng lng cung cp hng ngy cho c th con ngi cng nh cc sinh vt khc c ly t s
oxy ho cc hp cht hu c m quan trng nht l glucose. Khi oxy ho hon ton 1 mol glucose, bn cnh
sn phm CO2 v H2O, ta c nng lng gii phng ra l 2870kJ (Go = - 2870kJ). S nng lng ny
khng gii phng ra cng mt lc m din ra mt cch t t v phn ln c s dng tng hp phn t
ATP, mt hp cht c coi l tin t nng lng ca c th.
ATP (adenosin triphosphat) l mt hp cht giu nng lng, c vai tr nh mt ngi cung cp nng
lng ho hc cho cc hot ng ca t bo nh vn chuyn ion, vn ng t bo hay cc phn ng sinh tng
hp v.v..
V mt cu to, phn t ATP c hnh thnh t ba phn l:
Gc adenin.
ng ribose.
Ba gc phosphat lin nhau.
Ba gc phosphat l thnh phn mu cht quyt nh c tnh ca phn t ATP. Chng c ni vi nhau
bi cc lin kt giu nng lng. Nng lng c gii phng ra nhiu nht khi lin kt gia hai gc
phosphat ngoi cng b ph v, tng ng khong 30kJ/mol. Nng lng ny nhiu gp hai ln nng
lng hot ho trung bnh ca cc phn ng trong t bo.
Cc hot ng ca c th, su hn l ca t bo thc ra u l mt chui cc phn ng sinh ho khc
nhau v ATP chnh l ngun cung cp nng lng hot ho ph bin chng din ra. Nng lng ny c
gii phng thng qua s phn tch phn t ATP thnh ADP v gc phosphat. Thng thng, phn ng phn
tch ATP (phn ng to nhit) v phn ng cn nng lng (phn ng thu nhit) c kt hp vi nhau theo
cp, tc l din ra cng nhau, cng thi gian, ti cng a im v c xc tc bi cng mt phc h
enzym. Nng lng gii phng ra s c s dng gn gc phosphat vi cht tham gia phn ng, chuyn
chng thnh dng hot ng. Qu trnh ny c gi l s phosphoryl ho.
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Hnh 2.2. Cng thc cu to phn t ATP
Trong t bo, ATP c to ra theo hai cch l phosphoryl ha mc c cht v tng hp ho thm.
Phosphoryl ho mc c cht
L s hnh thnh ATP thng qua vic gn ADP vi gc phosphat ly t mt hp cht hu c. Phn ng
ny din ra di s xc tc ca enzym. V d:
Phosphoenolpyruvat + ADP Pyruvat + ATP
Phn ng trn c xc tc bi enzym pyruvat kinase. y l mt phn ng c G m. Nng lng
cha trong lin kt gia gc phosphat cao nng v phn t pyruvat l 62kJ/mol, trong khi lin kt gia gc
phosphat v ADP ch l 30kJ/mol. Bi vy, phn ng xy ra khng kh khn. Tng hp ATP theo phng
thc phosphoryl ho mc c cht l qu trnh c mi loi t bo.
Tng hp ho thm
y l qu trnh tng hp ATP thng qua s thm thu ca ion H+ qua mng cc bo quan chuyn bit
trong t bo (v d: ty th, lc lp).
Theo qu trnh ny, t bo s s dng cc in t cao nng bm v to s chnh lch v nng ion
+
H gia hai bn mng. Cc in t cao nng c s dng c th bt ngun t phn ng oxy ho cc hp
cht hu c trong qu trnh h hp hoc qua qu trnh hp th nng lng nh sng Mt Tri trong quang
hp. Sau khi thit lp s chnh lch v mt nng , cc ion H+ s c xu hng khuch tn ngc tr li, t
ni c nng cao quay v ni c nng thp hn. Qu trnh khuch tn ca ion H+ khng c thc hin
trc tip qua lp mng phospholipid m thng qua cc knh c bit c s kt hp vi enzym sinh tng hp
ATP. S vn ng ca cc ion H+ qua knh s to ng lc enzym gn ADP vi gc phosphat v c v
to nn phn t ATP.
So vi tng hp theo phng thc phosphoryl ho mc c cht, tng hp ho thm c hiu sut cao hn
rt nhiu. Phn ln ATP ca t bo Eukaryota u c tng hp theo phng thc ho thm v ty th l bo
quan chu trch nhim chnh trong vic thc hin qu trnh ny.
2. S TRAO I CHT TRONG T BO

S trao i cht c hiu l ton b cc phn ng ho hc din ra trong c th sinh vt. Tn gi
metabolism (trao i cht) xut pht t mt t Hy Lp c ngha l s thay i. Cc nghin cu cho thy
rng, gn nh ton b cc phn ng din ra trong h thng sng u c xc tc bi cc enzym.
Cc phn ng trao i cht khng din ra mt cch ngu nhin, v trt t. Chng c sp xp theo
nhng trnh t c hiu ph hp vi tng mc ch nht nh. Trnh t ny c gi l cc con ng trao
i cht. Mt con ng trao i cht c th gm t 2 cho n 20 phn ng ni tip nhau cho php bin i
mt hp cht ban u thnh sn phm mong mun ca t bo; cc thnh phn tham gia vo chui bin i
c gi l cc cht chuyn ho.
Qu trnh trao i cht trong c th din ra theo hai hng l ng ho v d ho. D ho l qu trnh
phn tch cc hp cht hu c ln, phc tp thnh cc sn phm n gin v c kch thc nh hn. Mt s
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phn ng trong qu trnh d ho gii phng ra nng lng v mt phn nng lng ho hc ny c s
dng tng hp ATP cho t bo. Ni cch khc, d ho cung cp cho t bo ngun nng lng v cc n
v cu trc cn cho hot ng sng. Ngc vi d ho, ng ho l qu trnh lp rp cc phn t n gin
thnh cc sn phm ln, c cu trc phc tp. Cc phn ng sinh tng hp ny cn c cung cp nng
lng ho hc, ch yu t ATP.
Nhn chung, cc con ng sinh tng hp ca mt hp cht khng n gin l s o ngc li ca qu
trnh phn gii cht . V d, s chuyn ho phn t glucose thnh pyruvat gm 10 phn ng, trong khi qu
trnh ngc li cn ti 13 phn ng v gia hai qu trnh ny ch c 9 phn ng l chung nhau. Cc con
ng trao i cht c th c lp vi nhau hoc cng chia s nhng cng on trung gian. Kt qu l chng
an xen vi nhau to thnh mt mng li trao i cht v t bo s dng mng li ny nh l phng tin
sng ca mnh. Khng c trao i cht, t bo khng th tn ti.
Tnh c lp ca hai mt phn gii v tng hp cht hu c cho php chng c cc c ch iu ho
ring. Tc sinh tng hp v phn gii c th c tng ln hoc gim i mt cch c lp da vo cc
phn ng ring ca mi qu trnh. Nu tt c cc phn ng u chung nhau th khi tc ng s khin cho c
hai qu trnh b thay i ging nhau v s iu ho tr nn khng c kt qu. S iu ho trao i cht
thng xy ra bc u tin hoc cc cng on ban u ca chui trao i cht, v cc phn ng c
iu ho thng khng phi l nhng phn ng thun nghch.
3. ENZYM
Nh trnh by trn, i a s cc phn ng trao i cht trong t bo u din ra vi s xc tc ca
enzym v y cng l i tng t bo thc hin s iu ho i vi qu trnh trao i cht. Cho n nay,
ngi ta bit v phn loi c khong 3500 enzym.
3.1. Thnh phn v cu to

Thnh phn ho hc ca i a s enzym l protein. Bn cht ho hc ca enzym ch c xc nh
ng k t khi ngi ta kt tinh c chng. Enzym u tin c kt tinh l urease ca u tng vo nm
1926, tip theo l pepsin v tripsin vo nm 1930 v 1931. Cho n nm 1977, ngi ta kt tinh c gn
200 loi enzym khc nhau. Nhng phn tch ho hc cho thy, cc tinh th enzym nhn c u l cc
protein. Tuy nhin, vo nm 1981, mt nh khoa hc tn l Thomas R. Cech lm vic ti i hc Colorado
pht hin thy rng, mt s on trn phn t ARN c kh nng xc tc cho s ct ri ca chnh n. iu
ny cho thy khng phi ton b cc enzym u l protein.
Da vo thnh phn ho hc, ngi ta chia enzym ra lm hai loi: enzym n gin v enzym phc tp.
Enzym n gin (hay cn gi l enzym mt thnh phn): khi thu phn ch cho ra thnh phn duy nht
l cc loi acid amin.
Enzym phc tp (hay cn gi l enzym hai thnh phn): ngoi phn acid amin ra (gi l apoenzym), sau
khi thu phn, ngi ta cn thu c mt hoc mt s phn khc khng phi l protein gi l nhm ngoi
(cofactor). Cc nhm ngoi c th l cc ion kim loi nh Mg2+, Fe2+ hoc l cc phn t hu c nh gi l
coenzym. Coenzym thng c quan h mt thit vi cc vitamin, cc vitamin l nguyn liu th to nn
coenzym. Vai tr ca nhm ngoi l h tr qu trnh xc tc ca enzym. Ngi ta thy rng, nhiu enzym s
khng th hot ng chnh xc khi thiu cc nhm ngoi. Chng hot ng nh l cu ni gia enzym vi c
cht v tham gia trc tip vo qu trnh xc tc. Phn ln cc nhm ngoi (khong 80%) ch gn vi enzym
bng nhng lin kt lng lo v s lin kt ny ch hnh thnh khi enzym hot ng xc tc. Cc nhm ngoi
c th c ti s dng nhiu ln.
V mt cu trc, enzym thng c dng cu, c khi lng phn t ln. Enzym c khi lng nh nht
hin nay l ribonuclease (12.700 dalton). Cn li a s nm trong khong 20.000 n 90.000 hoc vi trm
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nghn, mt s enzym c khi lng ln n mt triu hoc ln hn.

Trn enzym c mt vng c bit c gi l trung tm hot ng (TTH). TTH c cu to nh mt
vng lm trn b mt enzym v l ni tip xc trc tip vi c cht (cht tham gia phn ng, b chuyn ho
di tc ng ca enzym), thc hin vic ct t hoc hnh thnh cc lin kt mi trn c cht to thnh
sn phm. T kt qu nghin cu trn cc enzym, ngi ta rt ra mt s c im chung ca TTH l:
Ch chim mt t l th tch tng i nh trn enzym.
c cu to bi nhiu nhm chc khc nhau ca acid amin, cc phn t nc lin kt, trong nhiu
trng hp cn c c cc ion kim loi, cc nhm chc ca coenzyme.
S phn b ca cc nhm chc khin cho TTH c cu trc khng gian phc tp, c gi vng nh
mng li lin kt hydro. Cu trc khng gian ca TTH ch ph hp vi mt hoc mt s c cht nht
nh.
mt s enzym, ngoi TTH ra cn c mt trung tm khc gi l trung tm iu ho. y l ni
enzym kt hp vi cc cht iu ho. Khi s kt hp din ra, cu trc khng gian ca enzym s b thay i,
trong c cu trc ca TTH, do hot tnh xc tc cng b thay i theo. Nu s thay i din ra theo
hng tng hot tnh enzym th gi l iu ho dng, cn nu lm gim hot tnh th gi l iu ho m.
Cht iu ho mc d gn vi enzym nhng khng b enzym tc ng.
3.2. C ch hot ng ca enzym

Phng thc hot ng ca enzym l lm tng tc phn ng thng qua vic lm gim nng lng
hot ho. V d, phn ng phn hu H2O2 gii phng H2O v O2. Khi c mt enzym xc tc l catalase,
tc ca phn ng c th tng ln 10.000 ln.
xc tc, enzym s kt hp tm thi vi c cht ti TTH to nn phc hp enzym - c cht. Cc
nhm chc ca TTH s tc ng ln c cht ph v v hnh thnh cc lin kt mi. Phc enzym - c
cht rt km bn vng, c cht nhanh chng b chuyn ho v gii phng ra sn phm cng enzym t do. i
tng b bin i trong phn ng ch l c cht, cn enzym khng b tc ng v c ti s dng. Cc loi
lin kt ch yu c to thnh gia enzym v c cht trong phc l: tng tc tnh in, lin kt hydro,
tng tc Van der Waals. Mi loi lin kt i hi nhng iu kin khc nhau v chu nh hng khc nhau
khi c nc.
gii thch kh nng kt hp ca enzym v c cht, chng ta s nghin cu hai m hnh l gi thuyt
kho - cha kho v gi thuyt khp cm ng.
Gi thuyt kho - cha kho
Gi thuyt ny do Emil Fischer a ra nm 1890. Theo ng, TTH ca enzym vn c cu trc khng
gian tng ng vi cu trc phn t c cht (tng t nh s tng ng gia cha kho v kho). Do c s
ph hp nh vy nn enzym c th lin kt vi phn t c cht xc tc. V khng phi c cht no cng
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c cu trc ph hp vi TTH nn enzym ch xc tc cho mt s phn ng v c cht nht nh. Gi

thuyt kho - cha kho l mt im mc trong lnh vc nghin cu enzym v n vn cn gi phn ln
gi tr n ngy nay. Tuy nhin, gi thuyt ca Fischer cn n gin v cha th m t ht s hot ng ca
enzym.
Gi thuyt khp cm ng
Gi thuyt khp cm ng ra i vo nm 1958 do Daniel Koshland xut. Theo , phn t enzym
khng cng nhc m c cu trc ng. S tng ng v cu hnh khng gian ca TTH v c cht ch c
hnh thnh trong qu trnh enzym tip xc vi c cht thay v tng ng ngay t u nh m hnh ca E.
Fischer. Khi tip xc vi c cht, cc nhm chc trong TTH ca enzym s thay i v tr trong khng
gian, to thnh hnh th khp vi hnh th ca c cht, qu trnh ny c gi l khp cm ng. y chnh
l quan nim hin nay v c ch hot ng ca enzym.
3.3. Tnh c hiu ca enzym

Tnh c hiu chnh l mt trong nhng im khc bit ca enzym so vi cc cht xc tc v c thng
thng. Mi enzym ch c kh nng xc tc cho s chuyn ho ca mt hay mt s cht nht nh, theo mt
kiu phn ng nht nh. c tnh ny c quy nh do s phc tp v cu trc khng gian ca TTH ca
enzym. Ch nhng c cht hay nhm c cht no ph hp th mi c th gn vi TTH to nn phc h
enzym - c cht v b bin i. Tnh c hiu ca enzym c chia lm hai loi:
c hiu kiu phn ng
Tnh c hiu ny th hin ch mi enzym ch c th xc tc cho mt trong cc kiu phn ng chuyn
ho mt cht nht nh. V d, phn ng oxy ho kh, phn ng thu phn, phn ng ng phn ho v.v...
c hiu c cht
Mc c hiu c cht ca cc enzym l khc nhau. Ngi ta chia ra hai mc c hiu c cht:
c hiu tuyt i: enzym ch c tc dng trn mt cht nht nh v hu nh khng c tc dng vi
cht no khc. V d, urease hu nh ch c tc dng vi ure:
H2NCONH2 + H2O CO2 + 2NH3
Tuy nhin, v sau ngi ta thy rng, urease cn c tc dng vi hydroxyure (mt cht c cu trc
gn ging ure) nhng vi tc nh hn 120 ln so vi tc dng vi ure. Mt v d khc v tnh c hiu
tuyt i l enzym Lactat dehydrogenase. Enzym ny ch tin hnh xc tc vi ng phn dng L ca acid
lactic, cn ng phn dng D khng c tc dng.
c hiu tng i: enzym c kh nng hot ng trn nhiu c cht khc nhau c nhng c im
chung v mt cu trc. V d, carboxyesterase c th thu phn este ca nhiu loi acid carboxylic khc
nhau.
3.4. Cc nhn t nh hng n vn tc ca phn ng c xc tc bi enzym

Tc ca phn ng do enzym xc tc ph thuc vo nhiu yu t nh: nng enzym, nng c
cht, nhit , pH mi trng, cc cht c ch v.v... Trong , i a s cc yu t ny c lin quan n c
ch hot ng v c im cu to ca enzym.
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3.4.1. Nhit
Trong tt c cc phn ng ho hc, vic tng nhit s lm tng nng lng ng hc ca cc phn t
tham gia phn ng. Qua khin chng c tn s va chm tng ln v lm tc phn ng tng theo. Ngi
ta nh gi nh hng ca nhit n tc phn ng ho hc thng qua h s nhit Q10. Q10 c tnh
bng t s gia hng s tc ca phn ng mt nhit no so vi khi din ra nhit thp hn
nhit 10C.
Trong : kt l tc phn ng nhit t;

kt+10 l tc phn ng nhit t + 10.
Q10 ca phn ng ho hc thng thng l 2 n 3, trong khi ca phn ng enzym ch l 1 n 2. Khi
tng nhit , tc ca phn ng enzym cng tng nhng khi t n mt ngng nhit no , tc
s li gim xung v phn ng c th b dng li. S d c iu ny l do hu ht enzym c cu to t
protein. nhit cao, phn t protein b bin tnh v enzym khng cn hot tnh na.
Mi enzym c mt ngng nhit ti u (hot tnh mnh nht) v mt ngng nhit ti hn (mt
hot tnh hon ton). Thng thng, ngng nhit ti u nm khong 40 50C, cn nhit ti hn l
khong 70C. Tuy nhin, trong nhng trng hp c bit, nhit ti u ca enzym c th rt cao hoc rt
thp. V d, enzym mt s vi khun sng sui nc nng c nhit ti u trn 70C, hoc enzym ca mt
s loi c Nam Cc c th hot ng tt 2C. bn ca enzym c th tng ln khi kt hp vi c cht
hoc mt s kim loi.
Hnh 2.4. nh hng ca nhit ti tc ca phn ng enzym
3.4.2. pH
pH mi trng tc ng r rt n phn ng enzym v n nh hng n mc ion ho ca c cht v
enzym. pH 7 thch hp cho nhiu loi enzym. Khi vt qua khong ti u, cc ion trong mi trng s nh
hng ti cc nhm tch in ca TTH, khin chng mt i kh tng tc vi c cht. Nu pH nm xa
khong ti u, cc lin kt cu to nn enzym c th b ph v v lm enzym bin tnh.
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Hnh 2.5. nh hng ca pH ti tc ca phn ng enzym
3.4.3. Nng enzym v nng c cht

nh hng ca nng enzym, c cht ln tc phn ng c th hin trn hnh 2.6. T th c
th thy, vic tng nng c cht s lm tng tc phn ng. Tuy nhin, s tng ln ny khng ko di
mi m s t n mt trng thi bo ho. Ti y, d c cho tng thm nng c cht, tc ca phn
ng vn s khng tng ln c. Tc ca phn ng ti trng thi bo ho cao hay thp ph thuc vo
nng enzym. Tng ng vi nng enzym cao, ta c tc phn ng ti im bo ho ln. Bi vy, mun
tc phn ng vt qua im bo ho, cn b sung thm enzym.
Hin tng ny c th c gii thch mt cch n gin. Khi hm lng c cht thp, nhiu enzym c
TTH cn trng (lng enzym dng cha ht) nn tc ca phn ng ph thuc vo nng c cht.
Nhng khi lng c cht qu nhiu, ton b enzym u tham gia hot ng, lc ny tc phn ng t
trng thi bo ho. n y, nng enzym li l yu t quyt nh tc phn ng.
Hnh 2.6. nh hng ca nng enzym, nng c cht ln tc ca phn ng enzym
3.4.4. Cc cht c ch enzym

Hot ng ca enzym c th b thay i do tc ng ca cc cht c ch. Ngi ta chia cc cht c ch
enzym ra lm hai nhm ln l cht c ch cnh tranh v cht c ch khng cnh tranh.
Cht c ch cnh tranh
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Cc cht ny c cu trc phn t gn ging vi c cht. Chnh v vy, chng c kh nng cnh tranh vi
c cht gn vo TTH ca enzym. Mt khc, cht c ch cnh tranh li khng b enzym lm bin i nn
chng s gn lu di TTH, khin cho enzym khng c gii phng ra ti s dng. V th nn ch mt
lng nh cht c ch cnh tranh cng c th lm tc phn ng gim ng k.
Mt v d v s c ch cnh tranh l trng hp ca enzym succinic dehydrogenase. Enzym ny bin
acid succinic thnh sn phm l acid fumaric. Acid malonic l mt cht c cu to gn ging vi acid
succinic. Khi c mt cht ny, chng s cnh tranh vi succinic bm vo TTH, khin cho enzym b mt
tc dng.
Hnh 2.7. S c ch ca malonic i vi enzym succinic dehydrogenase
Cht c ch khng cnh tranh
Hnh 2.8. M hnh c ch tc ng ca cht c ch khng cnh tranh
Qu trnh c ch khng cnh tranh ch din ra cc enzym c trung tm iu ho. Cht c ch khng
cnh tranh khc vi cht c ch cnh tranh ch khng gn vo TTH ca enzym. Thay vo , chng s
gn vo trung tm iu ho, lm thay i cu trc ca phn t enzym, to nn hiu ng d lp th (allosteric).
S thay i khin cho enzym tr nn kh khn trong vic gn vi c cht v kh nng thc hin xc tc
b gim xung. Cc cht c ch khng cnh tranh ph bin ca enzym l cc ion kim loi nng nh Hg2+,
Ag+. Mt s cht c nh asen hay cyanid cng tc ng ln enzym theo cch ny. Tuy nhin, cng cn phi
lu rng, khng phi ton b cc cht gn vo trung tm iu ho u c tc dng c ch. Nhiu cht c
th c tc dng ngc li l lm tng hot tnh ca enzym, c gi l cht hot ho. Tc ng vo trung tm
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iu ho ca enzym l mt trong nhng cch quan trng t bo iu chnh s trao i cht ca mnh.
3.5. Cch gi tn v phn loi enzym
Theo quy c quc t, tn gi ca enzym c cu to t ba phn: tn c cht c hiu, tn loi phn
ng v ui ase. V d, glyceraldehyd 3 phosphat dehydrogenase; pyruvatkinase; triosephosphat
isomerase v.v...
Da vo tnh c hiu phn ng ca enzym, t nm 1960, Hi Ho sinh quc t (IUB) thng nht
phn loi enzym thnh 6 lp, nh s t 1 n 6. Cc s th t ny l c nh cho mi lp:
(1) Oxydoreductase: cc enzym xc tc cho phn ng oxy ho kh.
(2) Transferase: cc enzym xc tc cho phn ng chuyn v.
(3) Hydrolase: cc enzym xc tc cho phn ng thu phn.
(4) Lyase: cc enzym xc tc cho phn ng phn ct khng cn nc, loi nc to thnh ni i hoc
kt hp phn t nc vo ni i.
(5) Isomerase: cc enzym xc tc cho phn ng ng phn ho.
(6) Ligase: cc enzym xc tc cho phn ng tng hp c s dng lin kt giu nng lng ca ATP
v.v...
3.6. S iu ho trao i cht thng qua enzym
Trong t bo, cc enzym thng khng phn b mt cch t do trong dch t bo cht. Thay vo ,
chng c gn trn mng theo mt trnh t nht nh, ph hp vi chui phn ng trao i cht. Cc hp
cht hu c s ln lt i qua cc enzym trong chui, tng bc c bin i tr thnh sn phm hon
chnh.
Do i a s cc phn ng trong c th u din ra vi s xc tc ca enzym nn s iu ho qu trnh
trao i cht c thc hin thng qua h thng enzym ny.
Khi mt sn phm c tch lu qu nhiu trong t bo, sn phm c th tr thnh cht c ch khng
cnh tranh v bm vo cc enzym u tin ca chui phn ng, qua , lm ngng qu trnh chuyn ho li.
Hin tng ny c gi l s c ch lin h ngc. Phng thc iu chnh th hai c gi l s hot ho
khai mo. Hin tng ny xy ra khi trong t bo c tha c cht. Nhng c cht ny s ng vai tr hot
ho cc enzym v khi ng chui chuyn ho. Mt phng thc iu ho trao i cht khc l thng qua
iu chnh nng enzym trong t bo. Theo , enzym s c tng hp vi s lng v thi gian thch
hp, qua xc nh gii hn ca s trao i cht trong t bo.
Hnh 2.9. M hnh iu ha trao i cht thng qua enzym
4. H HP T BO
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4.1. i cng v h hp
H hp l qu trnh phn gii cc hp cht hu c gii phng ra nng lng cho t bo hot ng.
Phn ln nng lng ny c chuyn ho thnh ATP, tin t nng lng ca t bo. i vi phn ln c
th, hp cht hu c chnh c s dng trong h hp l glucose. Phng trnh tng qut nh sau:
C6H12O6 + 6O2 6CO2 + 6H2O (G = 2870kJ/mol)
Nu xt v phng trnh tng qut, c th thy rng s phn gii glucose trong h hp khng khc so
vi phn ng t chy glucose nhng v din bin li khc hn nhau. Qu trnh t chy glucose din ra mau
chng, nhit lng c gii phng ra phn ln di dng nhit. Trong khi , s phn gii glucose trong h
hp li l mt chui gm nhiu phn ng. Qua chui phn ng ny, nng lng s c gii phng t t v
phn ln tch lu trong cc lin kt ho hc ca ATP.
S kin ng vai tr mu cht trong vic chit rt nng lng t hp cht hu c l chui phn ng oxy ho
- kh. Trong mt s bc ca qu trnh h hp, hp cht hu c s c oxy ho m qua nng lng
c gii phng ra di dng in t cao nng (e-) i cng vi H+. Ngay tip oxy ho l s kh, trong
cht nhn in t v H+ ph bin ca t bo l phn t NAD+ (nicotinamid adenindinucleotid). Ngi ta gi
qu trnh ny l phn ng i km hay phn ng cp i (couple reaction).
NAD+ + 2e- + 2H+ NADH + H+ (G = + 219kJ/mol)
NADH cha phi l sn phm cui cng ca h hp. Chng s thc hin vai tr vn chuyn in t v
H+ n cho chui h hp ty th. Ti y mi tin hnh bin i nng lng ca in t cao nng thnh
nng lng c tch lu trong phn t ATP theo phng thc ho thm.
Tu vo s c mt ca oxy khng kh, ngi ta chia h hp ra lm hai dng:

H hp hiu kh: xy ra khi c mt oxy khng kh, hp cht c phn gii hon ton thnh CO2 v
H2O.
H hp k kh: xy ra khi khng c mt oxy khng kh, hp cht hu c khng c phn gii hon
ton thnh CO2 v H2O.
Trong chng ny ch yu cp ti qu trnh h hp hiu kh.
4.2. H hp hiu kh
y l qu trnh c mi sinh vt Eukaryota. Bo quan chuyn trch cho h hp hiu kh l ty th. Qu
trnh h hp hiu kh bao gm 4 giai on chnh l ng phn, decarboxyl-oxy ho acid pyruvic, chu trnh
Krebs v dn truyn in t to ATP.
4.2.1. ng phn
Qu trnh ng phn gm 10 phn ng vi hp cht u vo l 1 phn t glucose v sn phm l 2
phn t pyruvat, 2ATP v 2NADH. Din bin ca qu trnh ng phn c th hin hnh 2.10.
Mi phn ng ca qu trnh ng phn c chia lm hai phn: 5 phn ng u tin c gi l giai
on u t nng lng, 5 phn ng cn li gi l pha gii phng nng lng.
Trong pha u t nng lng, t bo bin i glucose thnh phn t fructose-1,6-diphosphat. Qu trnh
ny tiu tn 2ATP. Phn t fructose -1,6- diphosphat tip s c phn tch thnh 2 phn t c 3 carbon
l glyceraldehyd-3-phosphat (G3P). n y kt thc giai on u tin ca ng phn. Nu quan st biu
bin i nng lng ca chui phn ng, c th thy rng tng mc nng lng t do ca sn phm
c y ln cao hn ca glucose. iu ny t c chnh nh vic s dng 2 phn t ATP. Vi hai nhm
phosphat, phn t fructose-1,6-diphosphat c hot tnh mnh m hn so vi glucose. ng thi n c cu
trc gn nh i xng nn vic phn tch tr nn d dng v sn phm c tnh ng nht.
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Hnh 2.10. S bin i nng lng trong chui ng phn
n pha gii phng nng lng, mi phn t G3P c bin i thnh mt phn t acid pyruvic. giai
on ny, ba trong s nm phn ng l gii phng nng lng. Trong hai phn ng, nng lng c s
dng tng hp ATP theo phng thc phosphoryl ho mc c cht. Mt phn ng cn li c s tch in
t, H+ t c cht v kt hp vi NAD+ to ra NADH. giai on u, mi mt phn t glucose phn
tch thnh 2 phn t G3P nn sn phm thu c gm 2 phn t acid pyruvic, 2 phn t NADH v 4 phn t
ATP. Do s dng 2 ATP cho pha u t nng lng nn kt thc ng phn, lng ATP cn li l 2
phn t.
Nu quan st chui phn ng c th thy rng, O2 khng c vai tr g trong qu trnh ng phn ny.
Bi vy m ng phn c th din ra c qu trnh h hp hiu kh ln h hp k kh. Qu trnh ng
phn khng ph thuc vo vic c mt ty th trong t bo hay khng v n din ra trong t bo cht. V vy
nn n c c sinh vt nhn nguyn thy v nhn tht. Ngi ta thy rng qu trnh ng phn c nhng
vi khun c xa tn ti trn Tri t t 3,5 t nm trc v n nay qu trnh ny vn cn tn ti trong cc
c th sng. S khc nhau gia h hp hiu kh v k kh nm nhng giai on sau. Vi s c mt ca oxy
khng kh, acid pyruvic c phn gii tip thnh CO2, H2O v sinh ATP. Cn nu khng c oxy, acid
pyruvic s c chuyn ho thnh cc cht hu c khc. Lng ATP thu c trong h hp k kh ch yu
nm qu trnh ng phn.
Ngoi ng phn, sinh vt cn tn ti nhiu con ng khc bin i glucose thnh acid pyruvic
nh con ng KDPG (2-keto-3 deoxy-6-phosphat gluconat), con ng PP (pentose-phosphat). Vic s
dng con ng no ty thuc vo ngun hydratcarbon m t bo s dng.
4.2.2. Decarboxyl-oxy ho acid pyruvic
Sau khi c to ra trong t bo cht, acid pyruvic s c chuyn vo trong cht nn ty th. Ti y,
acid pyruvic s c bin i thnh acetyl coenzym A (acetyl-CoA) theo phng trnh sau:
CH3-CO-COOH + HS-CoA + NAD+ CH3-CO-S-CoA + CO2 + NADH2
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y l mt qu trnh gm nhiu cng on v c xc tc bi mt phc h a enzym c tn l pyruvatoxydase hay pyruvat-decarboxylase. Kt qu ca chui phn ng l t phn t acid pyruvic c 3 carbon, km
hot ng tr thnh phn t c 2 carbon l acetyl-CoA dng hot ho do c cha lin kt cao nng trong
phn t. Cht ny tip s i vo chu trnh Krebs. Bn cnh , qu trnh decarboxyl-oxy ho pyruvat cn
gip to nn NADH tng hp ATP v sau.
Acetyl-CoA l hp cht c vai tr quan trng trong t bo. N l im trung gian ni nhiu con ng
trao i cht khc nhau. Trong trng hp thiu nng lng, t bo c th phn gii cc acid amin hay cc
acid bo thnh acetyl-CoA ri t i vo chu trnh Krebs tip tc thc hin qu trnh bin i to ra
ATP. Acetyl-CoA cng c vai tr nh l mt n v cu trc t xy dng nn rt nhiu hp cht phc
tp cn thit cho t bo nh tng hp cc acid bo, steroid, aceto acetat v.v...
Hnh 2.11. Qu trnh bin i pyruvat thnh acetyl - CoA
4.2.3. Chu trnh Krebs

Tip sau ca qu trnh decarboxyl-oxy ho acid pyruvic, phn t acetyl-CoA s i vo mt chu trnh
khp kn. Chu trnh ny do mt nh khoa hc ngi Anh l Hans Adolf Krebs (1900-1981) xng. ng
t gii thng Nobel nm 1953 vi chu trnh v sau mang tn mnh, chu trnh Krebs (hay cn gi l chu
trnh acid tricarboxylic).
Chu trnh Krebs gm 9 phn ng chia lm hai giai on: giai on chun b gm 3 phn ng, giai on
chit rt nng lng gm 6 phn ng cn li.
Khi u ca chu trnh, acetyl-CoA s kt hp vi acid oxaloacetic to nn acid citric di s xc tc
bi enzym citrat synthetase. Trong phn ng ny, coenzym A c vai tr quan trng bi n tham gia vo
thnh phn cu to trung tm hot ng ca enzym. Nh c cha coenzym A, gc acetyl trong phn t
acetyl-CoA c th d dng gn vi enzym thc hin kt hp vi acid oxaloacetic. Tip , sau hai bc
loi v thm nc, acid citric c bin i thnh acid isocitric. n y kt thc giai on chun b. Trong
cc phn ng cn li, acid isocitric tng bc c phn gii v ti to li acid oxaloacetic. Mi mt phn t
acetyl-CoA khi i qua chu trnh Krebs s thu c 2CO2, 3NADH2, 1FADH2 v 1ATP.
2
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Hnh 2.12. Chu trnh Krebs
Nh vy, tnh n thi im ny, ton b phn t glucose c phn gii hon ton thnh CO2. Nng
lng gii phng ra trong qu trnh phn gii glucose, phn nh di dng nhit si m t bo, cn li
phn ln c tch tr trong cc lin kt ha hc. Trong cc sn phm quan trng ca chu trnh Krebs, ATP
ch chim mt lng nh, cn phn ln l NADH2 v FADH2. V sau, cc hp cht ny cng s c s
dng bin i thnh ATP, sn phm cui cng ca h hp.
Khng ch thc hin phn gii glucose, chu trnh Krebs cn l im n ca qu trnh phn gii cc acid
amin, acid bo. Bn cnh , chu trnh cn cung cp tin cht t bo tng hp cc cht hu c cn thit.
V d, -ketoglutarat v oxaloacetat thng l nguyn liu tng hp cc acid amin glutamat v aspartat;
NADH2 ngoi vic s dng tng hp ATP cn c dng kh cc lin kt kp, cc nhm carbonyl,
amin trong cc phn ng ca qu trnh trao i cht v.v... C th ni rng, chu trnh Krebs l ng hng
chnh phn gii cc hp cht hu c v cng l im giao lu ca nhiu con ng trao i cht trong t
bo.
4.2.4. Chui truyn in t ca ty th
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Hnh 2.13. Chui truyn in t ty th
Chui truyn in t c nh v ti mng trong ca ty th. Chng thc hin tip nhn in t v H+ t
NADH2 v FADH2 c sn sinh ra trong cc giai on trc tng hp ATP. Tuy nhin, nng lng
ca qu trnh tip nhn ny khng c dng tng hp trc tip ATP ngay m c dng bm ch
ng cc ion H+ t cht nn ty th vo xoang gian mng. Qu trnh ny to nn s chnh lch ln v nng
ion H+ cng nh in tch gia hai bn mng trong ty th. Nhng chnh lch v nng v in tch s
khin cho cc ion H+ c xu hng khuch tn ngc tr li cht nn thng qua cc knh c bit c dng ht
hnh nm (c cung) cng c nh v mng trong ty th. Trong cc ht ny c cha phc h enzym ATP
synthetase. Mi khi cc ion H+ i qua ht, phc h enzym s c kch hot lp rp ADP v gc phosphat
to nn phn t ATP. C 2 ion H+ qua mng th tng hp c 1 phn t ATP. y chnh l s tng hp
ATP theo phng thc ha thm.
Nm u tin ca chui truyn in t l phc h enzym NADH2 dehydrogenase. Enzym ny tin hnh
tip nhn in t v H+ ca NADH2, NAD+ gii phng ra c ti s dng li trong chui trao i cht. Sau
khi tip nhn, NADH2 dehydrogenase s a H+ vo xoang gian mng, cn in t s c chuyn n cho
mt cht mang gi l ubiquinon. n lt ubiquinon s chuyn in t cho mt phc h protein quan trng
gi l cytocrom b-c. Cc cytocrom trong phc h s ln lt truyn in t cho nhau. Bn trong cu trc ca
mi cytocrom l nhn hem cha ion Fe3+. Vic cho v nhn in t s khin cho cc nhn hem lun phin b
kh v oxy ha.
Fe3+
e-
Fe2+
Mt s bc trong chui oxy ha kh s gii phng nng lng v c dng bm ion H+ qua mng.
Sau khi i qua cytocrom b-c, in t s c chuyn n mt cht mang khc l cytocrom C. Cht ny s
em in t n cho phc h cytocrom oxidase. Ti y cng xy ra chui oxy ha kh dn n bm ion H+
qua mng nh cytocrom b-c. Tuy nhin, chui oxy ha kh s kt thc bng phn ng kt hp in t vi
O2 v H+ to nn H2O. y, O2 ng vai tr l cht nhn in t cui cng. Trong trng hp thiu O2,
chui truyn in t v chu trnh Krebs s b c ch v t bo khng th thc hin h hp hiu kh.
Nh vy, kt qu ca s dn truyn in t l vic bm ion H+ qua mng v dn n tng hp ATP. Khi
cht cho in t v H+ l NADH2, mi phn t NADH2 s gip tng hp nn 3 ATP. Cn khi cht cho l
FADH2, lng ATP tng hp c ch l 2.
4.2.5. Tng kt nng lng khi oxy ha hon ton 1 phn t glucose
Hnh 2.14 l s biu din sn phm to thnh qua cc giai on phn gii mt phn t glucose ca h
hp hiu kh. V tng cng, lng ATP c to thnh l 40 phn t. Trong qu trnh ng phn s
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dng 2ATP hot ha. Ngoi ra, c ti liu cho rng, khi acid pyruvic c chuyn t t bo cht vo
cht nn ty th, qu trnh ny tiu tn thm 2 ATP na. V vy m s ATP thu c cn 36 phn t.
Ta bit rng, khi oxy ha hon ton 1 mol glucose s thu c 2870kJ, tng ng vi 686kcal (1cal =
4,184J). tng hp ATP t ADP v gc phosphat cn 8 10kcal/mol. Vy tnh v mt l thuyt, khi oxy
ha hon ton 1 mol glucose s thu c khong 76 mol ATP. Trong khi , thc t thu c 36 ATP. Nh
vy, hiu sut ca h hp hiu kh l gn 50%. Hn 50% nng lng cn li c gii phng ra di dng
nhit.
Hnh 2.14. Lng ATP sinh ra khi oxy ha hon ton mt phn t glucose trong h hp hiu kh
4.2.6. S iu ha h hp hiu kh trong t bo

Cng ging nh nhiu qu trnh khc, h hp chu s iu ha ca t bo sao cho lng ATP c
sn sinh ra mc va cho cc hot ng sng, khng tha m cng khng thiu. Theo Atkinson (1970),
nhn t iu ha qu trnh h hp l t l ca 3 thnh phn ATP, ADP v AMP trong t bo. ng a ra khi
nim v t s in tch nng lng, vit tt l EC (EC - energy charge). EC c tnh theo cng thc:
Khi t bo trng thi cn bng, ta c EC = 0,85. Vi EC < 0,85, ta c lng ATP sn sinh ra thp hn
lng ATP tiu th, t bo trng thi thiu nng lng. Cn ngc li, vi EC > 0,85, t bo tng hp ra
nhiu ATP hn l s dng. Do ATP c to nn t ADP v AMP nn nu coi tng 3 thnh phn ny l mt
s khng i, ta c th thy rng khi t bo hot ng mnh, lng ATP s b gim i trong khi ADP v
AMP tng ln. Ngc li, khi ATP c tng hp mnh, lng ADP v AMP li gim i. Trong mi trng
hp, nng ca ba thnh phn trn s lm thay i cng qu trnh h hp theo nhng chiu hng
khc nhau.
Nng ATP qu cao s c ch mt s enzym ca qu trnh ng phn v chu trnh Krebs, v d
enzym phospho fructo kinase xc tc cho phn ng bin i fructose-6-phosphat thnh fructose-1,6diphosphat. ATP lc ny ng vai tr l cht c ch khng cnh tranh v lm gim hot tnh ca cc enzym
m n tc ng. Kt qu ca s c ch l lm gim h hp v lng ATP c to thnh t i. S c ch do
sn phm cui ny c gi l c ch lin h ngc, mt trong nhng hnh thc iu ha ph bin ca t
bo. Trong trng hp lng ADP v AMP cao, chng li tr thnh cht hot ha lm tng cng qu trnh
h hp b vo lng ATP b thiu ht. Bn cnh , nng AMP cao cn kch thch s bin i
glycogen thnh glucose, ngun nguyn liu quan trng sn sinh ra ATP.
4.3. H hp k kh
y l hnh thc phn gii cc hp cht hu c sinh nng lng m khng c s tham gia ca oxy
khng kh. Trong qu trnh ny, cc hp cht hu c khng c oxy ha hon ton thnh CO2 v H2O nh
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h hp hiu kh m thay vo l cc sn phm hu c nh acid lactic, ethanol. V phng din hiu

qu sinh nng lng, lng ATP thu c trong h hp k kh km hn so vi h hp hiu kh. Bi vy, i
vi nhiu sinh vt hiu kh, h hp k kh ch l mt gii php tm thi trong nhng trng hp cp bch. V
d, cc t bo c, khi phi hot ng mnh m lng O2 khng , t bo s chuyn sang hnh thc h hp
k kh sn sinh nng lng. Lc ny, sn phm phn gii khng cn l CO2 v H2O na m thay vo
l acid L-lactic. Tuy nhin, trong t nhin c rt nhiu sinh vt sng trong cc iu kin rt ngho hoc
khng c O2 nh di vng nc su, trong bn hay trong lng t v.v... Hu ht chng l cc vi sinh vt c
cu to n bo. nhng sinh vt ny th h hp k kh l phng thc sn sinh nng lng duy nht.
H hp k kh c giai on u ging vi h hp hiu kh. Glucose s i qua chui phn ng ng
phn, hoc cc con ng phn gii khc ty loi, bin i thnh acid pyruvic. Tuy nhin, giai on sau,
acid pyruvic s khng i vo chu trnh Krebs m tham gia vo cc phn ng to sn phm hu c, v d
lactic, propionic, acetic, ethanol v.v... S to thnh ATP ch c giai on u ca h hp k kh vi s
lng t i. Nu tri qua ng phn, sn phm thu c ch c 2 ATP. C th thy rng, hiu sut sinh nng
lng ca h hp k kh rt thp.
T ngun carbon ban u l glucose, qua qu trnh h hp k kh, t bo sn sinh ra rt nhiu cc dng
hp cht hu c khc. iu ny c ngha v cng quan trng trong lnh vc sn xut. Thay v phi thc
hin cc phn ng tng hp phc tp, ngi ta c th li dng qu trnh phn gii k kh thu c cc sn
phm mong mun. Trong acid lactic v ethanol l hai v d ni bt. Trong sn xut, ngi ta gi hai qu
trnh ny l ln men lactic v ln men ethanol. Ln men ethanol c ngha trong cng nghip sn xut cn,
sn xut ru, bia v.v... Cn ln men lactic c s dng trong sn xut phomat, sa chua v qu trnh mui
chua ca nhiu loi thc phm. C hai qu trnh ln men u c chung cng on ng phn, bin i
glucose thnh acid pyruvic. Tip , acid pyruvic s c bin i tr thnh cht nhn in t t NADH,
qua gip quay vng NAD+. Tm tt qu trnh ln men ethanol v ln men lactic c th hin trn hnh
2.15.
Hnh 2.15. Qu trnh ln men ethanol v lactat
5. QUANG HP
5.1. i cng v quang hp
Tri t l ngi nh chung ca v vn sinh vt khc nhau. Cc sinh vt u c mi quan h mt thit
vi nhau v mt dinh dng, to nn mt mng li thc n khng l. Trong mng li ny, mi sinh vt
va l k sn mi, va l con mi. Thc n c phn gii khng ch cung cp cho sinh vt cc thnh phn
cu trc m cn l ngun cung cp nng lng vn hnh c my c th sinh vt. Nng lng y l
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nng lng t do c gii phng ra khi ph v cc lin kt ha hc.

Nu chng ta xp cc sinh vt theo mt chui m trong sinh vt ng sau s n sinh vt ng trc
th sinh vt no s ng u tin ? l cc sinh vt t dng. Chng thu nhn nng lng khng ch t
vic n cc sinh vt khc m t hai ngun. Th nht l nng lng ha hc ca cc hp cht v c. Cc sinh
vt s dng ngun nng lng ny l cc loi vi khun nh vi khun st, vi khun lu hunh v mt s loi
vi khun khc. Th nhng sinh khi ca cc loi vi khun ny qu nh v khng cung cp cho chui
thc n khng l. Thay vo y, m nhim vai tr ny l cc loi thc vt. Chng s dng ngun nng
lng th hai, l nng lng nh sng Mt Tri. Thc vt s dng nng lng nh sng Mt Tri to
nn sinh khi t cc hp cht v c n gin. Nh vy, c th ni rng, nh sng Mt Tri chnh l khi
ngun nng lng cho mi hot ng sng ca cc sinh vt trn Tri t. i vi con ngi, nh sng Mt
Tri cn l ngun gc ca cc loi nhin liu c s dng hng ngy nh du m, than ,... Ngi ta c
tnh rng, nh sng Mt Tri cung cp ti hn 90% ton b nng lng m con ngi s dng hin nay.
Qu trnh hp th nng lng t nh sng Mt Tri v tch ly chng trong cc hp cht hu c din ra
thc vt c gi l quang hp (photosynthesis). Vic nghin cu v qu trnh quang hp v b my quang
hp c mt ngha ht sc to ln. Trong sn xut, t nhng hiu bit cn k v qu trnh quang hp, trc
mt, ngi ta c th ti u ha lm tng nng sut cy trng. Xa hn na, iu g s xy ra nu nh chng ta
to ra c mt h thng quang hp nhn to ? Khi , chng ta s nm c trong tay mt xch c bn
duy tr s sng.
5.2. L - C quan lm nhim v quang hp thc vt

Qu trnh quang hp thc vt c thc hin ch yu l. L c nhng c im v hnh thi cng
nh cu to gii phu thch nghi vi chc nng ny.
V hnh thi: l thng c dng bn mng v nga b mt v pha c nh sng Mt Tri. Cc l trong
mt cnh c sp xp sao cho khng che khut ln nhau. Nhng c im ny cho php l tip nhn c
nh sng nhiu nht.
Hnh 2.16. M hnh cu trc ca l cy

A. Lp biu b; B. Lp t bo m giu; C. B mch; D. Lp t bo m xp; E. Kh khng.
V cu to gii phu: bao bc hai b mt ngoi ca l l mt lp biu b mng. Di biu b l lp t

bo m giu, ni din ra phn ln hot ng quang hp ca l. c im ca cc t bo m giu l xp st
nhau theo tng lp nhm hp th c nhiu nng lng nh sng. Pha di ca m giu l cc t bo m
xp. Cc t bo ca m ny xp tha tht c nhiu khong gian bo gip cho vic lu thng kh (CO2, O2)
v hi nc din ra d dng. Trong l c mt h thng cc b mch l ni dn nc v mui khong phc v
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cho quang hp v vn chuyn sn phm ca quang hp n cc b phn khc ca cy. Nm ri rc trong

lp biu b l cc kh khng vi vai tr nh cc cnh ca cho CO2, O2 v hi nc lu thng qua li gia cc
t bo bn trong l v mi trng bn ngoi. Cc kh khng ny c kh nng ng, m ty theo iu kin
mi trng.
cp c th thc vt, l l c quan lm nhim v quang hp. Cn cp t bo, l cc lc lp.
Trong bo quan ny cha mt h thng sinh hc cho php bin i nng lng t photon nh sng thnh
nng lng tch tr trong cc hp cht hu c.
5.3. B my quang hp
5.3.1. Cc sc t quang hp
Sc t l cc phn t hu c hp th nh sng, thnh phn u tin ca b my quang hp. Hin nay,
ngi ta xc nh c 4 nhm sc t chnh c mt trong l cy: chlorophyll, carotenoid, phycobilin v
sc t ca dch t bo (antoxyan).
5.3.1.1. Chlorophyll
Chlorophyll hay cn gi l nhm sc t lc l nhm sc t quan trng nht ca qu trnh quang hp. Cc
sc t thuc nhm chlorophyll c cu to chung gm 4 nhn pyron ni vi nhau bng cc cu ni metylen (CH=) to nn vng porphyrin. gia vng porphyrin l mt nhn Mg2+. Gn vi vng porphyrin cn c
ru phyton v vng cyclopentan. Cng thc tng qut v cng thc cu to ca hai loi chlorophyll chnh
c th hin hnh 2.17.
Quang ph hp th ca chlorophyll nm trong vng bc sng nh sng nhn thy c (400 - 700nm).
Chlorophyll c nh hp th hai bc sng:
Chlorophyll a l 420 v 662nm.
Chlorophyll b l 435 v 643nm.
Do nh hp th ca chlorophyll l vng quang ph nh sng xanh lam v nn vng quang ph cn
li khng c hp th to cho chlorophyll c mu xanh, mu c trng ca thc vt.
Chlorophyll chim khong 4% trng lng ca lc lp. Trong , ch yu l chlorophyll a, cn li l
chlorophyll b c hm lng bng khong 1/3 chlorophyll a. Chlorophyll a c mt trong mi c th quang
hp gii phng oxy.
Theo mt s nh khoa hc, cholorophyll c nhng c tnh chng vim, chng oxy ha, cha lnh vt
thng. Mt s kho st s b cn cho thy, chlorophyll cn gip lm gim cc cht gy ung th trong c
th.
Hnh 2.17. Cu to phn t chlorophyll
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5.3.1.2. Carotenoid
Carotenoid hay cn gi l sc t vng. Da theo cu trc ha hc caroten c chia lm hai nhm nh:
Caroten (C40H56): trong thc vt thng c 3 loi caroten , , . Ph hp th ca caroten nm trong
khong 446 - 476nm.
Xanthophyll (C40H56On, trong n = 1 - 6 ): xanthophyll l dn xut ca caroten v gm nhiu loi
khc nhau ty theo tr s n. Quang ph hp th ca xanthophyll nm trong khong 451 481nm.
Hnh 2.18. M hnh phn t -caroten
Carotenoid c nhiu trong hoa qu hoc c quan gi yu ca thc vt. Hin nay ngi ta bit n mt
s vai tr ca carotenoid trong thc vt nh sau:
Lc nh sng, bo v chlorophyll.
Tham gia vo qu trnh quang phn ly nc v thi O2 (xanthophyll).
Tham gia vo qu trnh quang hp thng qua vic hp th nng lng nh sng v truyn cho
chlorophyll. Carotenoid c mt nhiu trong quang h II.
Cng ging nh chlorophyll, carotenoid c c tnh chng oxy ha rt mnh. Ngoi ra, mt s cht
carotenoid khi hp th vo trong c th ngi cn c chuyn ha thnh vitamin A. Trong s 600 cht
carotenoid bit th khong 30 50 loi c kh nng ny. Carotenoid c ng dng cha cc bnh nh
ung th (phi, da, ng tiu ha,...), bnh c thy tinh th hay cc trng thi bnh l gy nn do tia phng
x hoc qu trnh oxy ha mnh v.v... Trong s cc cht thuc nhm carotenoid th -caroten v -caroten
c nghin cu nhiu nht.
5.3.1.3. Phycobilin
Phycobilin c nhiu trong to v cc thc vt bc thp sng di nc. y l mt loi sc t a nc,
c cu to t 4 vng pyron lin kt vi nhau bng cu metylen ( = CH - ) thnh mt mch khng khp kn
nh chlorophyll. Quang ph hp th ca phycobilin nm trong vng nh sng lc v vng. Cng ging nh
carotenoid, phycobilin sau khi hp th nng lng nh sng s truyn n cho chlorophyll, qua h tr cho
qu trnh quang hp.
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Hnh 2.19. Cng thc cu to ca phycobilin
5.3.1.4. Antoxyan
Antoxyan l nhm sc t cha nhiu trong dch t bo vi nhiu mu sc khc nhau nh , xanh, tm,...
Quang ph ca hp th antoxyan b sung cho quang ph hp th ca chlorophyll. Nhm sc t ny khng
tham gia vo qu trnh quang hp. Nng lng nh sng m cc sc t nhm antoxyan hp th s c
chuyn thnh dng nhit nng si m cho cy. Chnh bi vy, antoxyan c nhiu trong cc thc vt sng
vng c kh hu lnh.
S hp th
Hnh 2.20. Quang ph hp th nh sng ca cc sc t quang hp
5.3.2. Cc thnh phn ca chui truyn in t

5.3.2.1. Quinon
Cng thc chung:
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Nhm quinon
R1
R2
R3
Ubiquinon (Coenzym Q)
-OCH3
-OCH3
-CH3
6-10
Plastoquinon
-CH3
-CH3
Cc quinon c quang ph hp th vng t ngoi 260-300nm.

5.3.2.2. Cc cytocrom
Cytocrom (Cyt) c cu to gn ging chlorophyll nhng khc ch nhn Mg c thay th bng Fe2+
v khng c vng xiclopentan. Cc Cyt c quang ph hp th trong khong 500 - 600nm. Cc Cyt quan
trng trong b my quang hp l Cyt b (gm Cyt b6 v b3) v Cyt c (gm Cyt f). Th nng oxy ha ca Cyt f
l +0,36V, ca Cyt b6 l 0,06V. T l gia Cyt v chlorophyll trong b my quang hp vo khong 1/300 1/400.
5.3.2.3. Ferredoxin v ferredoxin-NADP-reductase
Ferredoxin (Fd) l cc protein c th ha tan trong nc, thnh phn cha Fe v nhm sulfit v c. T l
gia Fd v chlorophyll trong b my quang hp khong 1/400. Fd-NADP-reductase l mt flavoprotein c
quang ph hp th cc i 275, 385 v 456nm.
5.3.2.4. Plastocyanin
L mt loi protein c cha 2 nguyn t Cu, lin kt cht ch trong cu trc ca lc lp. Plastocyanin
khi trng thi oxy ha c mu xanh tm, ph hp th cc i l 597nm, dng kh khng c mu. T l gia
plastocyanin v chlorophyll l 1/400.
5.3.3. n v quang hp
Cc sc t ch c th thc hin c cc phn ng quang hp hon chnh khi chng nm trong t bo,
ni cch khc l trong s lin kt cht ch vi cc thnh phn khc ca b my quang hp. n v quang hp
c th hiu l cu trc nh nht c kh nng thc hin bin i nng lng t photon nh sng thnh nng
lng trong cc lin kt ha hc. Mt cch chi tit, mt h thng c gi l n v quang hp khi n thc
hin c 3 chc nng:
Tip nhn mt phn t CO2 (hay gii phng mt phn t O2);
Chuyn mt in t;
Bin i nng lng mt quang t thnh dng nng lng ha hc.
Vy mt n v quang hp c t chc ra sao v s lng tng thnh phn nh th no ? Ngi ta thy
rng, trong mt n v quang hp lun c hai thnh phn chnh l phc h sc t v chui truyn in t. T
nm 1932, Emerson v Arnold khi lm th nghim vi to Chlorella thy rng, trong iu kin ti u,
c nh mt phn t CO2 cn c s tham gia ca 2500 phn t chlorophyll. Cc nghin cu v sau ca cc
tc gi khc cng cho kt qu tng t, s lng phn t chlorophyll cn thit l khong 2500 4200 phn
t. Da vo kt qu nghin cu ca Emerson, Arnold cng cc tc gi khc, nm 1969, Avron ngh
rng mt n v quang hp s gm 2400 phn t chlorophyll c sp xp theo 4 tiu n v nh. Mi n
v nh gm 600 phn t chlorophyll (trong 300 thuc h quang ha I, 300 thuc h quang ha II) v 1
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phn t P700.
5.4. Qu trnh quang hp

Quang hp l mt qu trnh phc tp. Trong bao gm c cc phn ng cn s c mt ca nh sng
ln phn ng enzym thng thng. Hin nay, ngi ta chia quang hp ra thnh hai giai on:
Pha sng: cn s tham gia ca nh sng, thc hin bin i nng lng cha trong photon nh sng
thnh dng nng lng ha hc cha trong ATP v NADPH2.
Pha ti: khng cn s tham gia trc tip ca nh sng, pha ny s dng ATP v NADPH2 sn sinh ra
t pha sng thc hin cc phn ng c nh CO2 thnh sn phm hu c.
5.4.1. Pha sng
Cc phn ng sng xy ra trn mng c gi l mng quang hp. vi khun quang hp, l mng t
bo. Cn t bo thc vt, l mng thylacoit. H thng thc hin pha sng chnh l cc n v quang hp.
C th chia cc phn ng sng thnh ba giai on:
H sc t hp th photon ng sng v lm bt ra cc in t trng thi kch thch.
in t t h sc t s i qua chui truyn in t, hot ha cc bm H+ to nn s chnh lch nng
H+ gia hai bn mng quang hp. Sau khi kt thc chui, in t s c quay vng hoc c tip nhn
bi NADP+ to thnh NADPH2.
S chnh lch nng H+ s dn n tng hp ATP theo phng thc ha thm.
5.4.1.1. S hp th nh sng
Cc sc t trong n v quang hp c t chc thnh cm gi l phc h anten. Phc h ny gm cc
sc t nm xung quanh mt sc t trung tm l mt phn t chlorophyll c bit. Ton b nng lng photon
do phn t sc t bn ngoi hp th s c tp trung li cho sc t trung tm.
Khi cc phn t sc t hp th nng lng t photon nh sng, in t ca sc t s chuyn t trng thi
nng lng c bn sang trng thi nng lng kch thch. Tuy nhin, trng thi ny khng bn vng, in t
s nhanh chng tr v trng thi ban u v gii phng ra nng lng d tha. Nng lng ny s c phn
t sc t bn cnh hp th v khin chng chuyn sang trng thi kch thch. Ngi ta gi y l phng
thc truyn nng lng theo kiu cng hng. Qu trnh dn truyn nh vy din ra lin tc cho ti khi nng
lng c chuyn cho phn t sc t trung tm.
Ngi ta thy rng, trong thc vt tn ti hai h quang ha, c gi l h quang ha I v II. Mi h
quang ha c mt trung tm phn ng v bc sng hp th hiu qu ring. H quang ha I, vi trung tm
phn ng P700, hp th hiu qu nh sng c bc sng di khong 700nm. Cn h quang ha II, vi trung
tm phn ng P680, li hp th hiu qu nh sng c bc sng ngn hn, khong 680nm.
Hai h quang ha I v II khng hot ng c lp m c s kt hp vi nhau. Nm 1940, Emerson tin
hnh th nghim o cng quang hp ca to Chlorella khi chiu nh sng c bc sng 650 v 680nm.
ng thy rng, cng quang hp khi chiu hn hp c hai nh sng ln hn so vi tng cng
quang hp khi chiu hai nh sng n sc cng li. Nh vy, hai h quang ha kt hp vi nhau lm tng
hiu qu quang hp. Ngi ta gi hin tng ny l hiu ng tng cng Emerson. Quan nim chung cho
rng, s tn ti v kt hp gia hai h quang ha I v II c to v tt c cc thc vt bc cao.
gii thch s kt hp gia hai h quang ha, vo nm 1960, Hill v Bendal xut mt m hnh
c gi l s Z. Theo s ny, hai h quang ha khng hot ng song song m c ni vi nhau
thng qua chui truyn in t. Cho n nay, s Z vn l m hnh c tha nhn rng ri, c nghin
cu ngy cng su sc v hon thin.
5.4.1.2. S dn truyn in t
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Sau khi ton b nng lng do phc h anten hp th c tp trung li trung tm phn ng, ti y,
in t ca trung tm phn ng s t mc nng lng cao v bt ra ngoi. in t ny s c tip nhn v
i qua mt h thng cc cht dn truyn in t. Kt qu ca s dn truyn ny l nng lng trong in t
s c chit rt mt cch t t v c s dng hot ha cc bm H+ nm trn mng quang hp. Qu
trnh dn truyn in t trong quang hp thc vt din ra theo hai phng thc l dn truyn theo vng v
dn truyn khng theo vng.
Dn truyn in t theo vng
Vn chuyn in t theo vng c thc hin bi h quang ha I vi trung tm phn ng P700 v bc
sng hp th cc i = 680 - 700nm. Thnh phn ca chu trnh bao gm ferredoxin (Fd); cytocrom b6-f
(Cyt). Qu trnh vn chuyn c th hin hnh 2.21.
Hnh 2.21. Qu trnh dn truyn in t theo vng
Sau khi tip nhn nng lng, in t ca trung tm phn ng P700 s chuyn sang trng thi kch thch
v bt ra ngoi. Chng c tip nhn bi Fd ri i qua Cyt v cui cng quay tr li P700. Nh vy, con
ng di chuyn in t l mt chu trnh kp kn.
im mu cht ca chu trnh nm s bin i mc nng lng ca in t khi i qua cc cht. T mc
nng lng c bn, photon nh sng s y in t ca trung tm P700 ln mc nng lng kch thch. Nng
lng ny s gim dn khi in t ln lt i qua Fd, Cyt v tr v trng thi c bn khi quay tr li trung
tm P700. Ti phc h cytocrom b6-f, nng lng chit rt t in t cao nng s thc y qu trnh bm ion
H+ vo bn trong ti mng thylacoit. Khi nng trong ti thylacoit tng cao, H+ s khuch tn ra cht nn
lc lp qua cc phc h ATP synthetase v dn n tng hp ATP. Qu trnh tng hp ATP nh vy gi l
quang phosphorin ha.
Qu trnh phosphoryl ha quang ha lc lp v phosphoryl ha oxy ha ty th ging nhau v c ch
hnh thnh ATP t ADP v phosphat v c. Chng ch khc nhau l phosphoryl ha quang ha thc hin
c nh nng lng photon nh sng, cn phosphoryl ha oxy ha thc hin c nh nng lng gii
phng ra t qu trnh oxy ha cc hp cht hu c. Mt s cng trnh nghin cu cho rng c mt photon
c hp th c kh nng hnh thnh c t 1 n 3 ATP v c th c hai im to ra ATP trong chui
truyn in t. l im t Fd n Cytb6 v im t Cyt b6 n Cyt f. Ti hai im ny, s chnh lch th
nng oxy ha kh gia cc cht truyn e trung gian l kh ln (0,4 v 0,39V).
Nh vy, t nguyn liu u vo l cc photon nh sng, thng qua dn truyn in t theo vng, ta c
sn phm u ra l ATP.
Dn truyn in t khng theo vng
Vn chuyn in t khng theo vng c thc hin bi c hai h quang ha I v II. Thnh phn ca
chui truyn in t gm: quinon (Q); plastoquinon (PQ); plastocyanin (PC); cytocrom b6-f (Cyt). Qu trnh
vn chuyn in t khng theo vng c khi u t trung tm phn ng P680. in t sau khi ra khi
P680 s ln lt i qua Q PQ Cyt PC P700. Ti P700, in t sau khi bt ra s n Fd ging vn
chuyn theo vng. Th nhng im khc bit ch, in t s khng ti Cyt m c dng sn sinh
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NADPH2 nh phc h enzym Fd-reductase. P680 sau khi b mt in t s c b sung t qu trnh

quang phn ly nc:
H2O 2H+ + 2e- + (1/2) O2
Bng phng php ng v phng x (O18), ngy nay ngi ta xc nh c rng O2 gii phng ra
trong qu trnh quang hp cy xanh l O2 t H2O ch khng phi t CO2 nh trc y quan nim. C ch
ca qu trnh quang phn ly nc nh tc dng ca nh sng c hp th bi chlorophyll c biu din
nh sau:
Chlorophyll hp th 4 photon nh sng tr thnh trng thi kch thch:
4Chl
4h
4Chl*
Chlorophyll trng thi kch thch s tham gia vo quang phn ly nc:
4ChlH +
4Chl* + 4H2O
4OH 2H2O +
O2
C th tm tt qu trnh nh sau:
4OH-
Hnh 2.22. Qu trnh dn truyn in t khng theo

vng
Dn truyn in t khng vng chnh l s Z m Hill v Bendal xut gii thch s kt hp

ca hai h quang ha. Trong s ny, in t cao nng sau khi bt ra t h quang ha II (P680) s i qua
chui truyn in t v c s dng hot ha cc bm H+ dn n tng hp ATP theo quang phosphoryl
ha. Sau khi chit rt ht nng lng, in t ny c nhp vo h quang ha I (P700) v tip tc c hot
ha ln hai. Lc ny, in t cao nng s c s dng tng hp NADPH2. Nh vy, qua hai ln hot ha
nh nng lng ca photon nh sng, s vn ng ca in t cao nng trong s Z s gip to ra hai sn
phm l ATP v NADPH2.
Trong hai con ng, dn truyn in t khng theo vng l c ch hnh thnh nng lng quan trng
thc vt. Ni cch khc, con ng ny tin ha hn dn truyn in t theo vng v ch c th thy thc
vt bc cao. Tuy nhin, nu ch din ra dn truyn in t khng vng, cy xanh s thiu ht ATP v qu
trnh hnh thnh glucose b nh hng, sn phm ch yu ca quang hp lc ny s l cc acid hu c, acid
bo. Bi vy, thng thng hai con ng dn truyn in t theo vng v khng theo vng din ra trong s
phi hp nhp nhng vi nhau.
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Hnh 2.23. M hnh chui truyn in t trong quang hp
5.4.2. Pha ti
Cc sn phm ca pha sng, bao gm ATP v NADPH2, s c pha ti s dng c nh CO2 thnh
cc phn t hu c. l cht dinh dng v nguyn liu xy dng cc cu trc t bo.
Vo nm 1946, bng vic s dng ng v phng x C14, Melvin Calvin v cng s chng minh mt
cch ton din v c ch ca phn ng ti - con ng carbon trong quang hp. Vi cng trnh nghin cu
c tnh t ph ny, nm 1961, Calvin c nhn gii thng Nobel ha hc v chu trnh carbon trong
pha ti v sau c mang tn ng - chu trnh Calvin.
Cc nghin cu cho thy rng, pha ti phn ln cc loi thc vt bc cao v cc vi khun quang hp
u din ra theo chu trnh Calvin (cn gi l chu trnh C3). Ngoi ra, mt s thc vt sng trong cc iu
kin c bit cn c thm chu trnh C4 v chu trnh CAM.
5.4.2.1. Chu trnh C3 (chu trnh Calvin)
Chu trnh C3 din ra ti cht nn ca lc lp. N bao gm mt lot cc phn ng enzym ni thnh vng
khp kn, nguyn liu u vo l CO2, ATP, NADPH2; cn sn phm u ra l hydratcarbon. C th chia
chu trnh C3 thnh hai giai on:
C nh carbon dn n tng hp ng.
Ti sinh cht nhn.
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Hnh 2.24. Chu trnh Calvin
C nh carbon dn n tng hp ng
Khi u ca chu trnh l vic CO2 kt hp vi mt phn t ng 5C l ribulose biphosphat (RuBP)
to ra sn phm l 2 phn t acid c 3C l acid phosphoglyceric (AGP). Phn ng ny c xc tc bi
enzym ribulose biphosphat carboxylase.
Sau khi to ra AGP, t bo s s dng ATP bin chng thnh 1,3 biphospho glycerat. Tip , 1,3
biphospho glycerat s tc dng vi NADPH2 tr thnh glyceraldehyd 3 phosphat (G3P).
G3P l mt sn phm trung gian quan trng ca chu trnh C3 v t c th tng hp glucose theo con
ng ngc vi chui ng phn trong h hp. Ngoi ra, t bo c th chuyn ha G3P thnh cc cht bo
hay acid amin hoc phn gii to thnh acid pyruvic v s dng cho h hp sinh nng lng. T G3P, t
bo to ra sn phm no ph thuc nhiu vo iu kin nh sng, nng CO2 v O2.
Ti sinh cht nhn
Mc d to ra G3P l im quan trng tng hp cc hydratcarbon song trong chu trnh C3, phn ln
G3P li c s dng cho vic ti sinh cht nhn CO2 l RuBP. y l mt chui phn ng enzym phc tp
v cn s dng nng lng cung cp t ATP.
im b v h hp sng
Xt v nhiu mt, quang hp v h hp l hai qu trnh ngc nhau. Trong cng mt t bo thc vt, khi
lc lp ang thc hin quang hp, ly CO2 vo v nh O2 ra th ti ty th ng thi din ra s hp th O2 v
gii phng CO2. Khi tng dn cng quang hp s dn n mt thi im thc vt khng c s trao i
kh gia t bo v mi trng na. Lng CO2 v O2 ca qu trnh quang hp v h hp lc ny cn bng
nhau. Ngi ta gi y l im b. i vi nhiu thc vt, im b thy cng nh sng khong 2%
nh sng Mt Tri ton phn. im b thay i tu theo loi v chu nh hng bi nhit v nng CO2
trong mi trng. Qu trnh quang hp ch xy ra hiu qu khi cng nh sng trn im b.
Trong trng hp cng nh sng nm di im b, thc vt s xy ra mt hin tng c gi l
h hp sng. Enzym u tin ca chu trnh Calvin, enzym ribulose biphosphat carboxylase ngoi kh nng
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xc tc cho phn ng gn CO2 vo RuBP to ra AGP, n cn c kh nng xc tc cho phn ng gn

O2 vi RuBP to ra phosphoglycolat, mt sn phm khng c ch li cho thc vt. RuBP s tham gia phn
ng vi CO2 hay O2 ph thuc rt nhiu vo nng ca hai thnh phn ny trong mi trng. Khi quang
hp din ra mnh, lng O2 sn sinh ra nhiu trong khi CO2 trong mi trng khng cung cp, O2 s
cnh tranh vi CO2 kt hp vi RuBP v gy nn hin tng h hp sng. Ngi ta tnh rng, h hp sng
c th lm gim nng sut ca quang hp n 30-40%. Bi vy, khi sng trong nhng iu kin c bit,
thc vt phi c nhng c ch thch nghi qu trnh quang hp din ra mt cch bnh thng.
Hnh 2.25. Sn phm to thnh khi quang hp din ra trong cc iu kin nh sng
v nng O2, CO2 khc nhau
5.4.2.2. Chu trnh C4 (chu trnh Hatch & Slack)

T khi pht hin ra chu trnh Calvin, ngi ta coi y l chu trnh c nht v con ng carbon trong
quang hp thc vt. Tuy nhin, n nm 1965, Hatch v Slack l hai nh khoa hc ngi Australia nhn
thy rng, mt s loi thc vt nhit i nh ma, ng, c lng vc, c gu, rau dn v.v..., con ng c
nh CO2 khng din ra theo chu trnh Calvin m theo mt con ng khc. trong lc lp ca nhng thc
vt ny, enzym ribulose biphosphat carboxylase (enzym ca phn ng u tin trong chu trnh Calvin) hot
ng rt yu hoc khng hot ng. Thay vo l mt enzym khc c tn l phosphoenolpyruvat
carboxylase hot ng rt mnh. V vy, sn phm u tin ca chu trnh ny khng phi l AGP m l cc
acid oxaloacetic, malic, aspartic. Cu to phn t ca cc acid ny gm 4 nguyn t carbon, v vy chu trnh
nhng thc vt ny c gi l chu trnh C4.
Qu trnh c nh CO2 thc vt C4 din ra theo hai giai on hai phn khc nhau trong l. u tin,
CO2 t kh quyn i qua l kh vo t bo cht ca t bo tht l v phn ng vi PEP (phosphoenolpyruvat)
to thnh oxaloacetat. Phn ng din ra vi s xc tc ca enzym PEP carboxylase. Do nng PEP
trong t bo tht l kh cao nn s to thnh oxaloacetat c th din ra iu kin nng CO2 thp. Tip
, nh c NADPH2, oxaloacetat s c kh thnh malat. giai on th hai, malat c chuyn t t bo
tht l vo t bo bao b mch. Ti y xy ra qu trnh decarboxyl ha bin malat thnh pyruvat v gii
phng ra CO2. Lc ny, CO2 s c c nh thnh cc hydratcarbon thng qua chu trnh Calvin
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Hnh 2.26. Chu trnh C4
Nh vy, c th thy rng, qu trnh c nh CO2 ca thc vt C3 v C4 khc nhau ch thc vt C4 c

thm chu trnh tch ly CO2 di dng malat. c im ny gip cho thc vt C4 c th sinh trng trong
mi trng kh mn, ni m thc vt C3 gp rt nhiu kh khn. So vi thc vt C3, thc vt C4 cn thm
mt s ATP vn ng chu trnh tch ly CO2. Tuy nhin, b li, chng li c cng quang hp cao.
Qu trnh ng ha CO2 khng b bo ha mc d cng chiu sng mnh.
5.4.2.3. Chu trnh CAM (Crassulacean acid metabolism)
Chu trnh CAM tn ti cc thc vt thch nghi vi mi trng sng kh hn ko di, v d da, xng
rng,... Nhng thc vt ny thng mng nc v c th c xu hng cu to tip xc vi mi trng
mt b mt nh nht. Cc l kh trn c th thng ng vo ban ngy v m vo ban m. Cc c im
nh vy s gip thc vt CAM gim thiu s mt nc. Song ng thi chng cng lm gim c s trao i
kh gia thc vt v mi trng. gii quyt mu thun ny, ngoi chu trnh Calvin, thc vt CAM cn
c thm mt chu trnh tch ly CO2 ging nh thc vt C4. Tuy nhin, khc ch, nu nh thc vt C4,
con ng c nh CO2 c phn bit v mt khng gian th thc vt CAM li phn bit v mt thi gian.
Chu trnh tch ly CO2 di dng malat xy ra vo ban m, khi cc l kh m, cn qu trnh tng hp
ng (chu trnh Calvin) li xy ra vo ban ngy.
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Hnh 2.27. Chu trnh CAM: A. Ban m, B. Ban ngy.
CU HI LNG GI
1. G l g ? G c nh hng nh th no n din bin phn ng ha hc ?
2. Nng lng hot ha c nh hng nh th no n tc phn ng ?
3. ATP c vai tr nh th no trong t bo ? Trnh by m hnh phn t v cc phng phc tng hp
ATP trong t bo.
4. Enzym l g ? Trnh by c im cu to ca enzym.
5. Trnh by c ch hot ng ca enzym. Gii thch tnh c hiu ca enzym.
6. Trnh by nh hng ca cc nhn t nhit , pH, nng enzym/c cht, cc cht c ch ln tc
phn ng do enzym xc tc.
7. Trnh by khi nim v h hp. Nhng im khc nhau c bn gia h hp hiu kh v k kh ?
8. Trnh by cc qu trnh h hp hiu kh.
9. Trnh by khi nim v quang hp. Mi quan h gia pha sng v pha ti ?
10. Trnh by cc din bin ca pha sng.
11. Trnh by cc giai on chnh ca pha ti. Qu trnh c nh CO2 thc vt C3, C4 v CAM khc
nhau nh th no ?
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Chng 3
DI TRUYN V BIN D
MC TIU
1.
2.
3.
4.
5.
6.
7.
8.
Trnh by c cu trc ca cht liu di truyn.

M t c cu trc nhim sc th Eukaryota v Prokaryota.
Trnh by c sao chp ADN Eukaryota v Prokaryota.
Trnh by c chu trnh t bo.
Trnh by c cc kiu phn bo.
Trnh by c di truyn nhim sc th v di truyn ngoi nhim sc th.
Trnh by c bin d.
Trnh by c cc kiu t bin.
1. C S PHN T CA CHT LIU DI TRUYN

1.1. Acid nucleic - vt liu di truyn
Vt liu di truyn ng vai tr ht sc quan trng trong sinh gii vi cc c tnh c bit m thin
nhin ban tng nh:
Hm cha cc thng tin cn thit i vi cu to, hot ng, sinh sn ca t bo trong dng bn vng
lu di.
T sao chp c mt cch chnh xc thng tin di truyn ca th h sau ging nh ca th h trc.
Thng tin trong cht liu di truyn c s dng to ra cc hp cht cn cho cu trc v hot ng
ca t bo.
Cht liu di truyn c kh nng bin i c.
Trong s cc polyme sinh hc hin hu ch c cc acid deoxyribonucleic (ADN) v acid ribonucleic
(ARN) c c tnh ph hp vi cc c tnh trn.
1.2. Cc th nghim minh chng acid nucleic l cht liu di truyn
C rt nhiu bng chng, th nghim chng minh chc chn rng cc acid nucleic l cht liu di truyn.
Di y l mt s v d:
Acid nucleic hp ph cc i nh sng t ngoi bc sng 260nm, v y l bc sng m nh sng
t ngoi c th gy t bin ti a cho cc t bo. Ngoi ra, acid nucleic cn hp ph cc i nh sng t
ngoi ca protein bc sng 280nm.
Nm 1928, F. Griffith pht hin thy ni S ca Diplococcus pneumoniae (khun lc nhn do c v bc
bn ngoi) lm cht chut khi em tim vo chut. Trong khi , ni R (khun lc rp do khng c v bc
bn ngoi) li khng gy hi g i vi chut c tim ni ny vo c th. Tuy nhin, khi tim hn hp cc
vi khun R cn sng vi cc vi khun S b cht do x l nhit vo chut th chut b cht v t mu ca
cc con chut b cht y ngi ta phn lp c cc chng S sng. Nh vy, c tc nhn nht nh (tc
nhn bin np) t vi khun S cht bin np vo vi khun R bin i vi khun R thnh vi khun S. Qu
file://C:\Windows\Temp\bhwcmokvfv\Chapter3.htm
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trnh ny gi l qu trnh bin np.

Nm 1944, O.T. Avery, C.M Macleod v M. McCarty chng minh c tc nhn bin np l ADN v
hin tng bin np vi khun R thnh vi khun S nu ch mt i khi x l tc nhn bin np tch ra c
t vi khun S vi deoxyribonuclease - enzym phn hu ADN.
Nm 1957, H. Fraenkel-Conrat v B. Singer cng b th nghim lp rp virus khm thuc l TMV
(tobaco mosaic virus) l virus cha li ARN v v protein. Virus ny c hai dng A v B. Cc nh khoa hc
lp c li ARN ca dng ny vi v protein ca dng kia v ngc li, to nn cc virus c v v li
thuc hai dng khc nhau, ri em nhim tng loi virus vo thuc l gy m khm. Kt qu phn lp
sau th nghim cho thy, tt c cc th h virus con phn lp c t cc m khm u mang cng mt
dng ca li ARN v v protein - dng ca ARN em nhim ch khng phi dng ca v protein. Nh vy,
thng tin di truyn trong TMV c cha trong ARN ch khng phi trong protein.
Ngy nay, chng ta bit rng phn ln sinh vt cht liu di truyn l ADN v mt s virus l ARN.
1.3. Thnh phn v cu trc ca acid nucleic

Cc acid nucleotid l cc acid 2-deoxyribonucleic (ADN) v acid ribonucleic (ARN). y l cc
polyme mch di c to bi cc monome l cc nucleotid. Mi nucleotid gm ba thnh phn l cc base
dn xut purin hoc pyrimidin, ng 2-deoxyribose (trong ADN) hoc ribose (trong ARN) v gc
phosphat. Cc dn xut base purin l adenin (A) v guanin (G) i vi c hai ADN v ARN, cn cc dn
xut pyrimidin bao gm thymin (T) v cytosin (C) l cc thnh phn ca ADN, trong khi ARN cha cytosin
(C) v uracil (U).
Uracil
Thymin
Cytosin
Base purin hoc pyrimidin gn vi ng ribose to thnh nucleozid. Nucleozid gn vi gc phosphat

to ra nucleotid (3- v 5- nucleotid) v polyme ho cc nucleotid to ra mch polynucleotid ca cc acid
nucleic.
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Phn t polynucleotid mang tnh phn cc: ng pentose mt u mang nhm phosphoryl hoc
hydroxyl v tr 5' (u 5'), cn u kia mang nhm hydroxyl v tr 3' (u 3').
Tuy nhin, cc ADN trong t bo sinh vt qua lin kt cc base vi nhau m to thnh mch p - chui
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xon kp. Hnh 3.1 gii thiu m hnh chui xon kp ADN ca Watson - Crick. Trong cu hnh chui
xon kp ca ADN mch khung polyphosphat-2'-deoxyribose chy bao bn ngoi, cc base purin v
pyrimidin chy bn trong v bt cp (sng i) vi nhau qua cc cu lin kt hydro to ra s gn kt tng
hp ca mch kp ADN (hnh 3.2). Ch cc cp base tng hp A - T v G - C mi c th to ra c cc
cu lin kt bn vng trong cu trc chui xon kp. Mi vng xon cha 10 cp base, c chiu di 3,4nm;
mi cp base cch nhau l 0,34nm. Nh tnh cht bt cp tng hp ny m kt qu l hai si n trong
chui xon kp c trnh t tng hp vi nhau. Tnh tng hp c th ny trong lin kt cp base c tm
quan trng c bit i vi chc nng ca ADN nh sao chp, phin m v dch m. c tnh quan trng
khc ca chui xon kp l hai si n ca chui c tnh phn cc ngc chiu nhau: u 3' ca si ny nm
cng pha vi u 5' ca si kia v ngc li (hnh 3.3).
Hnh 3.1. M hnh ADN ca Watson-Crick

a) M hnh phn t chui xon kp ADN; b) S chui xon kp ADN; c) S ct ngang mt si ca chui nhn dc theo
trung tm: khung polyme ng phosphat nm pha ngoi, cc base nm pha trong.
Trong ADN, t l (A + G)/(C + T) = 1. Tuy nhin, t s base (G + C)/(A + T + G + C) c gi l t s
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phn trm; cc base G + C thng khc nhau cc sinh vt khc nhau, nhng ging nhau mi loi.
Hnh 3.2. Sng i gia cc cp base trong ADN

a) Thymin v adenin; b) Cytosin v guanin bng cc cu hydro
Hnh 3.3. Cu trc chi tit mt phn on ADN

ADN cn c nhiu dng cu to khc nhau. Dng chui xon kp theo m hnh Watson Crick l ADN
dng B, dng ph bin ca ADN - mi vng xon cha 10 cp base thng gc vi trc ca chui xon. Bn
cnh cn cn k n cc dng A, dng C, dng D v dng Z. dng A, cc cp base nm bn ngoi v
nghing gc 19o so vi trc ca chui, c 11 cp trong mi vng xon. dng C, cc cp base nm tp trung
bn trong chui xon, nhng nghing gc vi trc chui xon ging dng A, tuy nhin vi gc b hn, mi
vng xon c 7,9-9,6 cp base. ADN dng D c thit din l hnh bt gic, cc cp base nm bn trong chui
xon v cng nghing gc vi trc xon, mi vng xon c 8 cp base. ADN dng Z (dng zigzag) c khung
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xon tri, mi vng xon cha 12 cp base nm bn ngoi chui xon. Dng Z ny tm thy mt s
sinh vt nhn chun hoc mt s on ADN m gen chu s iu ho (hnh 3.4).
Hnh 3.4. Cc dng khc nhau ca chui xon kp ADN
1.4. c im vt liu di truyn Prokaryota v Eukaryota

Vt liu di truyn sinh vt Prokaryota l ADN chui xon kp, mi gen c gn trn mt on ca
ADN v tt c cc phn on trn ADN ca cc Prokaryota u c cha ngha di truyn nht nh. Mch
ADN c s dng ti a chuyn ti thng tin di truyn.
Ngc li, ADN ca cc Eukaryota, nht l ca cc sinh vt bc cao cha lun phin cc phn on
exon v intron, trong , cc phn on exon m ho cc tnh trng di truyn ca c th, cn cc phn on
intron l cc phn on khng m ho (non-sense). Ngy nay, vai tr ca cc intron vn cn l b n i vi
cc nh nghin cu, c th vai tr ca cc intron ny nm trong vic n nh cc gen ca ADN.
2. CU TRC NHIM SC TH PROKARYOTA V EUKARYOTA

n nay, cc nghin cho thy nhim sc th Prokaryota v Eukaryota l ADN mch kp. Cc virus c
b gen l ADN hoc ARN mch n hay mch kp. B gen ca cc sinh vt khc nhau l rt khc nhau
(bng 3.1).
Bng 3.1. Chiu di b gen cc sinh vt
Sinh vt
Chiu di b gen n bi (s cp base)
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Virus
E. coli
Nm men
Drosophila
ng vt c xng sng
Ngi
Thc vt
103 - 105
4,5.106
5. 107
1,5.108
108 - 1010
3.109
1010 - 1011
2.1. Hnh thi nhim sc th

Khi nhum t bo ang phn chia bng mt s cht mu base, c th nhn thy di knh hin vi quang
hc cc cu trc hnh que nhum mu m, nn c gi l nhim sc th (chromosome), c ngha l th
nhum mu. Mi nhim sc th c hnh dng c trng, r nht k gia (metaphase) ca nguyn phn.
Tm ng (centromere) l im tht eo chia nhim sc th thnh hai vai vi chiu di khc nhau. Theo quy
c chung, vai ngn hn gi l vai p v vai di hn gi vai q. Da vo v tr tm ng c th phn bit hnh
thi cc nhim sc th: tm gia (metacentric) khi hai vai bng nhau, tm u (acrocentric) khi hai vai khng
bng nhau v tm mt (telocentric) khi tm ng nm gn cui (hnh 3.5). Trn nhim sc th c th thy cc
vt m hn, gi l vt nhim sc (chromomere).
cc t bo dinh dng (somatic cell t bo thn th), mi nhim sc th c mt cp ging nhau v
hnh thi, gi l cc nhim sc th tng ng hay ng ng (homologous). B nhim sc th c cp gi l
lng bi (2n) v b nhim sc th l n bi (n) khi mi nhim sc th ch c mt bn, khng c cp.
Ngoi ra, nhiu ng vt c s khc nhau gia th c v ci mt cp nhim sc th gii tnh (sexual
chromosme).
Hnh 3.5. S cc nhim sc th k gia v k sau
T bo mt s m c cc nhim sc th khng l nh tuyn nc bt ca rui gim (hnh 3.6) v
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nhim sc th chi n (lampbrush chromosome) mt s t bo trng ca mt loi lng c (hnh 3.7).
Hnh 3.6. Nhim sc th khng l tuyn nc bt ca Drosophila

a) Nguyn dng; b) Phng to ch mi tn ca a; c) Phng to mt on ca b.
Hnh 3.7. Nhim sc th chi n Triturus viridescens

a) Nguyn dng; b) Phng to mt on ca a; c) Phng to mt on ca b; d) Phng to mt on ca c.
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Tt c t bo ca mt loi ni chung c s lng nhim sc th c trng cho loi . V d, t bo rui

gim Drosophila melanogaster c 8 nhim sc th, t bo ng c 20 nhim sc th, t bo ngi c 46. Mi
loi nhim sc th c hnh dng c trng. S m t hnh thi ca nhim sc th c gi l kiu nhn
(caryotype) c trng cho mi loi. Kiu nhn c th c trnh din di dng nhim sc khi cc nhim
sc th c sp xp theo th t bt u t di nht n ngn nht.
2.2. Cu trc nhim sc th Prokaryota

ADN ca E. coli nu ko di thng ra s di 1,35mm, gp 1.000 ln chiu di t bo ca chnh E. coli.
V th, trong t bo ADN phi c xon cht v gi gn rt tinh vi sao cho gen vn biu hin c khi c
nhu cu.
2.2.1. Dng siu xon t nhin ca ADN
Genom ca E. coli cng nh ca phn ln cc sinh vt tin nhn l phn t ADN dng vng, khng gn
vi protein to phc hp nh nhim sc th ca Eukaryota. Nhim sc th c hiu l si ADN, c
dng cho c cc vi khun. ADN c th trong 3 dng cu trc t-p (topological structure):
Hnh 3.8. Cc dng thng, vng trn v siu xon ca ADN

a) Ly tm siu tc; b) in di trn agarose gel; c) Nhn di knh hin vi in t.
Dng siu xon, khi chui kp vn xon hnh s 8. y l dng t nhin (native) trong t bo vi khun
ca ADN. Dng siu xon ca ADN l dng ph bin trong t bo Prokaryota.
Dng vng trn, khi chui ADN cng trn. Dng ny xut hin khi ADN siu xon b ct t mt
trong hai mch kp.
Dng thng, khi ADN b t c hai si (hnh 3.8).
2.2.2. M hnh genom E. coli
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Mt s m hnh c xy dng m t cc dng bin i ca genom E. coli trong t bo. Hnh 3.9
trnh by mt s dng trong s .
Hnh 3.9. Genom E. coli thay i hnh dng trong t bo (Pettijohn v Hecht, 1974)
C th nhn thy s co rt kch thc ADN mnh (t ng knh 350m xung cn 2m), s cun li
c thc hin nh cc ARN ni (linker). Khi cc ARN linker b ct, mch ADN bung di ra thun tin cho
sao chp ADN. Nu ADNase ct mi si, chui ADN tho xon cng ra tng hp protein.
2.3. Cu trc nhim sc th Eukaryota

2.3.1. Kiu nhn
a s ADN ca Eukaryota c t chc thnh nhiu nhim sc th trong nhn t bo. Mi nhim sc
th cha 1 phn t ADN mch thng xon kp. T chc genom nh vy gi l kiu nhn bao gm cc nhim
sc th c s lng v hnh dng c trng cho t bo ca mi loi sinh vt, c bit l v tr ca tm ng.
ng vt, kiu nhn gii tnh ci v c thng khc nhau do c cc nhim sc th gii tnh X v Y. i
vi cc nhim sc th cn li, kiu nhn ca tt c cc sinh vt trong cng loi u ging nhau. Nhng loi
khc nhau c kiu nhn khc nhau.
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Hnh 3.10. Kiu nhn ngi

Pha trn: cc nhim sc th trong nguyn phn ca ngi n ng bnh thng.
Pha di: cc nhim sc th c sp xp thnh cp tng ng.
Hnh 3.10 gii thiu kiu nhn ngi nam gii, trong c 46 nhim sc th bao gm 22 cp nhim sc
th thng v hai nhim sc th gii tnh X v Y rt khc nhau. Cc nhim sc th ngi c phn thnh 7
nhm t A n G da trn hnh dng v kch thc ca chng. Nh k thut nhum bng (banding
techniques) c th lm hin ln nhng vch nang (bng) c th nm dc theo mi nhim sc th, nh vy c
th d dng phn bit c tng nhim sc th trong kiu nhn.
2.3.2. Cu to ca nhim sc th
Nhim sc th ca Eukaryota c t chc phc tp, bao gm ADN v nhiu loi protein gn vo. Trong
s cc protein gn vo , histon ng vai tr quan trng trong vic cun ADN li v iu chnh hot tnh
ca ADN. Cc histon l nhng protein nh cha nhiu acid amin mang in tch dng (lysin v arginin) nn
c th gn cht vi ADN in tch m.
S hnh thnh nhim sc th k gia t chui xon kp ADN qua h thng cc bc cu trc nh sau
(hnh 3.11):
Nucleosom l n v cu trc ca nhim sc th c to nn do si ADN di qun quanh cc
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protein histon, hnh thnh si 11nm. n v nucleosom l phc hp gm 146 cp base ca ADN qun
quanh 8 phn t histon (2 H2A, 2 H2B, 2 H3 v 2 H4). Cc nucleosom k nhau c ni qua mt phn t
nucleosom trung gian (H1, hnh 3.11b).
Si chromatin dy 30nm: Cc nucleosom xp kht nhau to thnh phc hp nucleoprotein.

Vng xp cun dy 300nm do si chromatin sau nhiu ln xon un khc to nn (hnh 3.11d).
Cht d nhim sc dy 700nm (hnh 3.11e): Cht d nhim sc l trng thi cun xon cao ca cht
nhim sc. nhng sinh vt khc nhau th cht d nhim sc phn b khc nhau, c trng hp tng phn
hoc ton b nhim sc th l cht d nhim sc.
Nhim sc th k gia dy 1400 nm (H. 3.11.f): Dng ph bin ca nhim sc th trng thi dn
xon, cn gi l cht nguyn nhim sc. C th c trng hp cht d nhim sc nm xen k vi cht nguyn
nhim sc v bc quanh cc tm ng.
V chc nng, cht nguyn nhim sc cha ADN trng thi hot ng c th phin m c, cn cht
d nhim sc th mang ADN dng khng phin m c. Cht d nhim sc sao chp mun hn cht
nguyn nhim sc trong chu trnh t bo.
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Hnh 3.11. Phc hp nucleoprotein cun li thnh nhim sc th
2.3.3. Cc trnh t lp li ca ADN

Cc th nghim v bin tnh (denaturation) v hi tnh (renaturation) ADN cc sinh vt khc nhau cho
thy nhim sc cc sinh vt nhn nguyn thy hu nh ch mang mt trnh t ADN duy nht, trong khi
nhim sc th sinh vt nhn tht li c nhiu on lp li. Nhng sinh vt nhn tht khc nhau c tn s cc
on ADN lp khc nhau. C nhng on (chiu di t vi cp n vi trm cp base) c tn s lp li n
hng triu ln trong h gen. Ni chung, nhng on ADN c tn s lp li cao thng nm quanh tm ng
v hai u ca nhim sc th, chng c sao chp mun hn cc on ADN khc v khng c phin
m, trong khi cc on c tn sut lp trung bnh (t vi chc n vi nghn ln) th c phin m. Cc gen
m ho cc rARN v tARN v cc histon nm trong cc on ny.
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2.4. Nhim sc th phage
Hnh 3.12. Phage T4 ca vi khun E. coli
Nh c cp, genom ca phage c th l ADN si n hoc si kp, hay ARN si n hoc si

kp, mch thng, ng vng hay chng lp. Hin nay, cc phage T xm nhim E. coli c nghin cu k
nht v mt di truyn (hnh 3.12). Nhim sc th ca chng l ADN trn, mch kp, khng c protein. C
phage T2 v T4 u c nhim sc th di hn h gen hon chnh ca chng. l kt qu ca s lp li d
tha hai u v hin tng i ch do quay vng i u, ging nh mi phage u cha nhng phn on
ADN c chiu di bng nhau c ct ra mt cch ngu nhin t mt vng trn nhim sc th ln. Trong khi
, cc phage T l nh phage T3, T5, T7 li ch c nhim sc th lp li d tha hai u. Cc kiu nhim
sc th ny c cc dng nh sau:
123456
H gen hon chnh.
12345612
Nhim sc th lp li d tha hai u.
456123
Cc nhim sc th i ch do mch vng.
345612
12345612
34561234
Cc nhim sc th lp li d tha hai u
61234561
v i ch do quay vng i u.
Bng chng v s tn ti cc kiu nhim sc th ni trn phage thu c trong cc th nghim x l

ADN phage vi enzym gii hn.
2.5. Cc yu t di truyn vn ng - gen nhy

Ngoi cc gen chim v tr c nh trn nhim sc th cc sinh vt tin nhn hay nhn chun u thy
tn ti cc yu t di truyn c bit c kh nng vn ng t mt v tr n cc v tr khc trong h gen. Cc
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gen di ng ny c t tn l cc yu t di truyn vn ng (transposable genetic elements - TGE).

Cc TGE l nhng on ADN c bit c kh nng chn (xen) vo mt hoc mt s v tr trong h gen v
cng c th c ct ri khi h gen. Chng c th to nn nhng bin i di truyn khi xen vo gen hoc
vo on k tip vi gen v nhng bin i ny s mt i nu cc TGE ri chuyn khi v tr m chng
chn vo. Yu t di truyn ng nh vy cn c gi l gen nhy.
2.6. Cu trc exon - intron ca gen

Cc gen ca ng vt c v, i khi ca c thc vt bc cao v nm men c cc on khng m ho acid
amin (non-coding sequences) c gi l cc intron, phn bit vi cc on m ho (coding sequences) c
gi l exon. C cc on intron v exon u c phin m to ra tin mARN (pre-mARN) v t phn t
ny qua qu trnh tch ghp (splicing) ct i cc intron, ri gn cc exon li tng hp nn cc phn t
mARN dng cho qu trnh dch m din ra trong ribosom t bo cht.
Cc intron u tin pht hin thy adenovirus (1977), sau tm thy virus SV40 v rt nhanh sau
ngi ta thy s tn ti ph bin cc intron cc sinh vt nhn chun.
Pierre Chambon v cs. Trng i hc Tng hp Strasbourg (Php) l nhng ngi u tin pht
hin cc on intron khi nghin cu iu ho di truyn trong sinh tng hp ovalbumin protein trong lng
trng trng g, cha 386 acid amin. Cc nh nghin cu tch mARN ca gen ovalbumin, sau cho phin
m ngc to ra cADN v pht hin thy cc phn t mARN u tin trong nhn l nhng phn t
mARN di hn mARN bnh thng c trong t bo cht. So snh ADN ca gen vi cADN c to ra t
mARN ly t t bo cht v gii trnh t c hai loi phn t, cc tc gi nhn thy c cc on polynucleotid
c trong ADN ca gen gc nhng li khng c trong cADN c tng hp t mARN ly t t bo cht. Cc
tc gi cn pht hin thy cc on polynucleotid ni trn (intron) khng c dch m, chng nm xen k
vi cc on c mang thng tin c dch m (exon). Gen ovalbumin c 7 intron xen k gia 8 exon theo s
di y:
77.000 cp base
7 intron: A, B, C, D, E, F, G.
8 exon: L, 1, 2, 3, 4, 5, 6, 7.
Exon L l on dn u 5 ca mARN trong gen ovalbumin.
Cc exon 1-7 m ho cc acid amin trong ovalbumin.
Intron u tin pht hin thy thc vt l gen phaseolin trong cy h u mang 3 intron. Ni chung,
cc gen thc vt bc cao u mang nhiu intron v exon. Ngi ta cng nhn thy khng phi tt c cc gen
nhn chun u mang intron. V d, gen histon nhm bin v 4 gen sock nhit Drosophila khng mang
intron.
Ni chung, cc intron pht hin c kch thc rt khc nhau t vi cp n hng nghn cp base.
Trong nhiu trng hp, mi exon m ho mt chc nng xc nh ca protein. V d, cc gen globin,
exon nm gia m ho im gn hema. nm men, trong gen cytochrom b, intron li l mt phn ca exon
thuc cc gen m ho cc enzym cn thit cho qu trnh phin m. Theo quan nim mi, cc intron khng
phi l cc on ADN d tha nh c ch ra trc y, chng ng vai tr quan trng trong qu trnh bit
ho ca sinh vt nhn chun a bo, v v vy chng l sn phm ca qu trnh tin ho. Nghin cu su hn
v chc nng ca cc intron c th thy r s tn ti ca cc intron l cn thit cho hot ng v biu hin
ca gen, c bit l cc sinh vt bc cao.
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3. SAO CHP ADN PROKARYOTA V EUKARYOTA

3.1. Sao chp ADN sinh vt nhn nguyn thy
T bo tng hp cc acid nucleic bng hai cch khc nhau: tng hp mi v tng hp cu cha. Trong
tng hp mi (de novo pathway) ng deoxyribose, mt s acid amin, CO2 v NH3 kt hp vi nhau trong
mt lot cc phn ng trc tip to ra cc nucleotid. Trong trng hp ny, purin v pyrimidin t do cng
nh cc nucleotid khng phi l cc cht trung gian trong con ng sinh tng hp. Trong tng hp cu
cha (salvage pathway) cc purin, pyrimidin, cng nh cc nucleotid sinh ra t cc acid nucleic b phn hu
li c th c chuyn ho tr li thnh cc nucleotid cn thit cho tng hp acid nucleic. C hai cch u
quan trng i vi t bo v mc quan trng ca chng tu thuc vo iu kin tn ti ca t bo.
3.1.1. Sinh tng hp purin v pyrimidin
Con ng sinh tng hp purin v c bn l ging nhau nhiu sinh vt khc nhau, v d, vi khun E.
coli, nm men v ngi. Nhiu hp cht c s dng to nn vng purin: aspartat, CO2, format,
glutamin, glycin. Cn cc cht c s dng to thnh vng pyrimidin l aspartat, CO2, glutamin.
Hnh 3.13. S sinh tng hp cc purin nucleotid ca ADN
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Hnh 3.14. S sinh tng hp pyrimidin nucleotid ca ADN
3.1.2. Sao chp theo khun

Trong chui xon kp ADN, adenin phi bt cp vi thymin v guanin phi bt cp vi cytosin nn trnh
t cc nucleotid trn mt mch s xc nh chnh xc trnh t c hiu cc nucleotid trn mch b sung ca
n. Nh vy, trong tng hp ADN nu tch ri hai mch ca phn t ADN ra, chng c th sp xp cc
nucleotid ca mch mi thnh hng theo trnh t b sung vi cc nucleotid trn mch c v sau gn cc
nucleotid li thnh mch mi b sung. Tc l, hai mch gc ca phn t ADN ban u c tch ra mi
mch lm khun tng hp mch mi. Kt qu mi phn t ADN ban u to ra hai phn t con ging ht
nhau. Mi phn t con u mang mt mch gc v mt mch mi. Kiu sao chp ny c gi l sao chp
bn bo tn (semi-conservative). Th nghim ca Meselson v Stahl c trnh by trong hnh 3.15. Nm
1958, M. Meselson v Stahl chng minh kiu sao chp bn bo tn. Hai ng nui cy E. coli nhiu th
h trn mi trng c ngun nit ng v nng N15. Nh vy, tt c ADN ca vi khun u mang ng v
nng N15 thay cho N14 bnh thng. Sau , t bo c chuyn sang mi trng ch cha N14 nh, ly mu
t bo sau nhng khong thi gian u n v chit tch ADN. Bng phng php ly tm siu tc trn
gradient nng CsCl, cc loi ADN nng, nh v lai c tch ra.
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Hnh 3.15. Th nghim ca Meselson v Stahl

Kt qu cho thy, ADN nng ban u (th h 0) cha N15, sau mt ln phn chia cho th h I E. coli vi
ADN lai c t trng nm gia ADN nng N15 v ADN nh N14. Nh vy, sau mt ln sao chp phn t
ADN mi cha mt si mang N15 v mt si mang N14. th h II c mt na s phn t ADN l lai, s
cn li l ADN nh vi N14. Nh vy, gi thuyt ca Watson Crick l ng, tc l hai mch ADN m tch
ra, mi mch lm khun tng hp nn mch mi b sung.

3.1.3. Qu trnh sao chp ADN
Cc nghin cu tip theo pht hin ra c ch phn t ca qu trnh sao chp ADN. y l qu trnh
phc tp v phi tri qua cc c ch chung nh sau:
Cc lin kt hydro gn hai si ca mch kp ADN v n nh cu trc xon phi b t ra v tch ri
hai si n;
Phi c on mi (primer), tc on ADN hay ARN mch n ngn bt cp vi mch khun;
4 loi nucleotid triphosphat (dATP, dGTP, dTTP v dCTP) bt cp b sung vi cc nucleotid
mch khun;
Mch mi tng hp theo hng 5P 3OH;
Cc nucleotid mi c gn li vi nhau bng lin kt cng tr to mch mi.
Mi bc sinh tng hp ADN c xc tc bng enzym c hiu v c thc hin mt cch nhanh
chng, chnh xc.
cc Prokaryota v c cc Eukaryota, tng mch ADN ring bit c sao chp ch theo mt hng:
cc enzym sao chp di chuyn trn mch m t u 3 n u 5 to nn mch mi b sung theo hng
5 3. Khi tch ADN mt u ra sao chp to ra ch ba sao chp (replication fork). Mch mi c
tng hp theo hng 5 3 nn trn mt mch khun qu trnh tng hp s hng t ngoi vo ch ba,
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cn trn mch khun kia qu trnh tng hp s din ra theo chiu t ch ba hng ra ngoi to ra cc
on ngn ri ni li vi nhau. Qu trnh sao chp nhim sc th vng trn vi khun E. coli c m t
trn hnh 3.16 bao gm hai giai on: khi pht (initiation) v ko di (elongation).
3.1.3.1. Khi pht
Hnh 3.16. Sao chp ADN vi khun E. coli
vi khun E. coli, qu trnh sao chp bt u khi protein B c hiu nhn bit im khi pht sao chp
(replication origin - vit tt l ori) v gn vo trnh t base c bit . Tip theo enzym gyrase (mt loi
topoisomerase) ct ADN lm tho xon c hai pha ca protein B. Trong khi hai enzym gyrase chuyn
ng ngc chiu nhau so vi im ori th hai phn t ca enzym helicase (cn gi l rep) tham gia tch
mch to ch ba sao chp. Helicase s dng nng lng ca ATP lm t cc hydro gia hai base bt cp vi
nhau. Cc protein lin kt mch n lm cng mch SSB (single strand binding protein) gn vo cc mch
n ADN lm chng tch nhau thng ra v ngn khng cho chng chp li ngu nhin hoc xon li sao
chp c d dng.
3.1.3.2. Ko di
ADN-polymerase III l phc hp gm nhiu enzym gn vi nhau bt u sao chp mt trong hai mch
bng cch gn vo mch khun v lp cc nucleotid b sung vo v tr tng ng. Tnh phc hp ca enzym
l cn thit v ADN-polymerase phi xc tc nhiu bc phn ng khc nhau v phi c kh nng s dng 4
loi nucleotid nh tc nhn phn ng ph thuc vo v tr c c trn mch khun. t nht enzym ny
phi c 4 trung tm hot ng rt linh ng, mi trung tm ng vi mt loi nucleotid (A, T, G, C). Ngoi
chc nng polyme ho theo hng 5 3, ADN-polymerase III cn c kh nng sa sai nh hot tnh
exonuclease (exonuclease l enzym ct ADN t u mt mt mch theo chiu 5 3 v ngc li 35).
Khi phc hp ADN-polymerase c theo mch khun v ko cc nucleotid b sung vo ng v tr n
lin gn cc nucleotid li lm mch mi b sung ko di thm ra. Trn ng di chuyn tng hp ADN
nu ADN-polymerase III bt gp v tr m nucleotid mi c lp vo b sai, hot tnh exonuclease 3 5
lin ct li li loi b nucleotid b sai, ri lp tc gn nucleotid ng vo tip tc sao chp. Cc nucleotid
trc khi gn vo u 3OH c hot ho tr thnh nucleosid triphosphat c mang nng lng. ADN-
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polymerase c tnh c hiu cao, v ch thm nucleotid vo u 3OH ca mch ang c tng hp.
Mch khun ADN c u 3 c ADN-polymerase III gn vo v tng hp ngay mch b sung 5
3 hng vo ch ba sao chp. Mch khun ny c gi l mch khun trc, cn mch mi c tng hp
gi l mch trc (leading strand).
Trong khi , mch khun sau c u 5 qu trnh tng hp phc tp hn v c thc hin t ch ba
sao chp hng ra ngoi duy tr ng hng 5 3.
Khi mch kp tch ra, gn ch ba sao chp enzym primase gn mi (primer) ARN cha khong 10
nucleotid c trnh t b sung vi mch khun vo. ADN-polymerase III ni nucleotid tip theo mi ARN
theo hng ngc chiu vi ch ba sao chp, tng hp cc on ngn khong 1.000 - 2.000 nucleotid, c
gi l cc on Okazaki. ADN-polymerase tip tc ko di on Okazaki n khi gp on mi ARN pha
trc th dng li, ri li ra sau tip tc tng hp t ARN mi mi c gn vo gn ch ba sao chp. Tip
theo, ADN-polymerase I nh hot tnh exonuclease 5 3 ct b mi ARN, gn cc nucleotid ca ADN
vo ch trng thc hin polyme ho hng 5 3. on ADN ngn 10 nucleotid ny cn h hai u c
ni lin ch h nh enzym ADN ligase. Mch c tng hp t ng ba sao chp hng ra ngoi c tng
hp chm hn nn gi l mch sau (lagging strand). Qu trnh sao chp ADN E. coli din ra rt nhanh, c
th t n hn 50.000 nucleotid/pht.
3.2. Cc nhn t tham gia sao chp ADN

Cc nghin cu n nay xc nh thm chi tit v cc protein v cc gen tng ng tham gia vo sao
chp ADN (bng 3.2).
Bng 3.2. Cc protein v cc gen tng ng tham gia vo sao chp ADN
Protein
M (kDa)
Gen
Chc nng
S bn sao/t bo
Rep
Helicase III
SSB protein
Protein i
Protein n
Protein n'
Protein n''
Protein dna B
Protein dna C
Primase
ADN-polymerase III
Tiu n v
Tiu n v
Tiu n v
Tiu n v
Tiu n v
Tiu n v
Tiu n v
65
75
74
66
28
76
17
29
300
60
760
140
37
52
32
25
10
83
Rep
Tch mch
Tch mch
n nh mch n
50
ADN-polymerase I
102
polA
Ligase
ADN topoisomerase I
ADN topoisomerase II
75
lig
Ssb
dnaT
Tham gia vo cu trc ca

prisom
dnaB
dnaC
dnaG
polC
dnaN
dnaZ
dnaX
dnaQ
Tng hp mi (primer) ARN
300
50
80
70
100
20
50
20
Ko di mch ADN ang

tng hp v kim sot s
chnh xc
Ct mi tng hp ADN ngn

k h
Hn kn mch
Ct h mt mch
Ct h hai mch
300
3.3. Sao chp nhim sc th Prokaryota
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theo di sao chp ADN, dn cht ng v phng x thymidin c s dng. Qu trnh sao chp c
khi pht t im ori (origine) gi l im khi pht v tin trin ra c hai pha (hnh 3.17). Khi ADN vng
ang sao chp quan st thy dng ADN hnh con mt (eye replication). Ch ba sao chp lan dn, cui cng
to ra hai phn t ADN lai: mt mch c mang cht nh du phng x T-H3 (thymidin H3). C trng hp
sao chp ch xy ra v mt pha.
Vi khun E. coli ch c mt im xut pht sao chp ori nn c ADN thnh mt n v sao chp thng
nht c gi l replicon. Nh vy, replicon c ngha l n v sao chp. Genom ca cc t bo tin nhn
thng ch c mt replicon.
Hnh 3.17. Sao chp replicon t bo vi khun
3.4. Sao chp nhim sc th Eukaryota

T bo nhn thc c s lng ADN ln hn nhiu so vi t bo tin nhn, to nn nhiu nhim sc th,
m mi nhim sc th bao gm si ADN thng c kt hp vi protein. Do , sao chp ADN ca t bo nhn
thc phc tp hn.
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Hnh 3.18. Sao chp nhiu replicon
im khc cn bn trong sao chp ADN t bo nhn thc v t bo tin nhn l ADN ca t bo nhn
thc c nhiu replicon. V d, t bo nm men bnh m Saccharomyces cerevisiae c ti 500 replicon, tc
l c 500 im ori khi pht sao chp. Qu trnh sao chp cng bt u t ori ri lan v hai pha nh trnh
by trong hnh 3.17. T bo c c ch kim sot nghim ngt qu trnh sao chp, im ori no sao chp
qua mt ln th s khng sao chp li trc khi ton b ADN c sao chp hon ton (hnh 3.18). Sau khi
sao chp xong, t bo c c ch phn chia u ADN vo cc t bo con.
3.5. Sa cha sao chp

S sao chp chnh xc ADN c ngha sng cn i vi hot ng bnh thng ca t bo. Nhim sc
th l ni lu gi cc thng tin cn thit cho vic xy dng v iu khin ton b hot ng ca t bo. V
thng tin cho s lp rp chnh xc hng ngn protein khc nhau, c chiu di hng trm hoc hng ngn base,
s sai lch mt vi base trong s c th d xy ra v c th s dn n nhng hu qu nghim trng. Bn
thn ADN di, mng manh, trong qu trnh sng thng xuyn chu tc ng ca cc nhn t bn trong v
bn ngoi t bo cho thy ch ring c ch sao chp chnh xc cha m cn thit phi n nh cu trc
ADN qua nhiu th h t bo.
3.5.1. Sa sai trong sao chp
S dng cc nucleotid v ADN-polymerase tng hp ADN in vitro sai st c pht hin l 1.10-5,
tc l trong s 100.000 nucleotid c mt nucleotid b sai. Tuy nhin, mc sai st ny cho thy sao chp ADN
in vitro c chnh xc cao, nhng nu em so snh vi sao chp ADN trong cc sinh vt th mc sai st
ny vn cn qu ln. V d, E. coli c 3.106 cp base, vy mi ln sao chp c 30 sai st, c th dn n
nhiu t bin, nhng trong thc t khng c nhiu t bin n nh vy qua mi ln sao chp.
ngi, ADN ca mt t bo c 3.109 cp base. Vi sai st 1.10-5 s c 30.000 sai st trong mt ln
sao chp. So vi thc t, mc sai st nh vy l qu ln. ng vt c v xut hin cch y 300 triu nm
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(3.108 nm), nu sai st khi sao chp ADN ln n nh vy th kh m c s ging nhau gia protein
ngi v cc protein c voi. Nhng thc t cho thy c nhng protein ging nhau gia ngi v c voi.
Bng cch nh gi tn s cc t bin mi xut hin trong qun th ln v theo di bin i enzym
trong nui cy m t bo c th tnh c rng trong c th sinh vt sai st khi sao chp ADN (in vivo) l
1.10-9, tc 1 sai st trn 1 t base. Nh vy, t bo ngi c 3.109 cp nucleotid mi ln sao chp c th c 3
sai st.
nh gi mc bin i trong tin ho cng khng nh mc chnh xc rt cao trong sao chp in
vivo ADN. Mc chnh xc cao trong sao chp ADN ca c th sinh vt c c nh cc c ch sa sai
sau:
Hng sao chp bao gi cng t u 5 3 vic sa sai c chnh xc.
Cc ADN-polymerase I v III va polyme ho, va c hot tnh exonuclease 5 3 v 3 5, nu
trn ng di chuyn polyme ho bt gp nucleotid b lp sai, ADN-polymerase s li li ct b theo hng
3 5 (hot tnh exonuclease 3 5).
3.5.2. Sa sai khi khng sao chp
Phn t ADN c th b bin i ngay c khi khng sao chp. Cc bin i t bin ny xy ra vi xc
sut kh cao. Nh c ch sa sai nn tn s t bin c duy tr mc thp.
Chin lc sa sai trn nhim sc th v c bn ging nhau khi sa sai trong sao chp v khi b t bin:
cc enzym nhn bit gn vo cc trnh t sai v ct ri cc on sai, ri s dng mch n ng lm khun
tng hp li on b ct cho ng. Hng lot enzym c hiu lm nhim v d tm v sa sai. C khong
20 enzym r sot dc nhim sc th d tm cc base c bin i ho hc, mi enzym c chc nng chuyn
bit cho mt loi sai lch. Khong 5 enzym khc c hiu cho cc lin kt cng tr sai gia base vi cc hp
cht khc hoc gia cc base k nhau trn mt mch. Mt s enzym khc pht hin s bt cp sai, nh
trng hp mt purin. Tng cng c khong 50 enzym chuyn pht hin v sa cha cc sai lch trn phn
t ADN.
4. CHU TRNH T BO
Phn chia t bo l c tnh c bn ca tt c cc sinh vt. cc t bo nhn nguyn thy, qu trnh sao
chp ADN c bit kh tng tn. V d, khi nui cy trong mi trng dinh dng, cc vi khun tng hp
ADN trong mt chu trnh t bo v sau hai t bo con c sinh ra nh thnh t bo mi hnh thnh v ct
ngang t bo. Ngc li, chu trnh t bo ca cc sinh vt nhn tht phc tp hn rt nhiu. S sinh sn v
tnh v tng trng ca cc m c th nh nguyn phn (mitosis) l kiu phn bo to ra cc t bo con c
kiu gen ging y nh t bo m. Sinh sn hu tnh nh gim phn l kiu phn bo to cc t bo sinh dc
hay giao t c s nhim sc th gim mt na vi kiu gen a dng.
4.1. Chu trnh t bo nhn tht
cc sinh vt nhn tht n bo cng nh a bo, s sinh sn ca t bo l mt qu trnh sinh trng,
phn nhn v phn bo mang tnh chu k. Qu trnh gi l chu trnh t bo (cell cycle). phn ln cc t
bo soma ca thc vt bc cao v ng vt, chu trnh t bo gm 4 pha (hnh 3.19) bao gm: G1 (gap 1), S
(synthesis), G2 (gap 2) v M (mitosis). Ba pha u gp li gi l giai on ngh hay k trung gian
(interphase). Tip sau G2, t bo phn chia nguyn phn cho hai t bo con. Sau , mi t bo con li bt
u mt chu trnh t bo mi. cc t bo ng vt khi nui cy, mt chu trnh t bo ko di khong 24
gi. i vi cc t bo nhn tht, thi gian ca mi pha cng nh ca chu trnh t bo nhng sinh vt khc
nhau c khc nhau (bng 3.3).
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Hnh 3.19. S chu trnh t bo

Bng 3.3. Thi gian chu trnh t bo v t l cc pha
Sinh vt
T bo
u Viciafaba
Chut
Chut ti
Thi gian chu trnh

(gi)
19,3
G1
(%)
25,4
S
(%)
38,8
G2
(%)
25,4
M
(%)
10,4
20,0
41,4
44,9
10,5
3,2
11,0
24,5
52,8
19,1
3,6
Nguyn bo
si
Nguyn bo
si
4.2. Pha G1
Pha G1 din ra tip sau nguyn phn, nhim sc th bin i t trng thi kt c trong nguyn phn
sang trng thi ko di ca pha ngh, ng thi din ra hng lot nhng bin i dn n s khi u sao
chp ADN. di ca G1 thay i ph thuc vo hm lng protein trong t bo, v c s khc bit gia
cc t bo cng th h, cng nh c nh hng ln n thi gian chu trnh chung. Nhim sc th trong G1 t
xon vn v hot tnh phin m tng ln. Mt phn hot ng phin m ny dn n vic sinh tng hp hng
lot phn t cn thit khi u tng hp ADN. Tuy nhin, vn c mt s sinh vt khng c G1 r rt, v
d, mc nhy Physarum polycephalum, nm men Schizosaccharomyces pombe, phi ca Xenopus v phi
chut. S tch lu vt cht ni bo t n im ti hn (restriction site) th t bo bt u tng hp ADN.
4.3. Pha S
4.3.1. Tng hp ADN khng lin tc
Trong pha S, vt cht di truyn ca nhim sc th c nhn i. Trong khi m nui cy ca cc t bo
ng vt, pha S ko di t 6 n 10 gi th t bo nm men in vivo Schizosaccharomyces pombe l 10
pht, hoc c th ko di ti 35 gi t bo da tai chut. S sao chp ADN din ra theo kiu na gin on:
cc nucleotid gn vi nhau to thnh cc on ngn (khong 100 nucleotid) ri cc on gn li vi nhau
to thnh nhng si di hn.
4.3.2. Cc ADN polymerase
cc sinh vt nhn tht, qu trnh tng hp ADN c khi pht bi cc on mi ARN di 9-10
nucleotid, sau cc on mi b ct b. cc sinh vt nhn chun bc cao c 3 ADN polymerase: ADN
polymerase alpha (), beta () v gama (), chng khc nhau v khi lng phn t, tnh mn cm vi cht
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c ch N-ethylmaleimid, tnh mn cm vi mui v kh nng sao chp nhng khun khc nhau (bng
3.4). C polymerase v u trong nhn, trong khi enzym polymerase li trong ty th. Cc enzym
polymerase v polymerase thng nh vi cc protein khc trong phc hp sao chp. Ni chung l
chng rt khc nhau trong cc sinh vt nhn tht. Mt s sinh vt c nhiu dng enzym khc nhau trong khi
mt s li thiu. Thc vt, protozoa, nm thiu ADN polymerase , cn cc vi sinh vt nhn tht ni chung
li c polymerase , mc d ADN polymerase ny rt khc ADN polymerase ca cc ng vt c v.
Bng 3.4. Tnh cht ca cc ADN polymerase ca ng vt c v
Loi enzym
Phn t lng
B c ch bi
N-ethylmaleimid
120.000 - 300.000
30.000 - 50.000
Khng
150.000 - 300.000
Tc dng vi mui
B c ch bi NaCl 25mM
c kch thch bi NaCl 100 - 200 mM.
B c ch bi phosphat 50mM
c kch thch bi KCl 100 - 250mM
v phosphat 50mM
cc sinh vt nhn tht bc cao ni chung ADN polymerase c s dng sao chp ADN ca
nhn, ADN polymerase c th hot ng nh enzym sa cha, cn ADN polymerase th chuyn trch
vic sao chp h gen ty th. Khng c enzym no trong s ny c hot tnh c sa. Ngc li, cc sinh vt
nhn tht bc thp, chng hn nh nm men c cc ADN polymerase trong nhn rt ging vi cc enzym ca
cc sinh vt nhn nguyn thy v c hot tnh c sa.
4.3.3. Cc n v sao chp
Cc nhim sc th nhn tht c nhiu n v sao chp (replication unit, RU), cng c gi l cc
replicon, mi replicon c mt im khi u v 2 im kt thc sao chp, y l cu to cn thit sinh vt
nhn tht c th sao chp s lng ADN khng l trong thi gian va phi.
Hnh 3.20. Khi u v kt thc sao chp ADN trong cc Eukaryota
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Qu trnh sao chp ADN xut hin bng vic xut hin chic vng trn ADN gm hai chc sao chp.
Cc chc ny chuyn ng ngc chiu nhau trong qu trnh tng hp ADN cho n khi chng t n cc
im kt thc c th. Cc tn hiu xut pht v kt thc tng hp ADN l nhng on ADN c hiu:
protein khi pht c hiu bm vo on ADN khi pht v nh h tr phc hp sao chp ADN bm vo
im khi pht ca qu trnh sao chp. Khong cch gia hai im khi pht sao chp thay i t 30.000 300.000 cp base nit tu thuc vo sinh vt c nghin cu. Trn nhim sc th ca t bo Hela (t bo
ung th ngi) c 100 n v sao chp. Tc chuyn ng ca phc h sao chp khong 1.000 n
15.000 nucleotid trong mt pht 37oC. Nhng tc chuyn ng ca tng chc sao chp rt khc nhau
ngay trong cng mt h thng v chu s kim sot di truyn v ph thuc vo bin i mi trng (hnh
3.18). Qu trnh bt u t O chuyn ng ngc chiu to thnh cc vng m. Qu trnh kt thc khi hai
chc sao chp k cn ho nhp vo nhau (n im kt thc t). ADN giu GC c sao chp trc, cn on
ADN giu AT c sao chp sau (hnh 3.20 trn).
4.3.4. Lp ghp ADN vo cc th nhn
Khi ADN sao chp th cc th nhn (nucleosom-bao gm histon v ADN) cng phi c nhn i, cc
histon phi c tng hp mi. Cc gen tng hp histon hot ng trong G1 v S to ra s lng ln cc
histon. ADN mi sao chp nhanh chng kt hp vi cc histon mi cn thit hnh thnh cc nucleosom.
Cc histon mi c s dng ngay ti cc chc sao chp to thnh cc nucleosom mi, v 2-15 pht sau
chng c sp xp li to thnh cc nhim sc th trng thnh.
4.4. Pha G2
Cc nhim sc th kt cht li chun b cho nguyn phn. Kt cht l qu trnh cun xon mc cao
hn ca cc nhim sc th. Vic tng hp c ARN v protein l cn thit hon thnh G2. Kt thc ca G2
bo hiu s bt u ca nguyn phn. Trong G2, protein tubulin c tng hp. Tubulin c trng hp
to ra cc vi ng ca b my thoi phn ly cc nhim sc th v cc nhn con trong nguyn phn v gim
phn.
4.5. Cc kha cnh phn t ca nguyn phn
Nguyn phn l s phn chia mt t bo thnh hai t bo con ging ht nhau v mt di truyn. Trc khi
nguyn phn, cc nhim sc th c nhn i. c im ca nguyn phn l c nhng bin i ln v cu
trc v chc nng ca t bo. Khi bt u nguyn phn, qu trnh tng hp cc cht trong t bo gim xung,
ri dng li k gia v li tip tc k cui. Tc tng hp protein b gim t ngt khi bt u nguyn
phn, sau li tng ln cui k nh tng hp ARN. Cng vi vic c ch cc qu trnh tng hp ARN v
protein th mng nhn v hch nhn b v, cc cu trc ny li hnh thnh tr li trong cc nhn con cui k.
5. CC KIU PHN BO
i vi vi khun, qu trnh tng hp ADN xy ra lin tc trong sut chu trnh ca t bo. Khi hm
lng ADN c nhn i th t bo to ln v bt u tng hp thnh t bo vng gia t bo. S kin ny
lm tch hai t bo con v hai pha ca t bo v khi thnh t bo mi hon thin, vi khun tch thnh hai t
bo con ring bit. Qu trnh lp li lin tc khi sinh vt cn trong chu k sinh trng. Chu trnh t bo
sinh vt nhn tht din ra vi nhng khc bit ng k. S phn chia cc t bo soma (khng to thnh giao
t) tri qua 4 giai on: Pha G1, pha S, pha G2 v pha M (nh trnh by trn). Cc nhim sc th c
sao chp trong pha S (n bi hoc lng bi) v trong qu trnh nguyn phn (pha M) phn ly v hai t bo
con.
5.1. Nguyn phn
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Hnh 3.21. S nguyn phn t bo ng vt

a) K u; b) K gia; c) K sau; d) K cui.
Nguyn phn l c ch duy tr s lng cc nhim sc th trong cc t bo soma (n bi hoc lng

bi) trong qu trnh phn bo bnh thng. Khi t bo pha ngh c nhum nhn t bo c th quan st
c di knh hin vi, c bao bc bi mng nhn. Trong nhn c mt hoc hai vng cha nhiu ARN gi
l hch nhn. Trong pha S, mi nhim sc th c sao chp nhng tm ng khng phn chia. Thc ra
ADN ca tm ng c nhn i, nhng ch mt cu trc nhn thy c. Sn phm sao chp ca hai
nhim sc th l hai bn sao chnh xc gi l cc nhim sc t ch em (sister chromatids), vn dnh vi nhau
bi tm ng cha phn chia. Nguyn phn c chia thnh 4 giai on gi l k u, k gia, k sau v k
cui (hnh 3.21). T bo trong hnh 3.21 l t bo lng bi, i vi t bo n bi nguyn phn cng din ra
tng t.
5.1.1. K u
Cc nhim sc th hin r dn ln do b xon li.
Mi nhim sc th gm hai nhim sc th ch em.
Cui pha u, hch nhn v mng nhn b tan v.
5.1.2. K gia
Cc si thoi xut hin v to ra t hai cc i din ca t bo. ng vt cc si thoi dnh vi cc cu
trc gi l trung t (centriole) nm hai cc; cc cu trc nh th khng thy c cc t bo thc vt.
Cc si thoi ln lt gn vi tm ng ca nhim sc th.
Cc nhim sc t ch em tp trung trn mt phng ct ngang t bo mt vng gi l tm k gia
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(metaphase plate).
5.1.3. K sau
Giai on ny bt u khi tm ng c phn i, cc nhim sc t ch em tch ri nhau ra thnh cc
nhim sc t con v tin v hai cc i din ca t bo. Nh vy, hai bn sao ging ht nhau ca nhim sc
th c tch v hai cc khc nhau.
5.1.4. K cui
Kt thc qu trnh vn chuyn cc nhim sc th con v hai cc ca t bo, vi s lng v thnh phn
ging ht nhau.
Mng nhn hnh thnh bao bc mi nhm nhim sc th.
Ti to li cc hch nhn.
Cc si thoi bin mt.
Cc nhim sc th dn xon v tr nn khng quan st c di knh hin vi c na. Lc ny thy
r hai nhn pha ngh in hnh.
Trong phn ln trng hp, tip theo k cui l qu trnh phn chia t bo cht (cytokinesis), tch hai
nhn mi v hai t bo con ring bit. cc t bo ng vt, qu trnh phn chia t bo cht din ra vi vic
to thnh mt eo gia t bo, eo tht dn li cho n khi hai t bo con c hnh thnh. Ngc li, cc t
bo thc vt phn chia bng cch to ra mng v thnh t bo mi gia hai nhn mi m sinh ra hai t bo
mi.
Nh vy, t quan im di truyn hc nhng im mu cht ca nguyn phn l:
Mi nhim sc th phn chia cho hai nhim sc t ging ht nhau trong pha S ca chu trnh t bo.
Cc nhim sc th gm hai nhim sc t sp xp mt cch ngu nhin trn tm k gia.
Nguyn phn duy tr thng tin di truyn ca t bo t th h ny sang th h sau.
5.2. Gim phn

Chu trnh hu tnh ca mt sinh vt lng bi bao gm s lun phin ca cc trng thi n bi v
lng bi:
Gim phn l qu trnh sinh ra cc giao t hoc bo t n bi t t bo lng bi sau hai ln phn chia.
Trong qu trnh gim phn, cc cp nhim sc th tng ng sau sao chp tham gia vo qu trnh sp xp
li sao cho mi sn phm trong s 4 sn phm ca gim phn nhn c mt i din ca mi nhim sc th.
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Hai ln phn chia nhn gi l gim phn I v gim phn II.

5.2.1. Gim phn I
Trong gim phn I lng nhim sc th gim i mt na, t lng bi xung n bi.
5.2.1.1. K u I
S lng nhim sc th lng bi hin r dn theo qu trnh cun xon. Khc vi pha u ca nguyn
phn, mi nhim sc th y xut hin n l.
Cc nhim sc th tng ng kt hp vi nhau thnh tng cp.
Mi nhim sc th phn i thnh hai nhim sc th ch em.
Hnh thnh cc th lng tr v din ra trao i cho gia cc nhim sc th bt ngun t cha v m.
cui k u I, do trao i cho xut hin cc cu trc hnh du nhn gi l th vt cho (chiasma).
5.2.1.2. K gia I
Vo thi im bt u pha gia I, hch nhn v mng nhn bin mt. Cc tm ng khng phn chia
nh vi cc si thoi v cc nhim sc t cn gn kt vi nhau nm trong vng ca tm pha gia.
5.2.1.3. K sau I
Cc nhim sc th ca mi th lng tr, tc l cc cp nhim sc th tng ng, tch nhau ra v tin
v hai cc i din, hnh thnh cc t hp mi ca nhim sc th c ngun gc t cha v m.
5.2.1.4. K cui I v pha ngh
S phn chia t bo cht din ra to thnh hai t bo con, mi t bo ch c mt b nhim sc th n
bi hon chnh (mi nhim sc th c hai nhim sc t nh vi nhau qua tm ng) v mi nhim sc th c
th xut x t cha hoc m vi xc sut bng nhau. Trong k ngh, cc nhim sc th ko di ra v mng
nhn hnh thnh tr li.
5.2.2. Gim phn II
5.2.2.1. K u II v k gia II (hnh 3.22)
Cc nhim sc th m dn ln v cc tm ng bt u phn chia trong k gia II. B my hnh thoi
hnh thnh, cc si thoi dnh vo tm ng, cc nhim sc th nm trn mt phng phn chia.
5.2.2.2. K sau II
Cc tm ng chuyn dch v hai cc i din ca thoi, ko theo cc nhim sc t m lc ny tr
thnh cc nhim sc th con.
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Hnh 3.22. S gim phn

A v B l hai cp nhim sc th tng ng khc nhau;
1 v 2 l hai kiu t hp ca cc nhim sc th khng tng ngkhi to thnh giao t.
5.2.2.3. K cui II
Mi t bo sinh ra sau gim phn I by gi phn chia. Do vy sinh ra 4 t bo n bi bt u t mt
t bo lng bi ban u trc gim phn.
Cc nhim sc th nht dn v mng nhn hnh thnh.
Nh vy, mi t bo trong s 4 t bo n bi mi c sinh ra mang mt na s nhim sc th ca t
bo lng bi bnh thng, tc l mang mt nhim sc th ca mi cp tng ng. V mi t bo n bi
nh th c cha rt ngu nhin cc nhim sc th c ngun gc khc nhau (t cha hoc m).
Hot ng ca nhim sc th trong gim phn lin quan trc tip n phn ly ca cc gen. C hai chi tit
cn ch lin quan n vic hnh thnh cc t hp nhim sc th mi trong gim phn I. l hai nhim sc
th tng ng (cng cp nhng khc ngun gc) phi tch nhau i v hai cc. Trong khi hai nhim sc
th khng tng ng (khc cp) th li c th cng i v mt cc, hoc i v hai cc khc nhau; xc sut
din ra hai s kin ni trn l bng nhau.
5.3. Cu trc phn t ca tm ng v im mt
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Tm ng ng vai tr rt quan trng trong nguyn phn v gim phn, do chng ta quan tm n
cu trc phn t ca tm ng, im nh ca cc nhim sc th vi cc si thoi, v ca im mt (telomere)
tc l u mt ca nhim sc th.
5.3.1. Tm ng
Trnh t ca mt s tm ng c tch ra t mt s sinh vt nhn tht bc thp c xc nh. Trnh
t ca cc vng li (core region) I, II v III ca ba tm ng ca nm men CEN3, CEN4 v CEN11 nh sau:
I
II
III
Cen3
ATAAGTCACATGAT...88 cp base, 93%AT,...TGATTTCCGAA
Cen4
AAAGGTCACATGAT...82 cp base, 93%AT,...TGATTACCGAA
Cen11 ATAAGTCACATGAT...89 cp base, 94%AT,...TGATTTCCGAA
Nh vy, mc d cc tm ng khc nhau nhng c chung mt chc nng trong t bo, cc trnh t ca
chng tng t nhau nhng khng ging nhau hon ton: vng I c trnh t tng i ging nhau nhng
tm ng nm men nghin cu, vng II l vng rt nhiu AT nhng khng c trnh t ging nhau, cn
vng III tuy ngn nhng c ging nhau rt cao. Vng I t quan trng hn i vi chc nng ca tm ng
so vi vng II v III.
Khi x l th nhn (nucleosom) ca tm ng vi nuclease cho thy, vng li I, II v III ca tm ng
khng chu tc dng ca ADNase I v nuclease ca cu khun. ADN bao quanh tm ng c bc gi
thnh cc th nhn theo mt kiu nht qun hn so vi cch gi ca phn ADN cn li ca h gen.
5.3.2. im mt
im mt l vng ADN u mt ca phn t trong ADN mch thng, cn thit cho sao chp v n
nh ca nhim sc th. Phn ln im mt nm ngay di mng nhn v chng thng kt vi nhau cng
nh dnh vi mng nhn. Cc s liu cho thy tt c cc im mt c chung mt trnh t nucleotid. Cc trnh
t c th chia lm hai loi:
(1) Nhng vng gn vi u mt ca nhim sc th thng cha nhng on trnh t lp li ca ADN,
nhng nhng vng ny rt phc tp, bao gm hng nghn cp base tnh t u ADN. Cc on ny l cc
on lin kt im mt (telomere associated), c th ng vai tr trung gian trong nhiu mi quan h c
hiu ca cc im mt, nh quan h gia cc im mt vi nhau v gia cc im mt vi mng nhn.
(2) Nhng on trnh t ngay st u ca phn t ADN nhim sc th gm nhng on trnh t n
gin, lp li hai nucleotid mt, gi l cc on u mt n gin, thc hin nhng chc nng quan trng ca
u mt, c th l to cho u nhim sc th tnh n nh. V d, Tetrahymena on lp li gm 5CCCCAA-3 v Tripanosoma on lp li l CCTAA.
6. DI TRUYN NHIM SC TH V DI TRUYN NGOI NHN

6.1. Cc nh lut Mendel
c im vn ng ca nhim sc th trong nguyn phn v c bit l gim phn c trnh by
trong cc phn trc. Gi y chng ta cn xem xt mi quan h gia quy lut phn ly ca cc gen v quy
lut vn ng ca nhim sc th trong gim phn m ln u tin c Gregor Mendel nghin cu v
cng b t th k XIX, t nn mng cho di truyn hc nhim sc th.
6.1.1. Hin tng tri ln trong sinh gii
i tng nghin cu l u H Lan Pisum sativum. Loi u ny t th phn, nhng cng c th buc
chng th phn cho. Mendel nghin cu cc ni thun (ng hp t), quan st tng tnh trng ring bit
trong mi th nghim v nhn thy u H Lan c nhng kiu hnh tng phn r rt, nh ht trn v ht
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nhn, ht vng v ht xanh, thn di v thn ngn, v.v... Mendel thc hin cc php lai gia cc ni
thun v nghin cu th h con theo cc kiu hnh ca cha m.
V d:
P
F1
trn
x
trn
nhn
V tt c cc ht u thu c t F1 u ging vi mt trong hai dng cha m v u trn nn Mendel

gi tnh trng trn l tri, cn tnh trng nhn l ln. Ngy nay, ngi ta thy tri ln l hin tng ph
bin i vi mi sinh vt.
Nh vy, tt c cc con lai ca th h th nht (F1) u ging nhau v u mang tnh tri. y l pht
hin ht sc quan trng ca Mendel.
6.1.2. nh lut phn ly
Tip theo, Mendel gieo cc ht u th h F1 cho pht trin thnh cy lai v cho t th phn, thu c
cc kt qu sau:
x
F1
trn
x
trn
F1
F2
5474 trn
1850 nhn
T l cc kiu hnh F2 l 2,96 : 1 hoc ni chung l 3 : 1 gia trn v nhn.

T nhng kt qu th nghim trn, Mendel rt kt lun ng ch sau: nhn c cc dng thun
nh dng cha m th c t bo trng v phn hoa u phi c cng mt cu trc di truyn. Khi cc cy u F1
ch mang tnh trng ca mt dng cha - m c t th phn, F2 xut hin cc tnh trng ca c hai dng
cha m, chng t rng cc cy u th h F1 phi mang c hai bn sao khc nhau, mi bn sao c ngun gc
tnh trng khc nhau; t l 3 : 1 thu c trong F2 l do cc loi giao t khc nhau c sinh ra vi xc
sut nh nhau v s kt hp cc giao t c v ci din ra mt cch ngu nhin. Theo quan nim ngy nay,
cc t bo soma lng bi mang mt b nhim sc th ca cha (b n bi) v mt ca m. Suy rng ra, c
mi cp nhim sc th tng ng trong t bo lng bi th mt l ca cha v mt l ca m. Cn cc giao
t l cc t bo n bi do gim phn sinh ra, ch mang mt nhim sc th ca mi cp tng ng. Cc gen
nm trn nhim sc th. V php lai trn nhn c th minh ho c bng cc k hiu di truyn sau:
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Cc con lai F1 l d hp t v c kiu hnh trn v alen S tri so vi alen s, cn cc cy F2 sinh ra do F1

t th phn c t l kiu hnh 3 trn : 1 nhn.
Cc cy ng hp t tri SS v d hp t Ss c kiu hnh ging nhau. phn bit SS v Ss ta s dng
php lai phn tch, ngha l lai hai dng tri ni trn vi dng cha m ng hp t ln:
Trn Nhn Giao t tt c trn 1/2 Ss (trn) 1/2 ss(nhn)
Kt qu l: 1/3 s c th F2 trn cho tt c cc c th th h sau trn, 2/3 s c th trn ca F2 cho 1/2 c

th ca th h sau trn, cn 1/2 l nhn. T tt c cc s liu ny, Mendel pht biu nh lut phn ly nh
sau (theo vn phm hin i):
Cc giao t l thun khit, tch bit nhau, chng ch mang mt trong hai bn sao ca mi gen, v th h
mi c sinh ra do t hp ngu nhin ca cc giao t bt ngun t hai cha m.
V bn cht, Mendel phn bit c cc nhn t to nn tnh trng (tc l gen) v bn thn cc tnh
trng.
Tri khng hon ton. Trong cc th nghim ca Mendel nu trn, mt tnh trng l tri hon ton so
vi tnh trng kia nn cc c th d hp t v kiu hnh khng phn bit c vi cc c th ng hp t tri.
Nhng khng phi bao gi cng nh vy. i vi nhiu tnh trng quan st thy hin tng tri khng hon
ton, ngha l cc gen AA, Aa v aa c ba kiu hnh khc nhau. V d v mu hoa ca cy Antirrhinum. Cy
AA c hoa , cy aa c hoa trng, cn cy Aa c hoa mu hng. V vy, khi lai hai cy c hoa mu hng
vi nhau cho cc cy hoa , hng, trng vi t l tng ng 1 : 2 : 1.
6.1.3. C ch phn t ca tnh tri
Trong trng hp tri hon ton, nu gen m ho mt enzym, cn alen ln ch sinh ra mt protein khng
c hoc c rt t hot tnh enzym. Qua ngn cn qu trnh iu chnh tc phin m v dch m, cc th d
hp t s ch sinh ra khong mt na s lng enzym c hot tnh so vi cc th ng hp t kiu di. Nu
s lng l c th thc hin cc chc nng ho sinh mt cch bnh thng, th s xut hin kiu
hnh bnh thng.
6.1.4. nh lut di truyn c lp
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Sau khi chng minh rng cc cp tnh trng tng phn u di truyn ging nh cp trn-nhn, Mendel
bt du nghin cu s phn ly ca hai cp gen trong cng mt php lai. ng chn nhng cp gen khng
lin kt, tc mi cp nm trn mt cp nhim sc th tng ng ring bit, ng tin hnh php lai gia
cy c ht trn vng vi cy c ht nhn xanh (hnh 3.23).
Cc c th F1 l d hp t kp v v vy sinh ra 4 loi giao t vi s lng bng nhau: SY, Sy, sY v ss.
Khi cho F1 t th phn th 4 cp giao t ny kt hp vi nhau mt cch ngu nhin trong tt c cc t hp c
th c sinh ra th h con lng bi. Trong bng Punnett trn hnh 3.23 c 16 t hp giao t. Do tc dng
ca tnh tri ch c 4 loi kiu hnh xut hin: trn-vng, trn-xanh, nhn-vng v nhn-xanh. Cc kiu hnh
ny phi xut hin vi t s 9 : 3 : 3 : 1, v l kt hp ngu nhin cc giao t v s phn ly c lp ca hai
cp gen cng vi nhim sc th mang chng trong qu trnh gim phn.
Trong th nghim ca mnh, Mendel nhn c cc kt qu c th nh sau: 315 ht trn-vng:108 ht
trn-xanh: 101 ht nhn vng : 32 ht nhn xanh. T l ny rt st vi t l 9 : 3 : 3 : 1 d on theo l
thuyt.
Trn, vng Nhn,xanh Kiu gen Giao t Trn,vng
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Cc kiu hnh F2:

9 trn, vng: 3 trn, xanh: 3 nhn, vng: 1 nhn, xanh
Hnh 3.23. Nguyn tc phn ly c lp
T l phn ly kiu hnh F2: 9 : 3 : 3 : 1 i vi hai cp gen khng lin kt (s dng bng Punnett).
Nh vy, cng thc phn ly theo kiu hnh F2 khi lai hai cp tnh trng trong cc th nghim ca Mendel
l: 9 A - B -: 3A - bb: 3 aaB -: 1aabb.
y, Aa v Bb l hai cp gen nm trn hai cp nhim sc th tng ng khc nhau.
T cc kt qu nhn c trong cc php lai hai tnh trng nu trn, Mendel c nhn xt rng, t l phn
ly 9: 3: 3: 1 khi xem xt ng thi hai tnh trng trong mt php lai chnh l s kt hp ca hai t l phn ly
3: 1 khi xem xt tng cp tnh trng ring bit. C th l t cng thc tng qut ni trn, nu ch xem xt cp
tnh trng Aa ta s c: 9A - + 3A -: 3aa + 1aa, tc l 12A -: 4aa hay 3A -: 1aa. Tng t nh vy, nu ch
xem xt cp tnh trng Bb ta s c: 9B - + 3B -: 3bb + 1bb tc l 12B -: 4bb hay 3B -: 1bb. T nhn xt ny,
Mendel pht biu nh lut th 2 ca mnh gi l nguyn l di truyn c lp: cc gen nm trn cc
nhim sc th tng ng khc nhau th vn ng c lp trong qu trnh to thnh giao t.
6.1.5. Cc cng thc lai a tnh
Vic phn tch s di truyn ca mt cp tnh trng trong lai n tnh gip Mendel v cc nh nghin
cu sau ny hiu c s di truyn ca hai cp v nhiu cp tnh trng trong cc php lai hai tnh v a tnh.
V ngy nay, chng ta hiu r rng cc quy lut di truyn l do quy lut vn ng ca cc nhim sc th
trong gim phn quy nh. T l phn ly kiu hnh F2 ca mi tnh trng l 3: 1. y l t l phn ly chnh
xc ca mt cp nhim sc th tng ng trong gim phn. Vi hai cp tnh trng, t l l 9: 3: 3: 1, tc
l (3 + 1)2, v vi n cp tnh trng th cng thc phn ly v kiu hnh F2 s l (3 + 1)n. Vi l lun tng
t, cc cng thc c bn trong trng hp lai a tnh (n tnh trng) c th c xc nh nh trong bng 3.5
sau y:
Bng 3.5. Cc cng thc c bn lai a tnh trng
Tn cng thc
Kiu lai
n tnh
Hai tnh
a tnh
S kiu giao t do F1 to ra
22
2n
S t hp giao t sinh ra F2
42
4n
S loi kiu hnh F2
22
2n
S loi kiu gen F2
32
3n
3+1
(3 + 1)2
(3 + 1)n
1+2+1
(1 + 2 + 1)2
(1 + 2 + 1)n
Phn ly theo kiu hnh F2

Phn ly theo kiu gen F2
6.1.6. Cc khi nim bt ngun t cc nh lut Mendel

a alen: i vi mt gen c th khng phi ch c hai m c th c nhiu alen, quan h tri - ln gia
cc alen rt khc nhau. V d, ba alen xc nh h thng nhm mu ABO ca ngi c quan h tri ln nh
sau:
IA = IB, IA > IO, v IB > IO.
ng tri: Cc th d hp t biu hin cc kiu hnh ca c hai dng ng hp t tri v ln. V d, cc
alen IA v IB l ng tri v cc c th IAIB mang nhm mu khc vi hai nhm kia.
Tng tc gen: Cc gen khng alen c th hot ng khng c lp, m ph thuc vo nhau xc
nh tnh trng ca c th. C nhiu kiu tng tc nh: b tr, c ch, a phn, v.v...
Alen gy cht: Khi biu hin lm c th b cht, c c ln v tri: chng ma git Huntington ngi
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l v d v gen gy cht tri ch biu hin tui trng thnh.

Mc ngoi hin (penetrance): T l m mt gen tri hoc mt gen ln c biu hin kiu hnh ca n
cc c th c gi l mc ngoi hin ca gen.
biu hin (expressivity): l mc biu hin kiu hnh ca mt gen hoc kiu gen ngoi hin.
biu hin c th nh, trung bnh hoc mnh.
Tnh trng s lng: Cc tnh trng m Mendel nghin cu u l cc tnh trng khng lin tc. Nhng
trong thc t c nhiu tnh trng biu hin tnh lin tc r rt nh chiu cao, mu mt hoc mu tc ngi,
chiu cao, sn lng ca cc cy trng. Cc tnh trng ny thay i v s lng v c gi l cc tnh trng
s lng, hoc cc tnh trng lin tc. c im nh sau:
Cc c th F1 nhn c t php lai hai dng cha m thun chng khc bit nhau r rt v kiu hnh
trung gian nm gia hai kiu hnh cha m. Cc c th F2 c bin d nhiu hn so vi F1.
Nh vy, c s ca cc tnh trng s lng l hng lot gen gi l nhng gen phc (multiple gen),
hoc cc dy a gen (polygenic series). C th c nhng alen ng gp hiu qu v c nhng alen khng
ng gp trong dy a gen, hot ng theo cch: khi s alen ng gp tng ln th hiu qu kiu hnh tng
ln. Ty thuc vo s lng gen c trong dy a gen m cc nhm kiu hnh c th gin on hoc lin tc.
6.2. Phn tch di truyn sinh vt lng bi

S t hp c lp ca cc gen trong qu trnh gim phn l mt trong nhng nguyn l c bn ca di
truyn hc. Tuy nhin, c nhiu gen thng di truyn cng vi nhau v cng nm trn mt nhim sc th; cc
gen nh th gi l cc gen lin kt v c coi rng chng thuc v mt nhm lin kt. Bng cc php lai
phn tch c th xc nh c s lin kt ca cc gen.
6.2.1. Lai phn tch v lin kt gen
T hp c lp gen chng ta nghin cu trn. Trong php lai phn tch AaBb aabb th t l kiu
hnh th h sau s l 1AB : 1Ab : 1 aB : 1aa. Lai phn tch l php lai hu ch xc nh s lin kt hay
khng gia hai gen, v nu c s sai lch ng k so vi t l 1 : 1 : 1 : 1 do c qu nhiu kiu gen cha m v
c qu t cc kiu gen ti t hp th hai gen ang kho st c coi l lin kt vi nhau, nh minh ho di
y:
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Trong s trn, cc nhm ti t hp ch c th sinh ra do trao i cho gia hai gen trong qu trnh
gim phn ca c th F1 d hp t kp. V trao i cho l s kin ngu nhin din ra dc theo nhim sc th
nn tn s trao i cho, tc l t l cc c th ti t hp trong tng s th h con, lin quan trc tip n 2
gen kho st. Hai gen cng xa nhau th tn s trao i cho cng ln. Do vy, c th dng tn s ny lp
bn di truyn.
T.H. Morgan l ngi u tin nghin cu trn i tng rui gim Drosophila melanogaster tm ra
nhiu gen khng phn ly c lp vi cc gen khc trong gim phn. Trn thc t, mt s gen kt thnh
nhm trong cc th nghim phn tch lai nh ni trn. ng xc nh c 4 nhm lin kt rui gim
ph hp vi 4 nhim sc th c trong b n bi ca sinh vt ny.
6.2.2. Nhim sc th gii tnh v lin kt gii tnh
B nhim sc th ca cc t bo ng vt bao gm cc nhim sc th thng v cc nhim sc th gii
tnh. Trong phn ln cc trng hp, c th c c hai nhim sc th khc nhau k hiu l XY, cn c th ci
mang hai nhim sc th gii tnh ging nhau, k hiu l XX. X v Y l hai nhim sc th khng tng ng,
mt s gen bn X khng c alen tng ng bn Y, ngc li, mt s gen bn Y li khng c alen tng
ng bn X. Nhng gen cn li th c c bn X v bn Y. Hnh 3.24 l s cu trc nhim sc th gii
tnh ngi v Melandrium.
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Hnh 3.24. S cu trc nhim sc th gii tnh ngi v Melandrium
S di truyn ca nhng tnh trng m gen quy nh chng nm trn cc nhim sc th gii tnh gi l s
di truyn lin kt vi gii tnh, mc d cc tnh trng c th khng lin quan vi gii tnh. Tu thuc vo
v tr ca gen nm phn no trong ba phn ni trn ca nhim sc th gii tnh m tnh cht lin kt vi gii
tnh c th khc nhau.
6.2.3. Ti t hp v trao i cho
6.2.3.1. Ti t hp
Khi cc gen lin kt khng hon ton xut hin cc giao t dng mi khng ging ca cha m, c s
sp xp li cc gen, gi l ti t hp (recombination) v cc dng mi xut hin gi l dng ti t hp
(recombinant). nh gi mc lin kt, Morgan et al. a ra khi nim tn s ti t hp, l phn trm c
th ti t hp so vi tng s c th trong th nghim. V d, trong th nghim lai rui gim, trong tng s
1000 c th c 170 c th dng ti t hp, tc 17% (1% l n v o ti t hp). Tn s ti t hp c tnh
theo cng thc:
ti t hp
Tn s ti t hp gia 2 gen kh n nh t ln lai ny qua ln lai khc, khng ph thuc vo s sp xp,
nh hai alen tri cng pha trn 1 nhim sc th (gi l v tr cis) AB/ab hay cho nhau trn hai nhim sc th
(trans) Ab/aB
hoc
Tuy nhin, tn s ti t hp gia cc cp gen khc nhau th thay i ng k.

V d, trong lai phn tch gia 2 gen pr (purple - mt thm) v vg (vestigial - cnh ngn), s nhn
c kt qu:
Pr vg / pr vg
165
Kiu cha m
+ + / pr
191
(khng ti t hp)
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vg
pr + / pr vg
23
+ vg / pr vg
21
Kiu bin th
(ti t hp)
400
V c th s dng phn trm ti t hp xc nh khong cch theo ng thng gia hai gen trn bn
di truyn, hay cn gi l bn lin kt gen.
6.2.3.2. Trao i cho
Hin tng ti t hp chnh l kt qu ca qu trnh trao i cho (crossing over), m trong hai
nhim sc th tng ng hon v hay trao i cho cho nhau nhng im nht nh.
Trong gim phn mi nhim sc th i c hai nhim sc th ch em tng t nhau. Cc nhim sc th
tng ng bt cp hay tip hp vi nhau v trao i cho xy ra gia cc chromatid khng ch em. S
sau m t trao i cho gia 2 gen A v B, gia chromatid 2 v 3.
Hnh 3.25 . Trao i cho gia cc chromatid
Cc nhim sc th t cc chromatid 1 v 4 khng c trao i, nn chng c gi l kiu cha m

(parental) v cc giao t nhn chng cng c gi l cc kiu cha m. Cn hai chromatid khc c xy ra
trao i cho gi l kiu ti t hp v giao t ti t hp.
V d: Cha m cis: AB / ab
Cc giao t: cha m:
AB v ab
con ti t hp:
Ab v aB
Cha m trans: Ab / aB
Cc giao t: cha m: Ab v aB
con ti t hp:
AB v ab.
6.2.3.3. Cc tn s hnh cho (chiasma)

Mt cp nhim sc th tng ng bt cp c 4 chromatid nn gi l b bn (tetrade). Trong mi b bn
c t nht mt hnh cho, biu hin nhn thy c ca trao i cho.
Khi mt hnh cho xut hin gia hai gen nh du, ch mt na sn phm ca phn chia gim nhim l
ti t hp (trong s 4 chromatid ch c 2 ti t hp). Tn s tng i ca cc hnh cho gp i tn s tng
i ca cc sn phm ti t hp.
6.2.3.4. a trao i cho (multiple crossing over)
Trao i cho gia hai chromatid c th xy ra nhiu hn mt ln, tc l 2, 3, 4 ln,... (hnh 3.26). Nu
hai trao i cho xy ra trn cng hai chromatid on gia hai gen nh du th sn phm cui cng u c
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kiu cha m (tc khng pht hin c). Kiu trao i cho ny ch c th pht hin c khi s dng
thm mt gen nh du th ba nm gia A v B. Nu xc sut trao i cho gia A v C v gia C v B
tng ng vi x v y th xc sut xy ra trao i cho i bng xy (tch ca hai xc sut).
a
Hnh 3.26. a trao i cho
6.2.3.5. Tng quan ng hnh gia gen v nhim sc th

Tng qut, cc c ch di truyn u gn cht vi cc nhim sc th v cc hot ng ca chng trong
gim phn. Khi hai gen A v B nm trn hai cp nhim sc th khc nhau th chng c kiu t hp t do theo
Mendel (hnh 3.27). Nu chng nm trn 1 nhim sc th, s c s di truyn lin kt hon ton (hnh 3.28)
hay khng hon ton khi c trao i cho (hnh 3.29).
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Hnh 3.27. Hai cp alen trn cc nhim sc th khc nhau
Hnh 3.28. Lin kt c trao i cho
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Hnh 3.29. Lin kt khng c trao i cho
6.2.4. Nguyn tc lp bn di truyn

Bn di truyn l s bao gm ton b nhim sc th n bi ca mt sinh vt v v tr cc gen trn
nhim sc th. Hnh 3.30 gii thiu bn di truyn ng. Trn bn c 10 nhim sc th c nh du
t I n X v khong 100 gen c xc nh v tr. Cc s nm bn tri nhim sc th ch khong cch t
gen tng ng n u trn ca nhim sc th c nh du bng s 0. Hai vic chnh cn xc nh l xc
nh nhm lin kt v xc nh v tr ca cc gen trn nhim sc th.
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6.2.4.1. Xc nh nhm lin kt
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Nhm lin kt l nhm cc gen nm trn cng mt nhim sc th. Xc nh nhm lin kt l cng vic
nh r gen no nm trn nhim sc th no. C th l xc nh xem hai gen no nm trn mt nhim sc
th hay nm trn hai nhim sc th khc nhau. C nh vy, cui cng s xc nh c c bao nhiu nhim
sc th (tc nhm lin kt) trong b gen n bi ca mt sinh vt v nhng gen no nm trn nhim sc th
no.
Bc u tin cn tin hnh l thc hin php lai hai tnh trng nh trong cc th nghim ca Mendel v
Morgan. Nu t l phn ly nhn c F2 l 9:3:3:1 th hai gen ang kho st nm trn 2 nhim sc th khc
nhau. Ngc li, nu t l l 3 : 1, hoc gn ging vi 3 : 1 th c th kt lun 2 gen kho st cng nm
trn mt nhim sc th.
Trong thc t thc nghim, thay v cc php lai nu trn, cc nh nghin cu thng dng php lai phn
tch, nh c m t trn thc hin mc ch ra. Cc c th F1 nhn c t php lai hai tnh
em lai vi c th ng hp t ln v hai gen kho st. Nu th h tip theo thu c t s phn ly kiu
hnh l 1 : 1 : 1 : 1 th hai gen kho st nm trn 2 nhim sc th khc nhau. Cn nu t l phn ly ni trn l
1: 1 th hai gen quan tm nm trn cng mt nhim sc th.
6.2.4.2. Xc nh v tr ca gen trn nhim sc th
C th xc nh c v tr ca cc gen trn nhim sc th nu bit khong cch gia chng. Nh
bit, khong cch gia hai gen t l thun vi tn s trao i cho, v thc t tn s trao i cho c s
dng o khong cch gia cc gen. Mt phn trm trao i cho c coi l mt n v chiu di ca bn
di truyn v c gi l centiMorgan (cM) t lng knh trng i vi T. H. Morgan, ngi chng
minh thuyt di truyn hc nhim sc th ni chung v hon thin phng php lp bn di truyn i vi
cc sinh vt lng bi bc cao ni ring.
Hy lm th nghim xc nh khong cch gia ba gen ca ng. Ngi ta lai hai dng ng khc nhau v
ba cp gen lin kt: mt dng c mm vng (v) v bng (gl), l b ca (sl), cn dng kia c mm xanh (v+) v
m (gl+), l bnh thng (sl+). Cy lai F1 c kiu gen
v c mm xanh v m, l bnh thng. Khi lai phn tch cy lai ny vi cy hon ton ln th FB (th
h nhn c sau lai phn tch) nhn c cc cy c tm kiu hnh ph hp vi s loi giao t c to ra
bi mt c th ba d hp t. Cc loi giao t ca cy lai F1, cc kiu gen ca cc cy FB v t l gia chng
c nu bng 3.6 di y.
Bng 3.6. Kt qu th nghim lai phn tch
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T bng trn cho thy, phn ln s c th (69,6%) c t hp cc tnh trng cha m. Cc c th nhn c
do trao i cho gia cc gen v v gl chim 18,3%
(16,8 + 1,5) tn s trao i cho gia cc gen gl v sl l 12,1 + 1,5 = 13,6%, cn tn s trao i cho gia v
v sl l 16,8 + 12,1 + (2 1,5) = 31,9%. Nh vy, trnh t sp xp ca 3 gen ni trn s l:
Bng cch tin hnh cc php lai ba tnh (lai theo ba im) vi tt c cc gen c trn nhim sc th theo
tng t hp tch hp ba gen mt, ngi ta sp xp c trnh t ca cc gen trn nhim sc th.
6.2.5. Nhiu (interference)
Nhiu l hin tng trao i cho din ra ti mt im trn nhim sc th ngn cn trao i cho din ra
cc im ln cn. Bi vy m kh nng din ra trao i cho kp trong mt on ngn ca nhim sc th s
rt nh. Hiu qu nhiu c o bng t l gia s trao i cho kp quan st c vi s trao i cho kp
tnh ton theo l thuyt.
S liu trn bng 3.6 cho thy, khong cch gia hai gen v v gl l 18,3%, cn khong cch gia gl v sl
l 13,6%. Nu khng c nhiu, ngha l trao i cho din ra trong cc on v gl v gl sl khng nh
hng ln nhau, th tn s trao i cho kp gia cc gen v v sl phi bng tch s ca cc tn s trao i
cho n din ra hai on ni trn (v xc sut din ra cng mt lc hai s kin c lp bng tch s ca
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cc xc sut din ra mi s kin ring l), c ngha l:

18,3 % x 13,6% = 2,5%.
Nhng trong th nghim ch nhn c 1,5% trao i cho kp, thp hn so vi tnh ton 1%, nh vy
y r rng c hin tng nhiu. Nh nu trn, i lng nhiu c o bng t s gia s trao i cho
kp quan st c trong th nghim v s trao i cho kp mong i theo l thuyt. T s ny c gi l
h s trng lp (coincidence) c o bng s thp phn hay phn trm. Trong v d nu trn, i lng
bng 1,5 : 2,5, tc l 0,6 hoc 60%.
Trn cc on ngn ca nhim sc th, ni m mi trao i cho din ra u nh hng n cc trao i
cho cc im ln cn th i lng trng lp s nh hn 1. nh hng ny ch tn ti trong mt khong
cch nht nh, cng xa cng gim dn v cui cng bin mt, khi i lng trng lp bng 1 hay 100%.
6.3. S a dng ca c ch di truyn

Cc nghin cu ca Mendel v Morgan c tin hnh v xc nhn s ng n ca cc quy lut di
truyn cc sinh vt bc cao m trong chu trnh sng c gim phn, chu trnh sng phn ln l lng bi. Xu
hng tin ho ca sinh gii l lng bi ho. Tuy nhin, chim phn ln sinh khi trn Tri t li l cc
loi nm v to, m chu trnh sng dng n bi chim u th.
Cc c ch di truyn lin quan cht ch vi sinh sn m trong , hot ng ca cc nhim sc th ng
vai tr ch yu, ht sc a dng.
6.3.1. Sinh sn v tnh
Sinh sn v tnh l t mt t bo hay nhm t bo m ch qua nguyn phn to ra cc c th con, gi
nguyn cc c tnh di truyn ca c th m ban u c th con. Nguyn phn l c s ca tng trng
cc sinh vt a bo v sinh sn v tnh cc sinh vt ni chung. Sinh sn v tnh c cc sinh vt n bi v
lng bi, l c s n nh b gen qua nhiu th h.
sinh vt a bo, nh nguyn phn m gia tng s lng t bo. ngi, hp t sau nhiu ln phn
chia nguyn nhim hnh thnh c th gm nhiu t t bo, nh vy m cc t bo ca c th u c b nhim
sc th nh nhau tr t bo sinh dc. Sinh sn v tnh c ng dng rng ri trong nhn ging v nui cy
m t bo thc vt v ng vt.
6.3.2. Sinh sn hu tnh
Sinh sn hu tnh l kiu sinh sn kt hp cc t bo sinh dc ca hai c th khc nhau. Sinh sn hu
tnh to ra s a dng di truyn, lm ngun nguyn liu cho tin ho. Mt trong nhng xu hng tin ho ca
sinh gii l sinh sn hu tnh. S a dng ca cc kiu sinh sn hu tnh th hin mt phn cc kiu xc
nh gii tnh. Tin ho to nhiu c ch duy tr s a dng.
6.4. Chu trnh sng hay vng i

Nhiu sinh vt c s thay i th h n bi v lng bi ni tip nhau thnh chu k gi l chu k sng
(hnh 3.31). Vng i in hnh Eukaryota bao gm: th tinh c hp t bo cht v hp nhn; sinh sn v
tnh ca t bo 2n nh nguyn phn; gim phn c gim nhim v ti t hp; cc t bo n sinh sn v tnh
hoc kt hp vi nhau (th tinh).
Nhng thay i trong chu trnh sng c lin quan cht ch vi thay i nhim sc th, c c s vt cht
l cc gen, dn n s a dng cc c ch di truyn lin quan n c im sinh sn ca cc loi.
S a dng cc chu trnh sng cc nhm phn loi khc nhau c th do chim khong thi gian di,
ngn khc nhau ca cc pha n bi v lng bi lun phin.
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Hnh 3.31. Chu trnh sng hay vng i

A. Chu trnh tng qut ; B. Chu trnh ring ca cc sinh vt ch yu:
a) Cc Eukaryota n bo (vi to, vi nm); b) Ru; c) Dng x; d) Thc vt bc cao; e) ng vt bc cao.
6.4.1. Cc Eukaryota n bo
Chu trnh sng ca to (Chlamydomonas reinhardi) c nghin cu. im c bit l cc giai on
n bi v lng bi c th tn ti mt thi gian di.
6.4.2. Cc ng vt bc cao
Cc ng vt bc cao l lng bi. Trong qu trnh sinh sn phn bo gim nhim to cc giao t n
bi, m s hp nht ca chng to thnh hp t. Cc giao t l cc t bo n bi c chuyn ho cho sinh
sn hu tnh. con c, qu trnh sinh tinh (spergetomatogenesis) to ra 4 t bo tinh t t mi t bo lng
bi. con ci, qu trnh sinh trng (oogenesis) ch sn sinh ra mt t bo trng trng thnh.
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6.4.3. Cc thc vt bc cao

Phn ln thc vt a bo c nm giai on trong chu k sinh sng. Ni chung, giai on n bi chim
u th thc vt bc thp, cn giai on lng bi li chim u th thc vt bc cao. Kh nng giao t
c sinh sn ra do mt cy kt hp vi nhau to ra th h con c sc sng v hu th l c tnh chung ca
nhiu loi cy c hoa (u, ng, i mch, u nnh, thuc l, c chua,...). Tuy nhin, mt s loi, t th
phn v th phn cho xy ra ng thi mc khc nhau nh cy bng vi v bo bo (Sorgum) hn 10%.
6.5. Phn tch b gen ngi

y l vn thi s ca khoa hc th gii th k XXI. S phn tch b gen ngi tin nhng bc
rt di v con ngi cng b xc nh c chi tit trnh t 3 t nucleotid ca b gen ngi. Phn tch b
gen ngi tri qua nhiu bc pht trin: lp bn lin kt gen, xy dng bn vt l ca b gen ngi v
cui cng l chng trnh xc nh trnh t nucleotid ca b gen ngi. Phn ny trnh by khi qut chng
trnh xc nh trnh t nucleotid ca b gen ngi, thnh tu khoa hc ni bt u th k XXI m thi.
Gii m chi tit v b gen ca ngi (human genome) l m c t lu ca cc nh sinh hc, nhng l
cng vic c khi lng s tng chng khng th thc hin c. Nh s ra i ca k thut di truyn
cng nhng thnh tu mi nh to c nhim sc th nhn to, hay in di trong trng xung (pulse field
electrophoresis) cho php tch cc on ADN di c triu nucleotid. Cui nm 1989, M chng trnh xc
nh trnh t nucleotid b gen ngi vi kinh ph 3 t USD c chp nhn gi l chng trnh b gen
ngi (Human Genome Project). Chng trnh d kin trong 15 nm:
Nm nm u lp bn gen trn nhim sc th c 50%, xc nh trnh t nucleotid 1% b gen, ci
tin k thut t 5 10 ln.
Nm nm tip theo lp bn gen 100%, xc nh trnh t nucleotid 10% b gen v ci tin k thut 5
- 10 ln na.
Nm nm cui cng, tc l n nm 2003 - 2004, xc nh c trnh t nucleotid ton b b gen
ngi.
Tham gia chng trnh ny c khong 80 phng th nghim ln trn th gii. Bc tin di trong xc
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nh trnh t nucleotid b gen ngi l vic s dng cc EST (expressed sequence tags) chuyn
mARN thnh cc cADN nh khuch i PCR. Nh k thut ny, s biu hin gen cc m c hiu c
ghi nhn. Trnh t cADN c gn vo vector plasmid to dng. S la chn cc dng xc nh trnh
t nucleotid v s so snh tip theo cho php nh gi mc biu hin ca gen cc m khc nhau. D
theo cc EST gip xc nh phn b ca cc gen m ho trn cc nhim sc th, xy dng bn di truyn
v vt l ca b gen ngi v nh r c cc gen gy bnh.
Trong s 87.983 trnh t EST khc nhau, 10.211 trnh t khi nh gi tip l nhng gen bit (cn c
s liu thng k ca database), s cn li l cc gen trc cha bit chc nng. Gn 40% cc gen c
xc nh ngi lin quan n trao i cht c bn, cu trc t bo, ni cn bng (homeostasis) v phn bo,
22% vi ARN v sinh tng hp protein, 12% lin quan n cc tn hiu v lin lc gia cc t bo.
Ngy nay, ngi ta gii m c hn 99% b gen ngi v cho rng s gen ngi ch nm trong
khong 30.000 - 40.000 (khng phi 100.000 nh d on trc y).
Nhng theo a s cc nh gi, trnh t m ho cho gen ch chim 3% ca hn 3 t nucleotid, nn phi
tip tc gii m nhng on khc mi thu hiu c hot ng ton vn ca b gen ch khng phi tng
gen ring l.
Phn tch chc nng b gen
Xc nh trnh t nucleotid ca b gen ch l bc u, bc thch thc tip theo l phi xc nh xem
mi gen tc ng nh th no. Phn tch chc nng b gen (functional genomic analysis) l mc tiu sp ti
ca cc nh khoa hc.
Mc d c nhiu k thut sinh hc phn t c s dng cho phn tch chc nng ca b gen, nhng cc
nh khoa hc cho rng tc hin nay rt chm v khi lng thng tin qu ln, v vy cn c tng cng:
S dng tin sinh: Tin sinh (bioinformatics) l cha kho x l cc s liu phn tch b gen lin
quan gia m ho v hot ng, pht trin, bnh l,... Tip theo l xy dng cc m hnh tng tc ca b
gen.
S dng cc h thng m hnh ng vt nh chut, Drosophila, tuyn trng v nm men. V d,
Caenorhabditis elegans c b gen c xc nh trnh t nucleotid hn 60% v khong 80% ca tt c
cc gen bit c gy bnh ngi c tng ng tuyn trng ny. Tt nhin, vic xc nh mt gen gy
bnh ch c gi tr khi bit c sinh l bnh ca gen m thi.
6.6. Di truyn ngoi nhim sc th

C nhng tnh trng trong sinh gii chu s kim sot ca cc gen ngoi nhim sc th. sinh vt nhn
tht c nhiu tnh trng di truyn theo kiu ngoi nhim sc th, cc gen m ho chng nm ty th v lp
th.
6.6.1. Ty th v lp th
Ty th l thnh phn quan trng ca cc t bo ng vt v thc vt a kh, y l nhng c quan t kh
phc tp c mng kp v c cha cc enzym tham gia vo chui vn chuyn in t (cc cytochrom) do xc
tc cc phn ng to thnh ATP.
Lp th ch c trong cc t bo thc vt, c loi khng mu, c loi c mu. Lc lp l loi lp th c
mu cha cht dip lc c l cy xanh tham gia vo qu trnh quang hp. Trung bnh mi t bo l ca
thc vt bc cao cha khong 40 - 50 lc lp.
Vt cht di truyn c trong ty th v lp th l cc ADN chui kp, trn, mch vng. ADN ny khc vi
ADN nhn, do c th tch c nh ly tm trong gradient t trng CsCl. Ngoi ra, cc c quan ny cn
cha ribosom v tng hp protein.
6.6.1.1. Cc gen ca ty th
ADN ca ty th m ho nhiu thnh phn ca ty th. Hai rARN tm thy trong cc ribosom ty th c
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m ho trong mtADN (ADN ca ty th), trong khi phn ln protein ca ribosom do cc gen trong nhn
quy nh, mtADN cng m ho cho 22-23 loi tARN v 10-12 loi protein c trong thnh phn ca mng
bn trong ty th.
6.6.1.2. H gen ca lp th
nhiu thc vt h gen lp th (cpADN) c hm lng GC khc bit so vi ADN nhn v ADN ty th,
cpADN ln hn mtADN ng vt ti 8-9 ln, tuy nhin c t l vi lng thng tin di truyn hay khng th
cn cha r. Ch c mt s gen lin quan n kiu hnh ca lp th. C nhiu bn sao cpADN trong mt lp
th. V d, trong l mt s thc vt bc cao c ti 6000 phn t cpADN trong mt t bo phn b trong
khong 40 lp th, c 4 n 18 vng nhn (nucleotid) trong mi lp th v mi vng nhn c ti 4 n 8
phn t cpADN.
H gen lp th c cha cc gen m ho rARN lp th, m ho nhiu tARN v c th c mt s protein
ca ribosom cng nh m ho cho ba tiu n v ca enzym lp th a hp phn, cng nh m ha cho mt
s protein mng lp th cn thit cho vn chuyn in t trong qu trnh quang hp.
6.6.2. c im di truyn ngoi nhim sc th
Cc tnh trng do di truyn ngoi nhn quy nh c kiu di truyn ring:
Kt qu lai thun v lai nghch khc nhau: cc con lun c kiu hnh ca m, cn gi l kiu di truyn
theo dng m.
Cc t bin ca gen t bo cht khng th a vo bt c nhm lin kt no.
Cc tnh trng do gen t bo cht quy nh vn s tn ti khi thay th nhn t bo bng mt nhn c
cu trc di truyn khc.
6.6.3. Mt s v d v di truyn ngoi nhn
6.6.3.1. Tnh bt th c bo cht ng
C nhng cy ng mang bao phn rng hoc c nhng ht phn bt th km pht trin do cc nhn t t
bo cht. Nu th phn cho cy b bt th c bng ht phn bnh thng ca cc cy khc th trong phn ln
trng hp c cc cy th h sau bt th c. Khi lp li php lai ny th chng bt th c khng b mt
i m di truyn theo dng m. Chng bt th c k hiu l cytS, cn cc t bo c ht phn hu th bnh
thng th gi l cytN. Tuy nhin, cc gen trong nhn cng nh hng n tc dng ca t bo cht bt th.
T bo cht cytS ch c th to nn ht phn bt th khi trong kiu gen ca cy gen ln rf (khng phc hi)
trng thi ng hp t rfrf. Cn nu gen ny l alen tri Rf (phc hi tnh hu th) th cc cy cytSRfRf v
cytSRfrf c ht phn bnh thng. T , cc cy cytNrfrf, cytNRf- v cytSRf c ht phn hu th, ch
nhng cy cytSrfrf mi c ht phn bt th hon ton. Tt c cc php lai nhiu ln cytSrfrf cytNrfrf
lun cho th h sau mang cc ht phn b bt th hon ton. Ch c php lai cytSrfrf cytSRfRf (hoc
cytNRfRf) mi c th cho cc cy th h sau vi tt c cc ht phn bnh thng mc d t bo cht l cytS.
Rf khng lm bin i cu trc v c tnh ca t bo cht cytS, m ch c ch s biu hin tc dng ca t
bo cht bt th.
6.6.3.2. Tnh thiu nng h hp nm men
Trong nui cy Saccharomyces cerevisieae to nn mt s nm lc nh vi ng knh bng 1/3 hoc
1/2 nm lc bnh thng. y l tnh trng theo di truyn ngoi nhn hay t bo cht, cc t bo b t bin
petite b thiu cc cytochrom b, c, a, +a3, v cytochrom oxydase, y l nhng enzym mng pha trong ty
th. Cc ni petite c th lai vi cc ni bnh thng v bng cch phn tch kiu hnh ca th h con ngi
ta phn bit c ba loi t bin petite, trong c hai loi l di truyn ngoi nhim sc th: t bin petite
trong nhn pet - , t bin petite ngoi nhim sc th mang nhn t [rho -(N)] khc vi th bnh thng
[rho+], v t bin petite c ch ngoi nhim sc th vi nhn t ngoi nhim sc th l [rho -(S)].
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7. T BIN V BIN D
7.1. t bin
7.1.1. Pht sinh t bin
t bin l nhng bin i trong kiu gen hay trong trnh t cc cp base ca ADN sinh vt ni chung.
Ngy nay c mt s phng php thay i h gen ca sinh vt (t bin, ti t hp). V tng th c hai
dng t bin: t bin h gen (genom) v t bin gen.
t bin h gen l lm thay i s genom ca sinh vt (v d, t n bi thnh a bi), c gi tr trong
bin i ging thc vt.
t bin gen l lm thay i cu trc ca gen - thay i trnh t cc cp base trong cu trc ADN ca
sinh vt. t bin ny c hai kiu chnh: t bin im v t bin chuyn (trt) khung. t bin im c
hai kiu chnh l t bin i dch (ng hon - transition) v t bin o dch (d hon - transversion). t
bin trt khung v thc cht l thm vo hoc mt i cc cp base nucleotid lm thay i khung c m ca
ADN. Phn ln nhng bin i ch din ra vi mt cp base ring l th c th d b hi bin. Hu qu ca
t bin ph thuc vo v tr ca nucleotid trong gen, v v vy, khng phi tt c cc t bin u dn n
kiu hnh b bin i ca sinh vt ang nghin cu. S sinh ra t bin gi l pht sinh t bin
(mutagenesis), cn tc nhn gy ra t bin gi l tc nhn t bin.
7.1.2. t bin ngu nhin
t bin pht sinh khng c s tham gia ca tc nhn t bin gi l t bin ngu nhin. C cc bin
i ca cc cp base cng nh cc sai hnh ca nhim sc th u c th xy ra mt cch ngu nhin. V d,
phn t adenin c th tn ti hai dng cu hnh. dng bn vng adenin to nn hai lin kt hydro vi
thymin trn mch ADN b sung, nhng khng th to lin kt hydro vi cytosin. Nhng nu adenin b h
bin lm cho nguyn t hydro chuyn t C6-amin sang 1-N th mi lin kt vi cytosin c th hnh thnh
hai v tr (hnh 3.33).
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Hnh 3.33. Adenin chuyn sang dng him dn n bt cp bt thng adenin-cytosin
Nu din ra s kt cp AC trong khi ADN ang sao chp th sau mt trong hai chui kp ADN s c
cp GC thay cho AT v tr . y l v d v t bin i dch (hnh 3.34). Hu qu ca t bin ny i
vi t bo ph thuc vo v tr ca cp base b thay th trong gen.
7.1.3. t bin gy to
Cc t bin c sinh ra bi cc tc nhn t bin vt l hay ho hc c gi l t bin gy to. Cc
tc nhn t bin vt l thng s dng l tia X, tia gamma v nh sng t ngoi. Hu qu ca vic chiu tia
X hoc tia gamma l lm gy nhim sc th dn n nhng sai hnh nhim sc th hoc lm cht t bo. Cn
cc tc nhn ho hc c th tc ng theo mt s c ch khc nhau ph thuc vo c tnh c ch phn ng
ho hc ca chng vi cc base ca ADN.
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Hnh 3.34. t bin ngu nhin do adenin h bin bt cp sai trong sao chp
7.1.3.1. t bin gy to do hydroxylamin
Hydroxylamin (NH2OH) ch tc dng vi cytosin, bin i khng thun nghch cytosin sang dng bt
cp c vi adenin, gy ra t bin t GC AT. Cc t bin do hydroxylamin gy ra khng th hi bin
khi x l vi chnh cht ny, trong khi cc hi bin c th sinh ra do x l 5-bromouracil, 2-aminopurin, acid
nitr, v cc tc nhn ny c th thc hin i dch thun nghch t GC sang AT v ngc li.
7.1.3.2. Cc acridin
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Hnh 3.35. Tc dng gy t bin ca acridin
X l acridin lm mt i hoc thm vo mt cp base trn ADN. Vic ny a n nhng hu qu

nghim trng v trnh t acid amin trong protein, do on gen b bin i m ho s b thay i c bn.
nng tng i thp, acridin xen vo gia cc cp base k cn trn ADN, lm cho khong cch gia hai
si ny di thm 0,68nm, gp i khong cch bnh thng. Nu v tr acridin xen vo nm trn si khun,
th mt base ngu nhin s c xen vo si ang c tng hp v tr i din vi phn t acridin. chu
trnh tip theo mt cp base mi s c b sung vo ADN v tr thnh t bin chn thm (t bin xeninsertion mutation). Nu acridin xen vo trn si ang tng hp s gy ra hi bin (hnh 3.35)
7.1.4. Sa cha t bin
C th sinh vt chu nhiu tc ng t mi trng gy tn hi n ADN v do cn ti nhiu c ch
sa cha v ang hot ng loi b cc t bin trn ADN. Trn m hnh tc dng ca nh sng UV
to thnh dimer thymin-thymin hay cytosin-cytosin ta nghin cu c ch sa cha ca c th. Cc dimer lm
sai lch ADN n mc cc purin trn si i din khng th kt hp c. Nu khng loi b c cc
dimer th t bo c th b cht, hoc nu cc nucleotid sai nh vo v tr i din cc dimer trong qu trnh
sao chp, c th dn n t bin. Hai c ch hot ng y l quang phc hot v sa cha ct b.
7.1.4.1. Quang phc hot
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Hnh 3.36. Quang phc hot i vi dimer thymin do UV to ra
Trong qu trnh quang phc hot cc dimer pyrimidin do nh sng t ngoi to ra trc tip phc hi tr
li trng thi ban u. Phn ng ny xy ra nu nh sng thng chiu vo t bo mi x l t ngoi, vy
nn phn ng c gi l quang phc hot(hnh 3.36). Qu trnh hi bin c xc tc bi enzym
photolyase, n phn ho cc dimer sau khi c hot ho bi photon nh sng c bc sng 320 - 370nm.
Cc enzym ny c th l rt ph bin.
7.1.4.2. Sa cha bng ct b
Cc dimer c ct ri khi ADN nh tc dng ca enzym nuclease v ligase (hnh 3.37). y l sa
cha trong bng ti v khng i hi phi c nh sng. C ch sa cha bng ct b rt phc tp E. coli.
Ni chung c nhng enzym nuclease hiu chnh c th tham gia sa cha.
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Hnh 3.37. Sa cha vi c ch ct b i vi dimer thymin trong ADN
Bc u tin l nhn bit dimer trn ADN, tip theo l s hnh thnh vt t trn si n di tc dng
ca endonuclease hiu chnh. Cc enzym ny thng l cc protein nh, phn t lng khong 30.000D. Tt
c cc enzym c bit cho ti nay u ct ADN v tr st cnh ch hng. u 5 ri ra v tr thnh c
cht ct b cho hot tnh exonuclease 5 sang 3 ca ADN polymerase I. Sau khi on ADN c cha dimer
pyrimidin c ct b, mt on mi c trng hp theo chiu 5 n 3 di tc dng ca ADN
polymerase I. Khe h cn li trn si n ADN c ni li bng enzym ligase. Qu trnh sa cha trong
bng ti l ph bin trong cc sinh vt khc v nhiu enzym lin quan cng c xc nh.
7.1.5. Tch cc th t bin
Cc t bin ngu nhin v k c cc t bin gy to ch xy ra vi mt s nh t bo hoc c th trong
qun th. V vy, mun tch cc th t bin cn phi c nhng phng php thch hp.
7.1.5.1. t bin trng thy
L nhng t bin m kiu hnh c th quan st bng mt thng hoc knh hin vi quang hc. Trng
hp sinh vt lng bi cn phi chuyn sang ng hp t th mi pht hin c t bin.
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7.1.5.2. t bin sinh trng

t bin sinh trng, t bin khuyt dng, t bin c iu kin l nhng t bin cn quan tm.
7.2. Bin d
Bin d l c tnh bin i ca sinh vt th hin kh nng c thm nhng tnh trng mi v mt i
nhng tnh trng c, biu hin tnh a dng mun hnh v ca th gii hu sinh. C di truyn v bin d
u biu hin chc nng v quy lut vn ng ca vt cht di truyn.
7.2.1. Phn loi bin d
C th phn loi bin d theo s nh trn hnh 3.38.
Hnh 3.38. Phn loi bin d
Nguyn tc phn loi bin d y l da trn nguyn nhn gy bin d.

7.2.2. Thng bin (modification)
Thng bin l s biu hin a dng ca cc kiu gen ging nhau trong nhng iu kin mi trng
khc nhau. Ni cch khc, cng mt kiu gen trong nhng iu kin ngoi cnh khc nhau s c nhng kiu
hnh khc nhau, s a dng kiu hnh chnh l thng bin.
Thng bin c khp ni trong sinh gii. V d, ging th Himalaya nui 35oC th lng hon ton l
mu trng, nui 5oC th b lng hon ton l mu en, cn nui 20 - 30oC th c b lng Himalaya in
hnh: thn trng, cn mi, tai, bn chn th en. nm men Saccharomyces cerevisiae t bin t bo di
rpm (rough pseudomycelial) c kiu hnh t bo thay i theo pH mi trng: pH = 7 th t bo rt di,
khi pH = 5 - 6 th t bo hnh elip. Cy t dng trng trn t chua c hoa mu hng nht, cn trn t kim
cho hoa mu xanh lam,...
Khng th dng thng bin lm nguyn liu chn ging. Nhng c th dng s thay i ca mi
trng iu chnh kiu hnh trong mt phm vi nht nh phm vi phn ng ca c th.
7.2.3. Bin d t hp
Bin d t hp l s biu hin kiu hnh ca cc t hp gen v t hp nhim sc th mi hnh thnh trn
c s hai qu trnh:
S phn ly c lp ca cc nhim sc th trong gim phn v s kt hp ngu nhin ca chng khi th
tinh.
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S trao i cho v ti t hp ca cc gen.

Trong kiu bin d ny, bn thn cc gen khng b bin i m ch bin i cc t hp v tnh cht tng
tc ca chng trong h thng kiu gen.
Nm vng quy lut hnh thnh cc t hp gen v t hp nhim sc th mi, nh chn ging c th to ra
cc t hp gen theo mun bng cch tin hnh nhng s lai thch hp. Cng trnh to ra ging la m
chu ng Bezostaia-1 ca P.P. Lukianhenco l mt v d: ging Bezostaia-1 tha k t ging Klein 33 ca
Argentina c tnh trng ngn cy, chn sm, chng g st; k tha t ging Effimovka 17 c tnh chu ng,...
y l ging la m c cng nhn l chu c nhng khc nghit ma ng tt nht th gii.
T bo ngi c 23 cp nhim sc th, khi gim phn s to ra 223 loi giao t. S t hp tng y loi
giao t trong th tinh ngu nhin l 423. Thm vo l cc bin d do ti t hp th con s cc bin thin s
v cng ln.
Bin d t hp gi vai tr quan trng nht trong tin ho v chnh n to nn s a dng di truyn n
mc khng c hai c th hon ton ging nhau. Cng tc lai to ging ch yu to ra nhiu bin d t hp
chn la cc dng ti u nht lm ging.
7.2.4. Bin d t bin
Cch phn loi t bin c tha nhn rng ri l da trn nguyn nhn dn n t bin, ngha l theo
nguyn nhn dn n bin i kiu gen. Nh vy, c 4 loi t bin:
1. t bin gen, cn gi l t bin im, khng nhn thy c qua knh hin vi.
2. t bin nhim sc th bao gm:
Cc bin i bn trong mi nhim sc th, nh mt on, thm on, o on.
Bin i din ra gia cc nhim sc th: mt on ca nhim sc th c th b t ra v ni vo mt
nhim sc th khc.
3. t bin h gen bao gm:
Tng hoc gim s lng h gen (b nhim sc th n bi y ), cn gi l a bi th v n bi
th.
Thay i s lng cc nhim sc th trong b nhim sc th lng bi (lch bi).
4. t bin t bo cht l nhng bin i xy ra trong ADN nm trong t bo cht nh ty th, lp th,
plasmid,...
Cc loi t bin u c cp, nn phn ny ch cp n cc t bin h gen, tc a bi th.
7.3. a bi th
a bi th l s gia tng s lng c b n bi ca cc nhim sc th. Cc t bo hoc sinh vt nu c
ba b nhim sc th n bi, k hiu l 3n v gi l tam bi; nu c bn b nhim sc th n bi, k hiu l
4n v gi l t bi, v.v...
a bi th ho thng lm bin i cc tnh trng ca sinh vt, v v th l ngun bin d quan trng
trong tin ho v trong chn ging, c bit l thc vt. c tnh chung ca cc dng a bi th l c kch
thc t bo ln. Cng vi s tng dung tch chung ca t bo thng din ra hng lot nhng bin i v
sinh l v ho sinh ca chng: tng lng nc, gim p sut thm thu, thay i hm lng ca nhiu cht
nh protein, dip lc t, vitamin v.v... lm thay i tnh chng chu ca cc dng a bi th i vi bin ng
ca cc nhn t ngoi cnh.
Cc dng a bi thng to thnh dy gi l dy a bi. V d, chi la m Triticum bao gm mt s loi
phn thnh ba nhm khc nhau v s lng nhim sc th, cc c tnh cng nh tnh trng ca cy. Nhm
th nht l T. monococcum c trong cc t bo soma, s nhim sc th lng bi bng 14. Nhm th hai l
la m cng T. durum c 28 nhim sc th. Nhm th ba l la m mm T. aestivum c 42 nhim sc th. S
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nhim sc th c bn la m l 7, th la m n ht (nhm u) l dng lng bi (7 2 = 14), la m

cng l cc dng t bi (7 4) = 28, cn la m mm l th lc bi (7 6) = 42. y chnh l dy a bi.
Hin tng ny pht hin thy nhiu thc vt khc v mi dy a bi c t hai thnh phn tr ln.
7.3.1. T a bi
Cc dng a bi xut hin trn c s tng h gen ca mt loi gi l t a bi. Nu k hiu s nhim sc
th c bn l A th A l th n bi, AA l th t lng bi, AAA l th t tam bi, AAAA l th t t bi,
v.v... Cc th t a bi xut hin vi cc hnh thc sinh sn khc nhau. Cc th t a bi cc loi sinh sn
v phi hoc sinh dng l c gi tr nh tnh n nh lu di. Trong sinh sn hu tnh, cc dng t a bi
sinh ra cc c th rt a dng v v th khng n nh.
7.3.2. D a bi
Cc dng a bi xut hin trn c s tng s lng gen thuc cc loi khc nhau gi l cc dng d a
bi. V d, con lai khc loi kt hp c cc h gen A v B th dng d a bi nhn c s l AABB. V
vy, ngi ta cn gi d a bi l a bi lai, c gi tr thc tin ln.
Thng cc dng lai xa l bt th. V d, cy lai ci c - ci bp mang trong mnh hai h gen R (ca ci
c) v B (ca ci bp) vi tng s nhim sc th l 18 (9 ca ci c v 9 ca ci bp). Khi cy lai gim
phn, v cc dng nhim sc th ca mt dng cha m (ci c hoc ci bp) khng c cc dng tng ng
thuc dng cha m kia, cho nn khng hnh thnh c th lng tr m c 18 nhim sc th u dng
ring r, nn cc giao t c sinh ra c s lng nhim sc th rt khc nhau t 0 n 18, v trng thi bt
cn bng, khng c kh nng sng (bt th).
Nhng cy lai c mt s giao t, c c v ci s c 18 nhim sc th (9R + 9B). Khi th phn vi
nhau, cc giao t nh th s to ra hp t mang hai b nhim sc th lng bi ca hai loi. Hp t ny gi
l dng song nh bi (amphidiploid) mang cu trc nhim sc th 18R + 18B v nh th rt hu th. Vic to
ra c cc th song nh bi m ra trin vng t hp c cc ging mi bng cch lai xa. Cy lai
Raphanobrassia gia ci c Raphanus sativus v ci bp Brassia oleracea rt kho v kt hp c cc tnh
trng c ca ci c v ci bp.
7.3.3. a bi th ng vt
ng vt phn tnh rt t thy c a bi th. Cn cc loi ng vt khc a bi th quan st thy
trong thin nhin v to ra c trong thc nghim nh: cc c th a bi th ca tm, triton v oxolotl. Cc
dng t bi ca tm Bombix mori cc con ci hu th, cn cc con c bt th. Nguyn nhn l do cc con
c ng giao t trong gim phn to thnh cc th a tr, v vy cc giao t hnh thnh c s nhim sc th
lch bi khng c kh nng sng. con ci d giao t s tip hp ca cc nhim sc th trong gim phn b
hn ch nn cc th a tr khng hnh thnh c v cc giao t c c to ra c kh nng sng.
nhng ng vt m sinh sn hu tnh b thay th bi trinh sinh th a bi th xy ra gn nh thc
vt. Cc dy a bi quan st thy giun a, giun t, bm v mt s ng vt khc. cc ng vt c v
pht hin thy cc hp t tam bi, nhng chng ch sng c n mt na giai on ca phi v thng
pht hin thy trong cc trng hp sy thai hoc ra cht.
Dng d a bi tm u tin t tm lai khc loi Bombix mori v B. mandarina. to ra dng d t bi
ngi ta dng phng php trinh sinh nhn to. B. L. Austarov chn ra cc dng c c v ci mang
hai b nhim sc th ca mi loi: 2n B. mori v 2n B. mandarina, tc l chn ra c cc dng song nh
bi, hay cn gi l d t bi.
Trong gii ng vt, a bi th tng i ph bin trong cc t bo soma, tc l trong trng hp t bo
sinh sn bng nguyn phn.
7.4. a bi th lch
S thay i nhim sc th ch lin quan n mt cp hoc mt s cp nhim sc th c gi l a bi
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th lch hay lch bi l hoc a nhim (aneuploidy).

7.4.1. Cc dng a bi th lch
7.4.1.1. Th n nhim hay th mt (monosomic)
Cc sinh vt lng bi b mt mt nhim sc th ca mt cp c gi l th n nhim vi cng thc
b gen l 2n - 1. Nhim sc th n c khng c cp tng ng trong tip hp gim phn I nn c th di
chuyn v bt k cc no ca t bo, nhng c th b mt v khng nm nhn no ca t bo con. Th n
nhim c th to hai loi giao t: (n) v (n - 1). thc vt mt mt nhim sc th (2n - 1) c th vn sng
vi mt s bin dng. ng vt, vic mt nguyn mt nhim sc th lm mt cn bng di truyn nn
thng b cht hoc bt th.
7.4.1.2. Th tam nhim hay th ba (trisomic)
Th lng bi nu c d mt nhim sc th c gi l th tam nhim vi cng thc b gen l 2n + 1.
Trong gim phn I, nhim sc th d c th tip hp vi hai nhim sc th kia to thnh b ba (tam tr trivalent). Nu hai nhim sc th v mt cc v nhim sc th l v cc kia s to ra hai loi giao t (n + 1)
v (n) tng ng. Tam nhim c th to ra cc kiu hnh khc nhau ph thuc vo nhim sc th no c b
ba. ngi, tam nhim nhim sc th 21 gy hi chng Down.
7.4.1.3. Th t nhim hay th bn (tetrasomic)
Khi mt cp nhim sc th ca sinh vt lng bi c thm hai nhim sc th th gi l t nhim, vi
cng thc b gen l 2n + 2. Trong gim phn I, 4 nhim sc th tip hp c th to thnh b bn, hay t t
(quadruvalent) v c s phn ly v cc cc nh t t bi.
7.4.1.4. Th tam nhim kp (double trisomic)
Th ny c c khi tam bi hai cp nhim sc th khc nhau, cng thc b gen c k hiu l 2n +
1 + 1.
7.4.1.5. Th v nhim hay th khng (nullosomic)
Sinh vt lng bi mt c hai nhim sc th ca mt cp th c gi l th v nhim, cng thc b gen
l 2n - 2. V nhim thng c hiu qu gy cht. Mt s a bi th thc vt c th sng khi mt mt cp
nhim sc th (v d, dng la m lc bi).
7.4.2. Mt s th a bi lch thc vt
7.4.2.1. Cc th tam nhim Datura stramonium
cy Datura stramonium cng chi c c dc c 2n = 24 nhim sc th pht hin c cc t
bin tam nhim 25 nhim sc th tun t tt c 12 cp nhim sc th, vi s khc nhau ch yu l qu v
gai lin quan n nhim sc th no l d tha.
7.4.2.2. Cc th v nhim la m
La m (Triticum vulgare) c 6n = 42. Mc d v nhim thng gy cht, nhng nh a bi th nn
la m nhn c tt c 21 loi v nhim ca tt c 21 cp nhim sc th.
S nghin cu cc dng v nhim gip hiu c cc nhm lin kt gen v vai tr ca tng cp nhim
sc th.
7.4.3. Cc th a bi lch ngi
ngi nhiu hi chng di truyn do cc th a bi lch lm thay i s lng cp nhim sc th gii
tnh a n cc trng bnh: hi chng Turner (XO), hi chng Klinefelter (XXY), siu n (XXX super
female), siu nam (XYY) v YO khng sng. Mt s bnh do tam nhim cc cp nhim sc th khc ca
b gen ngi ngy nay cng c quan tm.
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Hi chng Turner do Henry Turner miu t ln u tin l do mt mt nhim sc th X (XO) c cng

thc di truyn 2n - 1 = 45. Hi chng c mt s biu hin c trng nh: ln (chiu cao < 1,4m), c quan
sinh dc km pht trin (khng dy th), km thng minh,...
Hi chng Klinefelter c Henry Klinefelter m t nhng nam gii c d mt nhim sc th X
(XXY), cng thc b gen 2n + 1 = 47. Hi chng c cc biu hin c trng nh: bt th, ngn tay chn di,
pht trin ngc, c tnh n v suy gim tr tu.
Ngi siu n: do d mt nhim sc th X (XXX), vi b gen 2n + 1 = 47, thoi ho, suy gim tr tu.
Ngi siu nam c XYY, ngi nam bnh thng, nhng c th gi l siu nam v tnh tnh mnh bo
hn ngi thng. Nm 1965, khi nghin cu trong s nhng ngi phm ti, nhng ngi XYY chim ti
3,5%, nn c lc Y c gi l nhim sc th ti li (criminal chromosome). Thc t nhng ngi ny
mnh bo hn ngi thng nn d phm ti. Tuy nhin, nhng nghin cu gn y ph nh nhn xt trn.
Hi chng Down do Langdon Down pht hin, thng l tam nhim cp nhim sc th th 21 (47,
+21) ngi hoc t ra l do nhim sc th 21 ni vi mt phn nhim sc th 15. T l xut hin 1/700 tr
s sinh. Biu hin bnh c trng: thoi ho tr tu, mt trn, di. C mi lin h thun gia tui ca b m
v s tr sinh ra b mc bnh Down: 1/2500 cc b m 20 - 30 tui, 1/300 cc b m 40 - 45 tui v 1/60
i vi cc b m cao tui hn 45. Tui cha khng nh hng.
Dng tam nhim cp nhim sc th 13 (47, +13) c K. Patau pht hin nm 1960 vi t l
1/20.000 tr s sinh, biu hin bi teo no, mt tr, ic v nhiu d hnh khc ni ngoi tng, phn ln tr
cht 3 thng tui, mt s t sng c n 5 tui.
Th tam nhim cp nhim sc th 18 (47, +18) c J.H. Edwards et al. m t nm 1960, biu hin
mt tr, d hnh tai, thn, u v.v..., xy ra vi t l 1/8.000 tr s sinh, hay gp b m ln tui. Tr b hi
chng ny u cht sm, 90% cht 6 thng u.
ngi, cc n nhim kh thy hn, c th do mang nhiu gen ln nguy him nn trng thi bn hp
t kh sng.
CU HI LNG GI
1. Trnh by thnh phn v cu to ca cht liu di truyn ADN, cu trc nhim sc th Eukaryota v
Prokaryota.
2. Trnh by qu trnh sao chp ADN Eukaryota v Prokaryota.
3. Trnh by chu trnh t bo cc sinh vt Eukaryota.
4. Trnh by cc kiu phn bo.
5. Nu s khc bit ca di truyn nhim sc th v di truyn t bo cht.
6. Trnh by t bin, bin d, cc kiu t bin v bin d.
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Chng 4
NGUN GC S SNG
V A DNG SINH HC
MC TIU
1. K c tn v c im cc gii sinh vt trong h thng phn loi 5 gii ca Whittaker.
2. Trnh by c cc hnh dng thng gp v phn loi vi khun theo phng php nhum Gram.
3. Trnh by c cc c tnh chung ca virus.
4. Trnh by c c tnh ca cc nhm thuc phn gii Protozoa.
5. Trnh by c c tnh ca cc ngnh thuc gii nm.
6.
K c tn cc sinh vt thng k sinh gy bnh ngi ca 2 ngnh giun dp v giun trn
7. Trnh by c c im ca cc lp thuc ngnh chn khp v ngha y hc quan trng ca ngnh.
1. NGUN GC S SNG
1.1. S hnh thnh Tri t v kh quyn
V tr c l c hng chc t nm. Mt Tri v cc hnh tinh, trong c Tri t c hnh thnh
cch y khong 4,5 n 5 t nm. S ngng kt bi v tr to thnh Tri t vi cc lp khc nhau. Ban
u, Tri t c nhit rt cao v th lng. St v niken chuyn vo pha tm, nhng cht nh hn gn
b mt. Tuy nhin, ban u trng hp dn ca Tri t cn rt b cho nn cc kh cui cng thot vo
khong khng v tr v ch cn mt qu cu , khng c bin, cng khng c kh.
Nhng ri cng vi thi gian, sc ht ca trng lc cng vi s phn r phng x to nn mt lng
nhit ln v trong lng Tri t tr nn nng chy. Do to nn mt li st v niken, lp v dy khong
4.700km gm silicat nng ca st, magie v mt lp v ngoi dy t 8 - 65km gm ch yu l cc silicat nh.
Do nhit trong lng t tng ln lm thot ra ngoi cc kh khc nhau t hot ng ca ni la. Nhng kh
ny to thnh kh quyn ca Tri t.
Kh quyn ban u khng c oxy t do. C hai m hnh v thnh phn kh quyn ban u v u ph
hp vi gi thit hin nay v ngun gc v s sng.
Theo Oparin, kh quyn ban u rt t i, ch yu l H2, mt t nit dng NH3, oxy dng hi nc v
carbon c th dng CH4.
M hnh th hai cho rng kh quyn ban u do ni la phun ra nh H2O, CO, CO2, N2, H2S v H2.
Hn hp kh d hnh thnh nn hydro cyanid (HCN). Nh vy, c hai m hnh u cho rng kh quyn
ban u khng cha oxy t do.
Thot u, nc ca kh quyn dng hi, ri ngng t li to nn nhng trn ma ln. Sng sui hnh
thnh, ho tan v cun theo cc mui khong (nh st v urani) tch t dn trong bin c. Kh trong kh
quyn cng c th b ho tan trong nc ma vo i dng. Oxy t do c th c hnh thnh trong kh
quyn do s ho tan cc ion t do trong nc bin.
1.2. S hnh thnh cc phn t hu c nh b n t bo
file://C:\Windows\Temp\nyfwwpmgvd\Chapter4.htm
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Phng x tia cc tm, chp v nhit cung cp nng lng lin kt tng hp nn cc cht hu c
n gin. Thc nghim ca Stanley L. Miller xc nh gi thit tng bc pht sinh s sng trn Tri t
ca Oparin l ng. Miller thc nghim vi cc kh c th c trong kh quyn ban u ca Tri t vi
lng kh ln. Dng nng lng bng cch phng tia cc tm tc ng ln hn hp kh amoniac, metan,
hydro v nc trong mt thit b hon ton kn, trong thi gian mt tun tng hp c nhng cht hu
c quan trng nh cc acid amin; cc cht khc nh ur, hydro cyanid, acid acetic, acid lactic. Ngun nng
lng nh th trong kh quyn ban u khng thiu cho nn nhng cht hu c n gin ban u c
tng hp. S tch ly cc phn t hu c trn Tri t m khng b phn hu c th l do cha c cc vi
sinh vt v oxy t do trong kh quyn. Do vy, nhiu hp cht ny c tch ly dn trong bin hng triu
nm m khng b phn hy do oxy ho hay b phn r.
T sau nm 1955 c nhiu nghin cu dng hn hp cc kh c trng ca ni la vi hydro cyanid
t c kt qu dng tnh. ng ch l cc acid amin d dng tng hp c trong thc nghim iu
kin v sinh.
T hnh thnh cc cht trng hp. Cc cht trng hp c hnh thnh nh th no trong cc
i dng c xa cng ch mi dng li cc gi thit. Mt s nh nghin cu cho rng, s phong ph ca
cc cht hu c trong bin l qu thay i cc lin kt gia cc phn t n gin v tri qua hng triu
nm hnh thnh nn nhng lng tng i cc i phn t. Mt s khc li cho rng, cc c ch lm ng
c cng c th lm tng tc ca cc phn ng ho hc l yu t cn thit trng hp cc phn t thnh
cc cht trng hp. Qu trnh hnh thnh cc t hp phn t v cc t bo nguyn thu.
Oparin cho rng, nhng iu kin thch hp ca nhit , thnh phn cc ion, pH, trng thi keo ca
cc phn t a n s hnh thnh nn cc git coaxecva. l nhn ca cc phn t ln c bao quanh
bi mt v bc cc phn t nc. Cc git c cu trc bn trong xc nh v c th hp th vt cht mt
cch chn lc.
Cng nh Oparin, Sydney W. Fox cho rng, cc h thng trc sinh vt (prebionts) dn ti s pht trin
nhng t bo u tin nh nhng tiu cu dng protein mang nhiu tnh cht ca t bo sng. S lng cc
h thng trc sinh vt c th sinh ra bin. Mt s c th cha nhng t hp nguyn liu thch hp v
ln ln v kch thc. Nhng git nh mi cng c th c hnh thnh tng giai on. Bng cch no
cc chui nucleotid trong cc acid nucleic c m ho thnh cc chui acid amin trong phn t protein v
s phin m, dch m c tin hnh. H thng kim tra di truyn to kh nng sao chp v kim tra chnh
xc hn cc phn ng ho hc xy ra bn trong nhng git nc nh . Nhng git vi cc c tnh
ring pht trin thnh nhng acid nucleic trong cc t bo u tin. Nhng t bo u tin l t do v
c kh nng t sao chp. Nhng t bo gi l Prokaryota.
1.3. Tin ho ca s t dng

Nhng c th sng u tin l nhng c th d dng, nhn nng lng t nhng cht dinh dng c sn
trong i dng do cc cht hu c b oxy ho. Khi cc cht dinh dng cn dn gy nn s cnh tranh gia
cc sinh vt tng. S chn lc t nhin to cho bt c bin d no tng kh nng tm kim v ch bin thc n
ca sinh vt. ng thi, cc phn ng sinh ho khc nhau tin trin lm cho sinh vt s dng c cc thc
n khc nhau. Dng u tin ca s chuyn ho ATP rt c th l s ln men.
V cc sinh vt s dng ht cc cht dinh dng t do nn mt s ln trong chng tin ti kh nng
s dng ngun nng lng khc - nng lng Mt Tri. u tin c th l quang phosphoryl ho theo vng
ri sau mi n quang phosphoryl ho khng vng. T thi im ny tr i, s sng trn Tri t ph
thuc vo hot ng ca cc sinh vt t dng quang hp. Kh oxy c thi ra do quang hp lm thay
i kh quyn n mt kh quyn c oxy ho. O2 t do lm cho cc sinh vt c th tin ti s h hp hiu
kh c hiu qu. Oxy cng to nn mt lp ozon tng trn ca kh quyn, ngn cn cc phng x cc tm
xung b mt Tri t, to iu kin cho cc sinh vt chuyn ln i sng cn.
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1.4. S tin ha ca t bo nhn tht

D cho Eukaryota c cu trc phc tp hn v xut hin mun hn nhiu, nhng c hai nhm Eukaryota
v Prokaryota ging nhau bi nhiu hng sinh ho quan trng. Ni bt nht l ng phn (glycolysis) v
phn ng nh sng ca quang hp. V vy, ngi ta a ra thuyt cng sinh. Theo thuyt ny, cc c quan
nh ca t bo nh ty th, lc lp, lng, roi v trung t mi lc, mi ni u l cc c th Prokaryota c
lp m chng b ln t bi cc t bo ln hn v chng tip tc sng st bn trong t bo cht, cc t bo
nh mt thnh phn quan trng ca chng. Cng v th m chng c hot ng ring bit. ng h cho l
thuyt ngi ta tm thy ty th ging vi vi khun hiu kh v kch thc v cu trc, cn lc lp rt ging
vi mt vi loi c th Prokaryota quang hp nh vi khun lam. Chng hn, c ty th v lc lp c ADN
ring ca chng v c dng vng trn ging nh Prokaryota. Thm ch t bo ngi, cc c quan t
c ribosom ring, cc ribosom ny nh hn v hon ton tch bit vi phn cn li ca t bo cht. Cc c
quan t ny sinh ra cc protein ring ca mnh v sinh sn c lp vi t bo ch.
Nhiu c tnh cu trc t bo v sinh hc phn t ch r ty th v lc lp c nhiu nt chung vi sinh
vt khng nhn sng t do v cho rng lc lp c th xut hin t khun lam, cn ty th th t cc vi
khun hiu kh.
Sau y l mt s im vn tt trong gi thit trn:
1. C ty th v lc lp u c cha ribosom ring v nhim sc th ring. Nhim sc th m ho ADN
ribosom v protein ribosom ca chng v cho mt s enzym. C ty th v lc lp u c mng ring.
2. Cc nhim sc th bo quan cng ging vi nhim sc th ca Prokaryota, cun vng v khng c
mng nhn.
3. T chc bn trong ca cc gen bo quan cng ging cc gen ca Prokaryota nhng li rt khc vi
Eukaryota.
4. Ribosom ca ty th v lc lp ging vi ribosom ca Prokaryota hn l ca Eukaryota.
5. Nhiu vi khun quang hp ngy nay sng trong cc vt ch Eukaryota, cung cp cht dinh dng cho
vt ch v li c che ch. Cng vy, mt s vi khun khng quang hp sng cng sinh trong cc c th
Eukaryota, ly nng lng t cht dinh dng m vt ch ca n khng chuyn ho c.
Qua nhng im tm tt trn ta thy vi khun hiu kh b thc bo cch y 1,5 t nm, chu ng s
tiu ho, sng chung bn trong cng vt ch v phn chia ring r. V sau, mt s gen ca vt cng sinh
chuyn vo nhn t bo ch. T y, mi sinh vt khc tin ho. Sau , vi khun quang hp (c th l
Cyanobacteria hay vn c gi l to lam) lm nh vy v to s tin ho cho gii thc vt.
2. A DNG SINH HC
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Hnh 4.1. H thng 5 gii ca Whittaker
2.1. a dng gii

S sng v cng phong ph v c mt mi ni trn Tri t. Chng ta c th tm thy cc sinh vt
sng t hai cc cho n xch o, t y i dng mnh mng cho ti cao hng nghn mt trong khng
trung. T cc vng nc ng bng, cc thung lng kh cn, cc mch nc nng di y bin cho n cc
mch nc ngm su di mt t chng ta u bt gp cc c th sng. Do s phong ph v a dng ca
sinh vt lm cho vic phn chia sinh gii tr thnh mt vn rt phc tp. Mi ti gn y (nm 1753),
nh thc vt hc ngi Thy in Carolus Linneus chia sinh gii thnh hai gii ng vt v thc vt. T
th k XIX, do s pht hin thy nhiu bt hp l ca h thng 2 gii, nhiu nh nghin cu a ra cc
biu phn loi gm 3, 4 hay nhiu hn cc gii khc nhau. Tuy nhin, s phn loi c hu ht cc
nh sinh hc s dng l h thng ca Whittaker a ra nm 1969 (hnh 4.1). H thng gm 5 gii l: vi
khun (Morena), Protista, thc vt (Plantae), nm (Fungi) v gii ng vt (Animalia). i cng vi h thng
phn loi ny l mt s chia cc sinh vt sng thnh 2 nhm ch yu l nhm sinh vt c nhn nguyn
thu (Prokaryota) gm ch yu l vi khun v nhm sinh vt nhn tht (Eukaryota) gm ng vt, thc vt,
nm v n bo. Cc nghin cu gn y a ra s sinh gii gm 3 nhm (domain) l Archaea,
Bacteria v Eukaryota (hnh 4.2).
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Hnh 4.2. Ba nhm (domain) ca sinh gii
2.2. Gii Monera v Virus

2.2.1. Gii Monera
Gii ny gm cc sinh vt tin nhn, ch yu l vi khun, vi s lng c th ln nht trong sinh gii.
Chng tin ha thnh nhiu dng khc nhau v c mt hu nh tt c cc mi trng trn Tri t. Cc
bng chng ha thch cho thy, cc sinh vt tin nhn xut hin cch y 2,5 t nm. Khc vi cc nhm
sinh vt khc, vi khun c t s khc nhau v c im hnh thi, khng thay i nhiu v hnh dng v kch
thc phm vi rng. Phng php phn loi ch yu da trn c im cu trc t bo, c im sinh l,
c im cu trc phn t v s phn ng vi cc loi thuc nhum, t da trn mi quan h tin ha ca
chng. So snh trnh t ARN ribosom cc nh khoa hc thy c 2 nhm vi khun khc bit ln l gii
Eubacteria gi n gin l Bacteria. Nhm ny gm cc vi khun trong cc mi trng bnh thng xung
quanh chng ta. Nhm th 2 l Archaebacteria hay gi n gin l Archaea; nhm ny bao gm cc loi vi
khun c nht ca gii v chng c th thuc mt gii ring.
c im chung ca nhm vi khun l nhng sinh vt n bo, c thnh t bo, khng c dng a bo
v khng c cc bo quan c bao bc bi mng. Hu ht cc sinh vt tin nhn c kch thc nh hn sinh
vt nhn tht b nht.
2.2.1.1. Nhm vi khun c (Archaeabacteria)
L nhm cc vi khun sng cc mi trng khc nghit, khng bnh thng. Hin nay, nhiu nh khoa
hc xp vi khun c thnh mt gii ring do chng rt khc vi cc vi khun khc. Thnh, mng t bo v
ARN ribosom ca vi khun c khc vi vi khun tht. thnh t bo vi khun c khng c peptidoglycan
nh vi khun tht. Nhm vi khun c c th sng nhng ni m khng mt sinh vt no khc c th tn ti
nh cc sui acid nng, quanh ming ni la hay cc vng nc cc mn,... cng l nguyn nhn ti sao
qua mt thi gian di chng mi c pht hin bi mi trng nui cy v phn lp chng rt kht khe. Cc
nh khoa hc cho rng mi trng khc nghit m vi khun c ang sng tng t nh cc iu kin trn
Tri t khi cc dng sng u tin xut hin v bt u tin ha.
Da vo mi trng sng ngi ta chia vi khun c thnh 3 nhm sau:
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Nhm sinh metan (Methanogens): sng mi trng khng c oxy v sinh kh metan.
Nhm a nhit v acid (Thermoacidophiles): sng cc sui acid nng (230oF v pH <2), quanh
ming ni la..., iu kin ny c th git cht cc sinh vt khc.
Nhm a mui (Extreme halophiles): gm cc vi khun c sng cc vng nc cc mn nh Bin
Cht (Dead Sea),... Chng c th pht trin cc vng nc c hm lng mui gp 10 ln lng mui
cc vng nc bin bnh thng, ni m c th git cht hu nh tt c cc loi vi khun khc, th i vi
chng mui li l ngun nguyn liu tng hp ATP.
2.2.1.2. Nhm vi khun tht (Eubacteria)
Hu ht cc loi vi khun c xp vo nhm ny. Chng c nhiu hnh dng, kch thc, cc c tnh
sinh ha v di truyn khc nhau. Tip u ng EU c ngha l tht, nn nhm ny bao gm tt c cc vi
khun c bit bng cc phng php truyn thng nh cc loi vi khun gy bnh m chng ta thng
thy. Hnh dng thng gp ca cc t bo vi khun nhm ny l dng hnh cu (Coccus), hnh que
(Bacillus) v hnh xon (Spirillum) (hnh 4.3). Tn ca vi khun thng xut pht t hnh dng ca chng.
Hnh 4.3. Cc hnh dng thng gp ca t bo vi khun.
Theo mi quan h tin ha Eubacteria c th chia thnh 12 ngnh khc nhau. Trong phm vi cun sch
ny, cc tc gi ch trnh by nhng nt chnh ca 4 ngnh c a s cc nh khoa hc tha nhn. Cc
ngnh l:
Vi khun lam (Cyanobacteria).
Xon khun (Spirochetes).
Vi khun Gram-dng (Gram-positive).
Proteobacteria.
a) Phn loi theo phng php nhum Gram
Hu ht cc loi ca nhm vi khun tht c th c chia thnh 2 nhm gia trn s p ng ca chng
i vi mt k thut phng th nghim gi l phng php nhum Gram. Bng phng php nhum ny cc
vi khun tht c chia thnh 2 nhm l vi khun Gram (-) v nhm vi khun Gram (+). Tn ca cc nhm
ny xut pht t s p ng khc nhau i vi thuc nhum. y l phng php do nh vi sinh hc ngi
an Mch Hans Christian Gram xut nm 1884, v l phng php phn loi vi khun c ng dng
rng ri trong y hc cho n ngy nay.
b) c im ca mt s nhm c phn loi da trn mi quan h tin ha
Nhm vi khun lam (Cyanobacteria): l cc vi khun Gram m, c kh nng quang tng hp ging
thc vt v sn phm ph c gii phng l oxy. Chng c kch thc ln hn cc sinh vt tin nhn khc.
Do kh nng to oxy v l ngun thc n ca nhiu sinh vt khc nn chng c vai tr quan trng trong
nhiu h sinh thi. Nhm ny trc y c xp vo nhm to lam (blue-green Algae) do chng pht
trin thnh dng chui ging nh to. Hin nay chng thuc nhm Eubacteria v khng c nhn, c bao
bc bi mng v cc lc lp. Cc vi khun thuc nhm ny c th c cc mu sc nh vng, , nu, xanh l
cy (green), thm ch l mu en v mu lam (blue-green). Khng ging nh cc vi khun tht khc, cc vi
khun lam c bao bc bi mt cht dng gel v lm cho chng c kh nng dnh li vi nhau hnh thnh
cc qun th dng chui. Trong cc dng chui ca vi khun lam, hnh thnh cc t bo c chuyn bit ha
gi l d bo nang (heterocyst) (hnh 4.4), cc d bo nang ny c cha cc enzym c nh nit kh quyn,
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chuyn thnh dng d s dng cho cc loi thc vt.
Hnh 4.4. Hnh thi qun th Anabaena sp.
Nhm xon khun (Spirochetes): l cc vi khun Gram (-), d dng v c dng xon. Mt s l vi
khun hiu kh v s khc l k kh. Chng di chuyn bng cch t xoay quanh mnh nh xoay m nt chai.
Cc vi khun thuc nhm ny c th sng t do, cng sinh hay k sinh. Mt loi thuc nhm ny c
con ngi nghin cu kh k l Treponema pallidum, l tc nhn gy bnh giang mai (Syphilis) ngi v
lan truyn qua ng tnh dc.
Nhm vi khun Gram dng (Gram-Positive Bacteria): mc d tn ngnh trng vi tn ca nhm vi
khun gram dng (theo cch phn loi ca phng php nhum Gram) nhng khng phi tt c cc vi
khun ca nhm ny u l vi khun gram dng. Mt s t loi ca nhm vi khun Gram (-) cng c xp
trong ngnh ny, do chng c nhng c im sinh hc phn t tng t vi cc vi khun Gram dng. in
hnh thng gp l cc loi lin cu khun (Streptococcus) gy cc bnh hu v hng. Sa chuyn thnh
sa chua (yogurt) khi c mt trc khun Gram dng (Gram-positive Bacilli) no pht trin trong sa v
to acid lactic. Cc trc khun Gram-dng cng tm thy khoang ming, ng rut, ni chng cn tr s
pht trin ca cc vi khun gy bnh. Vi khun sinh acid lactic (Lactobacilli) l cc trc khun Gram dng
c tm thy rng. Chng l cn nguyn gy hng men rng do gii phng acid lactic. Mt nhm vi
khun in hnh khc ca ngnh ny l cc x khun (Actinomycetes). Chng l cc vi khun Gram dng,
c th c dng si, phn nhnh, pht trin nhiu trong t. Cc vi khun ny c kh nng sinh cc loi khng
sinh c tc dng c ch hoc git cht cc vi sinh vt khc.
Nhm vi khun Proteobacteria: y l ngnh ln nht v a dng nht trong nhm vi khun tht.
Chng c chia thnh cc ngnh ph: vi khun ng rut (enteric bacteria), vi khun ha t dng v vi
khun c nh nit (nitrogen-fixing bacteria).
Nhm ph vi khun ng rut: bao gm cc vi khun d dng Gram (-), sng hiu kh hoc k kh
rut. Loi in hnh v c bit kh r l Escherichia coli (E. coli), vi khun ny sng rut ngi.
Ti y, chng sinh vitamin K tr gip cc enzym tiu ha thc n. Mt s cc vi khun khc c kh nng
gy bnh nh Salmonella gy bnh ng c thc phm.
Nhm ph vi khun ha t dng: l cc vi khun Gram (), chng c th ly nng lng t cc cht
khong bng cch oxy ha cc cht ha hc c trong cc khong ny. Mt in hnh l cc vi khun oxy ha
st, chng sng cc ao, h nc ngt c hm lng mui st cao, oxy ha ion st trong mui thu nng
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lng.
Nhm ph vi khun c nh nit: in hnh l chi Rhizobium, y l cc vi khun Gram (-) sng cng
sinh cc nt sn cy h u, nh cy u tng, u nnh,... Do lng nit t do (N2) trong kh quyn
chim khong 80% nhng ng, thc vt khng s dng c trng thi kh ny, m phi nh vo cc vi
khun c nh nit chuyn trng thi kh thnh dng hp cht nh NH3 v chng ly nin dng hp cht
tng hp thnh cc hp cht nh protein, acid nucleic. Khng mt gii no khc c th c nh nit kh
quyn thnh dng hp cht v nhm Rhizobium l nhm thit yu ca chu trnh nit. Khng c chng v cc
tc nhn c nh nit khc th a dng s sng trn hnh tinh chng ta c th khng tn ti.
c) Mt s nt sinh hc c bn ca vi khun
Cu to:
T bo vi khun c cu to in hnh, gm 3 phn l thnh t bo (cell wall), mng t bo (cell
membrane) v t bo cht (cytoplasm). Mt s vi khun c nt c trng ring nh hnh thnh dng ni bo
t (endospore), lp v bc bn ngoi (capsule) v cc mng ngoi (outer membranes). S phong ph trong
cu to ca vi khun l do s thch ng vi cc iu kin mi trng khc nhau ca chng.
Thnh t bo: Tr mt s t ngoi l, c vi khun tht v vi khun c u c thnh t bo. Khng ging
vi thnh t bo thc vt, cc vi khun tht c thnh t bo c cu to bi peptidoglycan. Thnh t bo vi
khun c c cu to bi cc hp cht khc khng phi peptidoglycan. thnh t bo vi khun Gram (-) c
mt lp mng ngoi, l mt lp ca phc hp lipid v ng. Mng ngoi c tc dng bo v vi khun
chng li mt s khng sinh xm nhp vo t bo. Chnh v vy nhiu khng sinh khng c hiu qu khi iu
tr cc bnh do vi khun Gram (-) gy ra.
Mng t bo v t bo cht: Mng t bo vi khun c cu to bi mt lp mng lipid kp tng t
mng t bo sinh vt nhn tht. mng c cha cc enzym thc hin cc phn ng h hp t bo do vi
khun khng c ty th. Cc mng t bo ca vi khun quang tng hp c cc np gp bn trong c gi l
thylakoid ging vi thylakoid trong lc lp. Cc thylakoid vi khun c cha cc cht mu quang hp v thc
hin chc nng quang hp. Cc t bo vi khun khng c cc bo quan c bao bc bi mng. T bo cht
ca vi khun l mt dung dch c, dnh ca cc ribosom v ADN. ADN vi khun c sp xp thnh mt
vng trn kn. Mt s vi khun cn c cc plasmid, l cc ADN vng, nh c kh nng t sao chp trong t
bo cht ca chng.
Lp v ngoi v khun mao (capsule & pili): Nhiu loi vi khun to ra mt lp v bao bc ngoi c
bn cht polysacarid, lp v ny dnh cht vo b mt t bo v bo v t bo chng li cc iu kin bt li
nh kh nng, cc cht ha hc c hi vi t bo vi khun cng nh s tiu dit ca bch cu k ch. Khi
lp v c cu to bi mt lp v xp ca cc ng dnh c gi l glycocalyx. Glycocalyx ny gip vi
khun bm vo cc m v t bo vt ch. Cc khun mao (lng nh) ging nh tc, ngn, c bn cht
protein, c trn b mt ca mt s t bo vi khun. Cc lng nh ny gip vi khun bm vo t bo vt ch
v chuyn cc vt liu di truyn t vi khun ny sang vi khun khc.
Ni bo t (endospore): Mt s vi khun c kh nng hnh thnh cc loi t bo c bit khi gp cc
iu kin sng bt li gi l ni bo t, c cu to bi mt lp v ngoi dy bao bc ly ADN ca vi
khun. Trong khi cc t bo vi khun bnh thng c th b cht th dng ni bo t ca chng vn tn ti khi
gp cc iu kin bt li. Chng gip vi khun chu ng c nhit cao, cc cht ha hc c hi, phng
x, kh v cc iu kin bt li khc.
Cc cu trc gip vi khun chuyn ng: Nhiu vi khun s dng cc cu trc ging roi (flagellum)
di chuyn. Cc roi ny c dng di chuyn cc t bo vi khun trong nc. Vi vi khun khng c roi c
cc phng php di chuyn khc. V d: Vi khun nhm Myxobacteria to ra mt lp nhy v sau di
chuyn nh nhng qua n. Cc xon khun di chuyn theo kiu xoay trn ging nh xoay m nt chai.
Dinh dng v pht trin:
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C hai khc bit c bn gia cc nhm vi khun l ngun nng lng ca chng v s dng hay khng
s dng oxy trong h hp t bo. Hu ht cc vi khun l cc sinh vt d dng. Chng ly nng lng bng
cch s dng cc vt liu hu c nh mt ngun nng lng. Vi khun s dng cc vt liu cht, su mc
gi l cc vi khun hoi sinh. Mt s vi khun khc l cc sinh vt t dng, chng c c ngun nng
lng bng cch to ra thc n ring cho mnh t nh sng Mt Tri hoc cc cht khong. Cc vi khun
quang t dng l cc vi khun s dng nh sng Mt Tri nh mt ngun nng lng, trong khi cc vi
khun ha t dng c c thc n ring cho chng bng cch oxy ha cc hp cht v c thay cho nh
sng Mt Tri. S khc bit v chuyn ha quan trng th 2 gia cc nhm vi khun l vn s dng oxy
trong h hp t bo. Vi khun no s dng oxy trong qu trnh h hp t bo c gi l cc vi khun hiu
kh (aerobe), vi khun khng s dng oxy c gi l vi khun k kh (anaerobe), in hnh l vi khun ly
nng lng cho qu trnh h hp t bo thng qua s ln men k kh. S c mt hay khng c mt oxy trong
mi trng sng ca chng l iu kin quyt nh s sng ca mt s vi khun: cc vi khun khng th
sng khi khng c oxy gi l vi khun hiu kh bt buc v vi khun khng th sng khi c mt oxy gi l
cc vi khun k kh bt buc. Mt s vi khun khc c gi l vi khun k kh ty tin, chng c th s
dng oxy khi thun li hay khng s dng ty thuc vo oxy mi trng. Cc vi khun c yu cu v nhit
cho s pht trin vi mt phm vi kh rng: mt s pht trin tt nhit thp 0 - 20oC, s khc pht
trin tt nhit 20 - 40oC. Mt s vi khun a nhit pht trin tt 40 - 110oC. Hu ht vi khun pht
trin tt pH 6,5 - 7,5.
Sinh sn v ti t hp gen: Hu ht vi khun sinh sn bng qu trnh phn i (trc phn) (hnh 4.5), l
qu trnh trong nhim sc th (chromosome) c sao chp, sau t bo s phn i. Qu trnh phn i
ny l mt dng ca sinh sn v tnh.
Hnh 4.5. Trc phn t bo vi khun E. coli (Dennis Kunkel)
iu kin l tng, cc vi khun phn chia nhanh cho s lng gp i ch trong vng 20 pht. Tt c
cc t bo vi khun u l th n bi v ADN ca hu ht vi khun l mt si xon kp duy nht, gn vo
mng t bo v ch nhn ln trc khi t bo phn chia. Mt s loi vi khun cn c cc vng ADN nh hn
c gi l cc plasmid, chng c s dng trong cng ngh gen mang cc gen mi vo cc sinh vt
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khc. Cc vi khun c th vn chuyn cc gen bng mt trong ba cch l tip hp (conjungation), bin
np (transformation) v ti np (transduction).
2.2.1.3. Vi khun v con ngi
a) Cc vi khun c hi
Mt s vi khun c hi gy ra cc bnh ngi. Chng ta c mt mn khoa hc chuyn nghin cu
v cc bnh do cc vi sinh vt ni chung v vi khun ni ring gy ra l mn Bnh hc (Pathology). Mt
s bnh do vi khun gy ra c tm tt nh trn bng 4.1.
Bng 4.1. Tm tt mt s bnh do vi khun gy ra
Bnh
Tc nhn gy bnh
C quan b bnh
Phng thc ly nhim
Clostridium botulinum
Thn kinh
Thc phm bo qun khng

hp l
Bnh t (Cholera)
Vibrio cholerae
ng rut
Nc b ly nhim
Su rng (tooth decay)
Streptococcus mutans,
S. sanguis, S. salivarius
Rng
Vi khun t mi trng
vo ming
Bnh lu (Gonorrhoea)
Neisseria gonorrhoeae
Niu o, vi Fallope,
mo tinh hon
Qua ng sinh dc
Bnh lyme (Lyme disease)
Borrelia burgdorferi
Da, khp, tim
Ve t
St chm ni Rocky (Rocky

mountain spotted fever)
Rickettsia ricketsii
Mu, da
Ve t
Ng c thc phm do
Salmonella
Salmonella
ng rut
Thc phm v nc b
ly nhim
Vim hng do lin cu (Strep

throat)
Streptococcus pyogenes
ng h hp trn,
mu, da
Ht hi, ho, tip xc

trc tip
Bnh un vn (Tetanus)
Clostridium tetani
Synap thn kinh
Vt thng b nhim
Bnh lao (Tuberculosis)
Mycobacterium
tuberculosis
Phi, xng v cc
c quan khc
Ho
Bnh ng c tht
(Botulism)
Mt s vi khun gy bnh bng cch sinh cc c t v cc c t ny gm hai loi: ngoi c t

(exotoxin) l cc c t c cu to t protein, do cc vi khun Gram(+) tit ra mi trng bao quanh
chng. Bnh un vn l bnh gy ra bi ngoi c t. Loi c t th hai l ni c t (endotoxin) cu to
bi lipid v hydratcarbon, cc c t ny c lin quan ti mng ngoi ca vi khun Gram (-) nh E. coli. Ni
c t khng gii phng cho n khi vi khun cht. Ni c t gy st, au c, kit sc v ph hy cc mch
mu ca h thng tun hon.Cc vi khun cng gy bnh bng cch ph hy cc m c th nh cc loi lin
cu khun (Streptococcus). Khi cc vi khun bm cht vo cc t bo, chng tit ra cc enzym tiu ha gip
chng xm nhp su vo cc m.
b) Cc vi khun c li
Bn cnh nhng tc hi vi khun cng mang n cho con ngi nhiu li ch c trong t nhin cng nh
cc ng dng trong cuc sng hng ngy: nhiu khng sinh c ngun gc t vi khun, ng dng vi khun
trong x l nc thi. Vi khun l tc nhn tham gia phn hy xc ng, thc vt cht mt mt xch quan
trng trong chu trnh vt cht ca t nhin. Chng ng vai tr nh nhng tc nhn ti ch hon tr li cc
cht dinh dng cho mi trng. Mt s vi khun c ng dng trong sn xut thc phm, nh vi khun
gip chng ta lm nc sa (buttermilk), kem chua (sour cream), sa chua (yogurt), phomt gy (cottage
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cheese), da ci bp (saucerkraut) v cc loi chua (pickles). Vi khun cng c ng dng trong sn

xut ha cht cng nghip, nh sn xut cc nhin liu v hp cht hu c. Cc vi khun cn c s dng
tr gip lm sch cc thm ha mi trng do con ngi gy ra nh cc s c trn du v ha cht.
2.2.2. Virus
H thng phn loi 5 gii khng phn loi virus do chng khng c xem nh mt sinh vt. Virus
khng sinh sn c lp cng khng s dng nng lng. Ngy nay, ngi ta nh ngha virus l nhng thc
th, sng trong mt vt ch nh nhng k sinh ni bo bt buc, gy bnh v c kch thc siu nh.
2.2.2.1. Cc c tnh chung ca virus
Qua mt s lng ln cc loi virus m con ngi bit cho thy chng c s a dng v hnh thi v
kh nng xm nhim cc loi sinh vt khc nhau. Mc d c s a dng v hnh dng v cu trc, nhng tt
c cc loi virus c mt s c im chung sau:
L nhng thc th gy bnh v c kch thc siu nh.
Khng c t chc t bo v cng khng c b my chuyn ha ring.
C cu trc n gin vi thnh phn c bn l acid nucleic c bao bc trong mt v bng protein.
Acid nucleic ch gm mt loi, hoc ADN hoc ARN, m khng bao gi c c hai.
L cc k sinh ni bo bt buc do chng b bt hot hon ton khi bn ngoi vt ch.
Nhn ln trong k ch bng cch s dng b my chuyn ha ca t bo k ch.
Hot ng c tnh c hiu, ngha l chng thng xm nhim cc c quan v sinh vt thch hp.
Khng c kh nng pht trin v phn chia c th.
C th c kt tinh v ngay dng ny chng vn gi c kh nng xm nhim.
Khng b tc ng bi cc khng sinh.
C th tri qua qu trnh t bin.
2.2.2.2. Kch thc v hnh dng
Virus l nhng thc th rt nh, nh hn vi khun c kch thc nh nht. Chng ch c nhn thy
qua knh hin vi in t nh cc vt th nh c gi l cc virion. Chng c o bng n v nm (10-9 m).
Ni chung, cc virus c kch thc thay i t 10 - 300nm.
Virus c th c mt s hnh dng chnh nh trn hnh 4.6.
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Hnh 4.6. Mt s hnh dng khc nhau ca virus

+ Hnh khi a din hoc hnh cu nh Adeno virus, Herpes virus, ...
+ Hnh xon c hoc hnh que nh virus khm thuc l, virus di, ...
+ Hnh dng phc tp hoc khng u nh bacteriophage, vaccinia, ...
Vic phn loi virus thng da vo loi acid nucleic v hnh dng v protein ca virus.
2.2.2.3. Cu to ca virus
Virus c cu to bi hai phn chnh: v capsid v acid nucleic (hnh 4.7).
a) V capsid
L lp v bo v bn ngoi, hu ht c cu to bi protein c bit. N bo v acid nucleic trnh khi
tc dng gy bt hot ca enzym nuclease trong mi trng. V ny thng c cu to bi nhiu n v
ging nhau v c gi l capsome. Hnh dng v s sp xp ca cc capsome quyt nh hnh dng ca
virus. Mt s virus rt c bit nh virus cm, virus quai b cn c mt lp v ngoi cng c gi l
envelope, lp v ny c cha mng t bo k ch m virus nhn c trong qu trnh ra khi t bo ch.
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Hnh 4.7. Cu trc chung ca mt virus
b) Acid nucleic
Phn ny nm bn trong v capsid gi l li (genome), khng ging nh cc sinh vt khc, acid
nucleic ca virus ch c mt loi phn t duy nht hoc ADN, hoc ARN m khng bao gi c c hai phn
t ny. Phc hp gm acid nucleic v capsid gi l nucleocapsid. Mt s virus cn c mt lp v bao quanh
capsid c gi l mng bao (envelope).
2.2.2.4. Cc bnh do virus gy ra
Virus gy ra kh nhiu bnh cho tt c cc nhm sinh vt sng. Ring con ngi, chng gy ra mt
s bnh nh cm (flu), cm lnh, herpes, si (measle), bnh thy u (chicken pox), bnh u ma (small
pox), vim no (encephalitis) v c bit l cc virus ang c con ngi rt quan tm nh HIV,
Hantavirus v virus ebola. Cc virus ny xut hin, hoc l chng chuyn i k ch mi (con ngi) hoc
l cc virus m s tn ti v tc ng ca chng n by gi cc nh khoa hc mi nhn ra.
Khng sinh khng c hiu qu chng li virus. Tim chng s l bin php bo v i vi cc c th
cha b ly nhim.
2.3. Gii Protista

Theo phng php phn loi truyn thng, gii Protista gm cc sinh vt nhn tht, n hoc a bo
n gin v c chia thnh 2 phn gii l Protozoa v Algae.
2.3.1. Phn gii ng vt nguyn sinh (Protozoa)
ng vt nguyn sinh l cc c th ch gm mt t bo, thng c kh nng di ng, c kch thc
trung bnh t 50 - 150m. Hnh dng c th hoc khng nht nh nh amip (hnh 4.8), hoc tng i n
nh nh trng roi, trng lng (hnh 4.9 v hnh 4.10). Mt s loi c v cng bao bc nh trng Mt Tri,
trng phng x.
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Hnh 4.8. Hnh thi ca Amoeba sp.
D ch l mt t bo nhng ng vt nguyn sinh l nhng c th c lp, ton vn. Ba thnh phn

chnh ca t bo l mng, t bo cht v nhn, khng c thnh t bo.
amip trn, mng t bo cn dng s khai, c gi l mng sinh cht (plasmalemma). Cc loi trng
roi, trng lng c mng t bo dng pellicula, mng cellulose ch c mt s loi trng roi thc vt.
Hnh 4.9. Hnh thi ca trng roi Euglena sp
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Hnh 4.10. Hnh thi ca trng lng Paramecium sp.
T bo cht thng c chia thnh 2 lp. Lp ngoi mng, qunh v sng hn gi l lp ngoi cht
(ectoplasma hay plasmagel). Lp trong lng, c mu ti hn gi l lp ni cht (endoplasma hay plasmasol).
Trong t bo cht c nhn, cc c quan t v cc ht vt cht khc. S lng nhn thay i tu loi (1, 2
nhn hoc c th nhiu hn).
m bo mi hot ng sng bnh thng, trn c th ng vt nguyn sinh xut hin nhng cu trc
nh b (c chc nng nh nhng c quan ng vt bc cao) gi l c quan t (organelles).
C quan t vn ng: chn gi ( trng chn gi), roi hoc mng un trng roi, tim mao hay lng
nh, mn ( trng lng). C quan t tiu ho v hnh thc dinh dng: chn gi, roi, ngoi chc nng vn
ng cn tham gia nhim v bt mi. S tiu ho xy ra bn trong t bo (tiu ho ni bo) do cc khng bo
tiu ho m nhim. Tu loi m hnh thc dinh dng c th l d dng (kiu dinh dng ng vt), t
dng (kiu dinh dng thc vt i vi nhng loi m bn trong c th c cha ht dip lc), hoc hoi
sinh (dinh dng theo kiu thm thu i vi nhng loi k sinh).
C quan t bi tit v iu ho p sut thm thu do cc khng bo co bp ph trch.
C quan t bo v v tn cng: cc bo gai hay ti lng (trichocyst) trng lng.
C quan t phn x: im mt (stigma) trng roi c kh nng cm quan, trng lng mang si thn
kinh nguyn thu (thc cht l mt h thng mng si cht nguyn sinh) iu ho hot ng nhp nhng ca
cc tim mao.
ng vt nguyn sinh c th c hnh thc sinh sn v tnh, hu tnh hoc xen k gia hai hnh thc sinh
sn ny.
Sinh sn v tnh: mt s loi trng roi, trng lng, hnh thc sinh sn v tnh xy ra mt cch n
gin bng cch phn chia c th ban u thnh 2 c th mi. mt s loi khc (trng l, trng bo t) kt
qu ca hnh thc sinh sn v tnh khng phi ch l 2 m l nhiu c th mi. l hin tng lit sinh
(schizogenesis).
Sinh sn hu tnh: xut hin cc giao t (gametes) v kt hp vi nhau tng i to thnh hp t
(zygote); t pht trin tip tc thnh c th mi (trng l, trng bo t).
Trong chu trnh sinh sn v pht trin ca mt s loi (chng hn Plasmodium spp., Elphidium spp....)
hai hnh thc sinh sn v tnh v hu tnh xut hin k tip nhau. l hin tng xen k th h
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(metagenesis).
Vi khong 30.000 loi, sng trong nhiu mi trng khc nhau, ng vt nguyn sinh rt a dng v
hnh thi v sinh hc. Chng c phn thnh 4 ngnh khc nhau: ngnh trng chn gi (Sarcodina), ngnh
trng roi (Mastigophora), ngnh trng bo t (Sporozoa) v ngnh trng lng hay trng tim mao
(Ciliophora). Bng 4.2 di y gii thiu tm tt cc c tnh ca cc ngnh thuc phn gii Protozoa.
Bng 4.2. c tnh ca cc nhm thuc phn gii Protozoa
Ngnh
Tn thng
gi
Trng roi-
Trng roi
Mastigophora
(Flagellates)
Trng chn gi-
Trng amip
Amoeba
Sarcodina
(Amebas)
Entamoeba
Trng tim mao-
Trng lng
Balantidium
Ciliophora
(Ciliates)
Paramecium
Trng bo t-
Trng bo t
Plasmodium
Sporozoa
(Sporosoans)
Toxoplasma
i din
Phn b
Trypanosoma
Giardia
Bnh thng gp
- Bnh ng chu Phi.
Nc ngt, k sinh
ng vt
Trichomonas
- Bnh do Giardia.
- Bnh do Trichomonas.
Nc ngt, mn, k
sinh ng vt
Kit l do amip (amebiasis).
Nc ngt, mn, k
sinh ng vt.
Kit l
K sinh ng vt,
cn trng
St rt, bnh do
Toxoplasma
2.3.2. Phn gii to (Algae)

To l nhm sinh vt c cu to n bo, tp on hay a bo n gin hoc phn ha thnh thn, r, l
gi v phn ln sng nc. Dinh dng l t dng nh c sc t quang hp, d dng ch c mt s sng
trong iu kin c bit. Sinh sn thng l sinh sn hu tnh v c s lun chuyn th h n bi v th h
lng bi trong chu trnh sng. C quan sinh sn chnh l n bo c th n bo, hoc a bo c th a
bo. Nu c quan sinh sn l a bo th tt c cc t bo u l t bo sinh sn (tr nhm to vng).
Hin nay pht hin c khong 40.000 loi to trn ton th gii. Ngi ta d on s lng ny ch
chim khong 11% s loi thc c. Vit Nam, s loi to c pht hin khong trn 2.000 loi. Vic
phn loi to l mt vn phc tp do s a dng v sinh hc ca chng v thc t c rt nhiu quan
im khc nhau. y ch trnh by mt s ngnh to da theo cu trc t bo, kiu sng v s c mt ca
cc sc t quang hp v cht d tr, mc d hin nay mt s ngnh nh to , to nu v to lc c
xp vo gii thc vt.
2.3.2.1. To gip (Pyrrophyta)
Ngnh ny c khong 1.000 loi. To gip l nhng to nh thng l n bo, c dip lc a v c, c
mu vng lc n nu nh c cc carotenoid khc v peridinin. Tuy nhin, c nhiu loi khng mu. Dinh
dng bng cht hu c. Cht d tr l tinh bt. Mng t bo l cellulose gm nhiu mnh ghp li vi nhau
v c hai rnh: mt rnh ngang v mt rnh dc. Ti ch gp nhau ca hai rnh c hai roi khng bng nhau
i ra, mt nm trong rnh ngang v mt nm trong rnh dc.
Vi khun lam l loi cng sinh trong to gip nhit i v to gip cng l loi cng sinh trong nhiu
ng vt khng xng sng bin.
Mt s to gip c kh nng pht sng, mt s khc tit ra cht mu rt c gy ra hin tng thy
triu , git cht hng triu con c.
2.3.2.2. To vng (Chrysophyta)
C khong 10.000 loi thuc ngnh ny. Phn ln cc loi to c xp vo ngnh ny. c im c
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trng l c mu vng hoc nu do c cht ch o l carotenoid, c dip lc a v c, khng c dip lc b,

c cht d tr leucosin thay cho tinh bt. To silic l loi ph bin nht, vch t bo kp v c bo ha
silic, c hai roi khng u nhau, tuy nhin c mt s trng hp mt roi hoc hai roi u nhau.
Phn ln Chrysophyta l n bo hay tp on, ch yu sinh sn v tnh cn sinh sn hu tnh him gp,
sng c vng nc ngt v nc mn.
2.3.2.3. To nu (Phaeophyta)
Nhm ny gm cc loi to nu tng t thc vt, sng c nh bin, c cu trc a bo dng si hay
dng bn, c t 2 n nhiu lp t bo phn ho thnh thn, r v l. Vch t bo l cellulose v alginic. Th
mu hnh a, hnh ht hay hnh tn c sc t dip lc a, c, fucoxanthin v cc carotenoid khc. Cht d tr l
laminarin.
S sinh sn thng gm hai qu trnh: sinh sn v tnh bng cc bo t chuyn ng 1-2 roi v sinh sn
hu tnh bng cc hnh thc ng giao, d giao, non giao.
To ny to thnh rng bin nn l ngun thc n, ni tr n v sinh sn ca nhiu ng vt bin. To
cha hm lng iod cao nn c dng lm thuc. Chit acid alginic v chuyn thnh alginat lm t dc
dnh trong ngnh dc. Mt s lm thc n v phn bn.
2.3.2.4. To (Rhodophyta)
Ngnh ny gm cc loi to sng bin, c cu trc a bo, dng si phn nhnh, dng bn v mt s
phn thnh thn, r, l gi tng t thc vt. Vch t bo l cellulose, pha ngoi c bao bi cht nhy, vi
hay mui silic. Th mu dng a, hnh sao hay hnh si. Sc t dip lc l a (chlorophyll a), phycocyanin,
cc phycobilin v c trng l sc t mu phycoerythrin, cht d tr l tinh bt.
Sinh sn v tnh bng bo t bt ng, cn sinh sn hu tnh bng non giao. To va c dng c s
chuyn giao th h va c dng khng.
To thng pht trin su di y bin v nh c sc t m c kh nng thu nhn nh sng Mt
Tri. Nhiu loi trong ngnh ny dng lm rau n nh rau dong (Hypnea), rau cu (Gracillaria). nc ta
c loi ph bin l Gracillaria stemispicata.
2.3.2.5. To lc (Chlorophyta)
L ngnh rt a dng: n bo, tp on, hoc a bo hnh si, hnh bn, c khi phn ha thnh thn, r
v l gi, mt s dng cng bo. To thng c mu xanh lc do c sc t chnh l dip lc v sng ph
bin nc ngt.
c im ni bt trong chu trnh sng l c giai on bo t chuyn ng nh roi. T bo c sc t
quang hp ging thc vt l chlorophyll a, b. Ngoi ra cn c cc sc t carotenoid khc, cht d tr l tinh
bt. Sinh sn thng gm hai qu trnh: sinh sn v tnh bng cc bo t chuyn ng c hai roi, t cc bo
t chuyn ng pht trin thnh c th to mi; v sinh sn hu tnh l qu trnh to ra c th mi c s kt
hp ca hai giao t to nn hp t 2n. Ngay trong hp t, qu trnh phn chia gim nhim xy ra to nn
nhng t bo n bi v cho ra cy mi. Sinh sn hu tnh gm 3 dng: nu hai giao t chuyn ng, c hnh
dng ging nhau (ng giao t) th gi qu trnh l ng giao; nu hai giao t chuyn ng, khc nhau v
hnh dng (d giao t) th gi qu trnh l d giao. Trng hp th 3, khi c nhiu giao t b chuyn ng
v mt t giao t ln khng chuyn ng (bt ng) th qu trnh kt hp gi l non giao. Mt s trng
hp hai si to gn nhau hnh thnh ng tip hp v khi gp nhau ni cht cho nhau to thnh hp t gi
l sinh sn bng tip hp.
2.4. Gii thc vt (Plantae)

Thc vt l nhng sinh vt nhn thc, a bo, khng di chuyn v c kh nng quang hp. C t bo
c bao bc bi thnh t bo bng cellulose. Theo quan im hin i, thc vt bao gm ba nhm chnh:
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Thc vt nc n bi-to (Algae) bao gm 3 ngnh l to lc, to nu, to c trnh by

phn gii to, thuc gii Protista.
Thc vt cn khng mch gm ngnh ru.
Thc vt cn c mch gm ngnh dng x, ngnh c thp bt, ngnh thng t, ngnh ht trn v
ngnh ht kn.
Thc vt c chia thnh cc ngnh khc nhau cn c vo hai c quan chnh l c quan dinh dng v
c quan sinh sn.
i vi c quan dinh dng, cn c vo hai dng chnh l c th dng tn hay c th dng chi. C th
dng tn cn c vo hnh dng tn, cu trc t bo, cc cht mu quang hp v cht d tr trong tn phn
chia ra cc ngnh nh phn gii to c trnh by phn trn trong gii Protista. Loi c th dng chi
cn c vo s phc tp ha dn ca chi m phn loi:
Chi c thn v l nhng cha c r tht.
Chi c thn, l v r tht, tc bn trong cc c quan hnh thnh cc mch dn truyn chnh.
i vi c quan sinh sn, cn c vo hai dng chnh l sinh sn bng bo t v sinh sn bng ht. Vi
bo t c hai loi chnh l bo t v tnh v bo t hu tnh. Vi ht: thc vt c ht c th c hoa cha hon
chnh, tc cha hnh thnh nhy v ht cn trn, c gi l thc vt ht trn. Thc vt c ht li c th c
hoa hon chnh, tc hoa hnh thnh nhy v ht c bc kn, c gi l thc vt ht kn. Thc vt ht
kn li c hai loi: loi mt l mm nu trong ht c mt l mm v hai l mm nu trong ht c hai l mm.
Ngi ta chia cc ngnh thc vt thnh cc n v phn loi nh hn nh lp, b, h cn c vo cch sp
xp cc b phn ca hoa, cm hoa, bu, nh v nhy. Trong hai c quan dinh dng v sinh sn th c quan
sinh sn l c im t b mi trng chi phi hn v l c im phn loi quan trng nht ca thc vt.
Nh ngnh ru l cc thc vt cn c c th cu to chi gm thn, l, r nhng l r gi v trong thn, l
cha c mch. Ngnh dng x c r tht, trong thn v l c mch, sinh sn bng bo t. Ngnh ht
trn gm cc loi thc vt sinh sn bng ht nhng l non cha khp kn che ht. Ngnh ht kn sinh sn
bng ht v l non khp kn che ht.
Thc vt l ngun cung cp oxy, thc phm, vi, vt liu xy dng, cc cht mu, gia v, thuc nhum
v dc liu.
2.5. Gii nm (Fungi)

2.5.1. c im chung ca nm
Nm l nhm sinh vt c nhn tht (Eucaryotae), c th nm khng c mch. Nm sinh sn bng bo t
v thng pht tn vo mi trng nh gi. Ring cc loi nm n bo thng sinh sn theo kiu ny chi
v mt s t loi sinh sn theo kiu trc phn. Bo t ca nm gm 2 loi: bo t hu tnh v bo t v tnh.
Cc loi bo t ny c sinh ra ty thuc vo loi v cc iu kin ngoi cnh. Nm ging thc vt, u c
s thay i th h. Nm khng c kh nng di ng ngoi tr mt s rt t thuc b nm roi (Chytridiales) c
pha di ng. Dng sinh dng ca nm c th l n bo (nm men) (hnh 4.11) hoc dng si (nm mc).
Dng si c thnh phn si n c gi l hypha, nhiu si gi l hyphae. Cc si c th c vch ngn
(septate hyphae) hoc khng c vch ngn, c gi l cc si cng bo (coenocytic hyphae) v mt h si
c gi l mycelium v nhiu h si l mycelia.
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Hnh 4.11. Hnh thi t bo nm men
i vi nm men, cc t bo thng tch ri nhau ra nhng cng c trng hp cc t bo dnh li vi

nhau to thnh si, nhng khng c s trao i cht gia cc t bo c gi l si gi (pseudomycelium).
Mt in hnh thng gp l cc si gi ca nm Candida albicans, mt c im quan trng trong chn
on xc nh loi nm thng gy bnh ny.
Thnh t bo ca nm c cu trc tng t thnh t bo thc vt nhng khc nhau v hp phn ha hc:
thnh t bo nm cu to ch yu l cht chitin, thnh t bo thc vt cu to ch yu l cellulose. Nm
khng phi l cc sinh vt t dng m l cc sinh vt d dng. Cng l sinh vt d dng nhng nm khc
vi ng vt: ng vt nut thc n vo d dy ri mi tiu ha cn nm th ngc li, tit ra cc enzym vo
c cht tiu ha ri mi hp thu vo c th nm.
Nm tch tr nng lng di dng glycogen nh ng vt v khc vi thc vt tch tr nng lng
di dng tinh bt. Cc mng t bo nm cu to bi sterol duy nht l ergosterol, cht thay th cho
cholesterol mng t bo ng vt c v. Hu ht nm c nhn t bo rt nh v vi ADN t trng lp. Qu
trnh nguyn phn ni chung c kt thc m khng c s phn hy mng nhn.
Ty thuc vo loi v iu kin ngoi cnh, nm c th c cc trng thi dinh dng: hoi sinh, k sinh
v cng sinh.
2.5.2. Phn loi nm
Da vo phng thc sinh sn bo t hu tnh, cc nh nm hc chia gii nm thnh 4 ngnh v mt
nhm sau y:
2.5.2.1. Ngnh nm roi (Chytridiomycota)
Hu ht cc loi thuc ngnh ny l nm thy sinh v sng d dng. Mt s c th k sinh gy cc
bnh thc vt. Cc bo t hu tnh v c bo t v tnh u c roi v c kh nng di ng.
2.5.2.2. Ngnh nm tip hp (Zygomycota)
L ngnh rt ph bin v thng c gi l mc bnh m. Hu ht sng hoi sinh, nhng cng c kh
nhiu loi sng k sinh gy bnh ngi v thc vt. Ngoi ra trong ngnh ny c b Glomales sng cng
sinh kh ph bin vi khong 70% thc vt trn th gii. Cc bo t hu tnh l dng bo t ngh, c thnh
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dy c gi l bo t tip hp (Zygospore). Cc bo t v tnh c sinh ra trong mt nang gi l nang

bo t (sporangium).
c im khc ca ngnh ny l si nm khng c vch ngn, cha nhiu nhn n bi. Bo t v c
giao t khng chuyn ng. Sinh sn hu tnh bng s tip hp ca hai si nm c gii tnh khc nhau.
i din in hnh l mc bnh m Mucor, Rhizopus. Chng c cu to gm ba loi si nm lm thnh
mng trn b mt bnh: si hnh r (Rhizoid) xuyn su vo bnh m hp th thc n, si kh sinh c bit
hnh thnh nang (Sporangium) cha cc bo t nh si v cc si c bit nm trn b mt c gi l
Stolon sinh cc cung nang bo t (Sporangiophore) (hnh 4.12). Khi cc bo t chn pht tn gp iu kin
m, bo t ny mm cho ra si nm mi, kt thc chu trnh v tnh. Sinh sn hu tnh do s kt hp hai si
nm: thnh t bo hai si bin i thnh ti sinh giao t. Hai ti giao t kt hp vi nhau to thnh hp t,
cc nhn b m kt hp vi nhau tng i mt to thnh nhn lng bi. Cc nhn bn trong tiu bin ch
cn li mt nhn. Nhn ny tip tc phn chia gim nhim to thnh 4 nhn n bi, 3 trong s chng mt i
cn li mt nhn. Cu trc c gi l bo t tip hp, c bo v bi mt mng dy thm cc th canxi
oxalat v sau ng trong mt thi k di. Khi bo t tip hp ny mm sinh ra ti bo t. Ti lc chn gii
phng hng ngn bo t n bi v gp iu kin m, mi bo t sinh ra mt c th nm mi.
Hnh 4.12. Hnh thi nm mc bnh m Mucor sp.
2.5.2.3. Ngnh nm ti (Ascomycota)
Hnh 4.13. Nhm nm Ascomycetes

a) Cc loi ti (ascus): hnh tr, hnh chy, hnh cu; b) Pha u ca sinh sn hu tnh;
c) Mt ct ngang ca th qu hnh qu l c cc ti bo t hnh tr.
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Ngnh ny bao gm cc loi nm chn, nm morel v hu ht nm men. a s cc tc nhn gy bnh

thc vt u thuc ngnh ny. Cc bo t hu tnh c sinh ra trong mt ti gi l Ascus (hnh 4.13). Cc
bo t v tnh c sinh ra bn ngoi (bo t trn) c gi l conidi.
Khc vi cc ngnh khc, ngnh ny si nm c cu trc vch ngn. Kt qu ca qu trnh sinh sn hu
tnh, hnh thnh ti trong cha cc bo t ti (ascospore), tc l cc nhn kt hp lng bi c bao bc
trong cht nguyn sinh. Kt qu ca sinh sn v tnh l bo t nh (conidium) c sinh ra nh cc si
nm chuyn ho. Mi mt bo t nh s sinh ra mt cy nm mi khi gp iu kin thun li.
Nm ti c th khng c th qu nh nm men. y l nm n bo sinh sn sinh dng bng ny chi,
sinh sn hu tnh bng cch kt hp hai t bo li thnh hp t.
Nm ti c th qu do cc si nm (1n + 2n) bn li, c 3 loi chnh: th qu ng kn, v d nm gy
bnh hc lo, vy rng, bnh cy; th qu m l, v d nm gy bnh m l chui, l lc v th qu hnh
a nh nm tai mo trng ging nh ci chn, nm gy bnh ci bp, c chua, c rt.
2.5.2.4. Ngnh nm m (Basidiomycota)
Hnh 4.14. Cc loi m sinh bo t m ca nhm nm Basidiomycota
a) m dng chy; b,c) Qu l; d) Hnh tr

Cc bo t hu tnh (Basidiospore) ca ngnh ny c sinh ra bn ngoi, trn mt cu trc hnh chy
c gi l m (Basidium) (hnh 4.14). Bo t v tnh thng khng c.
Nm m l nhm ph bin rng ri nh nm rm, nm hng, mc nh, nm trng,... Nm ny phn
ho thnh thn v m (tn) (hnh 4.15). l nhng si nm c vch ngn v song hch ken cht li. Phn
chnh ca si nm nm di mt t v sng hoi sinh.
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Hnh 4.15. Nhm nm m: thn v tn ca Amanita sp (Nathan Wilson, 1992)
Nm m c im khc c bn so vi cc nm khc l si a bo v song hch l ch yu, si n hch

ch gp lc ny mm v sinh sn bng bo t m hay cn gi l bo t ngoi sinh, tc l cc bo t n bi
c hnh thnh nm trn cc u li tch bit khi u ca si nm.
Qu trnh hnh thnh bo t m din ra nh sau: trn cc phin di m nm c cc u si nm hai
hch nhn (1 nhn n bi c v mt nhn n bi ci ng cnh nhau) kt hp to thnh nhn hp t
lng bi. Nhn phn chia gim nhim cho 4 nhn con. Bn u li mc ra tn cng v 4 nhn con i vo
. Cc vch hnh thnh tch 4 u li to thnh 4 bo t m. Bo t gi tung ra, ny mm to nn cc si
n bi. Hai si n bi kt hp vi nhau to thnh si song hch v ken li thnh th qu nm.
Nm ny c loi n c nh nm rm, nm s, mc nh. Nm dng lm thuc nh linh chi, nhng
cng c nhiu loi nm c nh nm lim, nm c en v nm c thuc chi Amanita c th gy cht
ngi khi n phi.
2.5.2.5. Nhm nm bt ton (fungi imperfecti hay deuteromycetes)
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Hnh 4.16. Hnh thi vi hc ca loi Penicillium notatum
Nhm ny gm cc loi nm khng bit trng thi hu tnh. Trng thi v tnh chnh l qu trnh hnh
thnh conidi (bo t nh). Bt k mt loi nm no khng bit trng thi hu tnh hay qu trnh gim phn
trong chu k sng ca chng, cho tin li ngi ta u xp vo nhm ny. Hu ht cc loi ca nhm ny
c mi quan h thn thuc vi ngnh nm ti v theo Hawkswarth (1983), c khong 1.680 chi vi 17.000
ngn loi c xp vo nhm ny. a s cc loi ca nhm ny sng trn cn, hoi sinh hoc k sinh
thc vt. Mt s t k sinh gy bnh ng vt, trong c con ngi. Mt i din in hnh ca nhm ny
l chi Penicillium, vi loi nm ni ting P. notatum (hnh 4.16) sinh khng sinh penicilin, m ra mt k
nguyn mi cho loi ngi trong vic chinh phc cc loi bnh do vi khun gy ra.
2.5.3. Vai tr ca nm
Nm hoi sinh l mt nhn t quan trng trong chu trnh vt cht ca s sng, n tham gia phn hu
xc ca thc vt, ng vt cht, cng nh cc loi rc thi khc.
Nm c dng trong sn xut bnh m, ru, bia. Chng c dng lm thc phm, lm thuc v l
ngun sn xut cc cht khng sinh...
Nm k sinh l tc nhn gy bnh ngi, ng vt v cho cy trng.
Nm hoi sinh gy h hng lng thc, thc phm, c bit l cc sn phm ng cc, ng thi sinh
c t (mycotoxin) gy tc hi n sc khe v kinh t cho con ngi.
2.6. Gii ng vt (Animalia)
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Da trn nhng c im v cu to sinh hc hoc c cng ngun gc, hoc biu hin tnh cht phn
ho v c tin b hn (c im t chc v i xng c th; ngun gc l ming; s hnh thnh, mc
phn ho v pht trin ca th xoang,...) ng vt a bo c sp xp theo mt h thng sau y:
ng vt a bo bc thp (Parazoa), ch c mt ngnh: Porifera.
ng vt a bo chnh thc (Eumetazoa) gm cc ngnh cn li:
+ ng vt c c th i xng to trn (Radiata): ngnh Coelenterata v Stenophora.
+ ng vt nguyn khu, cha c th xoang (Protostomia, Acoelomata): ngnh Plathelminthes v
Nemertini.
+ ng vt c th xoang gi (Pseudocoelomata): ngnh Nemathelminthes v Entoprocta.
+ ng vt c th xoang tht, c th khng phn t (Coelomata, Inarticulata): ngnh Priapulida,
Sipunculida, Mollusca, Echuirida.
+ ng vt c c th phn t (Articulata): ngnh Annelida, Tardigrada, Onychophora, Pentastomida,
Arthropoda.
+ ng vt hu khu (Deuterostomia), gm tt c cc ng vt cn li: ngnh Phoronida, Bryozoa,
Ectoprocta, Brachiopoda, Echinodermata, Chaetognatha, Pogonophora, Hemichordata v Chordata.
Sau y gii thiu mt s ngnh c lin quan nhiu ti lnh vc y - dc hc vi nhng c im c bn
nht.
2.6.1. Ngnh giun dp (Plathelminthes)
Giun dp l nhm ng vt a bo u tin c i xng hai bn (Bilateria), xut hin l phi th ba hay
trung b (mesoderm), cha c th xoang (acoelome). C khong 12.700 loi; sng trong nc ngt, bin, mt
s sng trong t m; nhiu loi k sinh gy bnh ng vt v ngi.
C th dt theo hng lng - bng, hnh l hoc hnh di di. Phn ln cc loi sn tim mao c kch
thc trung bnh t 0,4 - 5mm, trong khi cc loi sn dy c chiu di c th rt ln nh sn mp
(Dyphyllobotrium latum) di 15m, thm ch 20m.
Thnh c th l mt bao c gm cc lp c dc, c vng v c lng - bng. Bc ngoi c th l mt lp
biu m n. cc loi giun dp sng t do, trn lp biu m ny (t mt bng) c ph mt lp tim
mao; cc loi k sinh, lp biu m tim mao c thay bng lp cuticun (nhn, c sc chu ng cao trong
mi trng k sinh).
ng tiu ho c cu to n gin (c l ming, rut, cha c hu mn). cc loi k sinh, c quan tiu
ho hon ton tiu gim (dinh dng theo li thm thu); ngc li, c quan bm pht trin (gic bm, mp
bm, mc bm). Thuc v h thn kinh - cm gic c hch no v cc dy thn kinh xut pht t . sn
tim mao, cu to h thn kinh t nhiu vn mang tnh cht i xng to trn. Thng c cc thu cm gic,
mt, bnh nang trn phn u c th cc loi c i sng t do. Chc nng bi tit do nguyn n thn
(protonephridium, cn gi l t bo ngn la), ln u tin xut hin giun dp, cha c c quan tun
hon v h hp.
Hu ht cc loi giun dp l lng tnh v hnh thc sinh sn chnh l hu tnh: trng. C quan sinh
sn c cu trc phc tp. Trng sau khi c th tinh, mt s loi (ch yu l sn tim mao) s pht trin
trc tip thnh sn trng thnh; cc loi khc, phi tri qua giai on u trng vi mt vi dng cu to
khc nhau. nhiu loi sn dy, giai on u trng tn ti trong c th vt ch di hnh thc cc nang sn
(Cysticercus). Trong chu trnh sinh sn v pht trin ca mt s loi giun dp (v d sn l gan) c hin
tng xen k th h, thay i vt ch vi cc hnh thc sinh sn khc nhau. Hnh thc sinh sn v tnh (ch
yu bng cch ct on c th) cng gp mt s t loi (Microstomum lineare).
Ngnh ny c 3 lp:
Lp sn tim mao (Turbellaria), lp sn l (Trematoda) i din l: sn l gan (Fasciola hepatica) (hnh
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4.17); lp sn dy (Cestoda) vi i din l sn dy ln (Taenia solium) (hnh 4.18) v sn dy b

(Taenia saginata).
Hnh 4.17. Sn l gan Fasciola hepatica
Hnh 4.18. Sn dy ln Taenia solium
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2.6.2. Ngnh giun trn (Nemathelminthes)

Ngnh ny c khong 12.500 loi, giun trn sng trong mi trng nc hay t; nhiu loi sng k
sinh trong c th ng vt, thc vt ( ngi khong 50 loi). Giun trn c xoang c th nguyn sinh; ng
tiu ho hon chnh, nhng cha c c quan tun hon v h hp.
Khc vi cc loi giun dp, c th giun trn thng ko di v c dng hnh a, c xoang c th
nguyn sinh hay th xoang gi (Pseudocoelome). Thnh c th gm cc lp: cuticun (l mt lp v nhn,
khng mang tim mao), bao bc ton b mt ngoi c th; lp biu b (Epidermis), cu to kiu hp bo;
trong cng l lp c (c phn thnh 4 di dc). ng tiu ho c cu to n gin, c rut sau v hu
mn. Cc loi giun sng t do c th thu nhn thc n di dng lng hoc rn; trong khi , cc loi k sinh
thng hp th cht dinh dng qua b mt c th hoc tit men tiu ho m ca vt ch. Mt s loi
giun khng c c quan bi tit; s khc, pht trin dng tuyn hoc ng bi tit. H thn kinh gm vng hu
(bao quanh phn trc thc qun) v cc ng dy thn kinh chy dc c th, trong 2 dy ln hn nm
trong g lng v g bng ca lp biu b. Cc gic quan giun trn km pht trin. Tuyn sinh dc c dng
hnh si di v mnh; sn phm sinh dc c thi ra qua l sinh dc. Cc loi k sinh thng c kh nng
sinh sn ln (Ascaris c th ti 20 triu trng). Cha c c quan tun hon v h hp chuyn trch.
Hu ht giun trn l phn tnh, th tinh trong, trng v ch c sinh sn hu tnh; mt s t loi con
(giun ch). Trong chu trnh sng ca mt s loi giun k sinh (giun ch, giun xon), u trng c pht trin
qua vt ch trung gian; mt s loi khc (giun a, giun kim) c ly nhim trc tip (khng thng qua vt
ch khc).
Tui th trung bnh ca cc loi giun sng t do thng l ngn; trong khi giun a k sinh ngi c
th sng n 1 nm, hay 17 nm i vi giun ch.
Ngnh giun c phn thnh 6 lp (Rotatoria, Gartrotricha, Nematoda, Nematomorpha, Kinorhyncha
v Acanthocephala), trong lp giun trn (Nematoda) chim hu ht s lng loi ca ngnh (trn 10.000
loi), cng l lp c nhiu loi k sinh ng vt v ngi. i din l giun a (Ascaris lumbricoides)
(hnh 4.19), giun kim (Enterobius vermicularis) (hnh 4.20), giun ch (Wuchereria bancrofti, Brugia malayi)
gy bnh chn voi (Elephantiasis) (hnh 4.21).
Hnh 4.19. Giun a Ascaris lumbricoides
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Giun kim (Enterobius vermicularis)

Hnh 4.20. Giun kim ci v c
Hnh 4.21. Bnh chn voi do giun ch (Wuchereia brancrofti) gy ra
2.6.3. Ngnh chn khp (Arthropoda)

y l ngnh c s lng loi ln nht trong gii ng vt (hn 1 triu loi, trong khong 850.000
loi l cn trng). ng vt chn khp c th tm thy mi ni, mt nhm ng vt a dng v cu to,
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phong ph trong li sng, biu hin mt s phn ho v thch nghi cao i vi mi trng. Mt khc,
cng xp trong nhm ng vt phn t (Articulata), nhiu du hiu cho thy chn khp c quan h ngun
gc vi giun t.
Nt ni bt ng vt chn khp l s hon thin v mt t chc c th: phn t d hnh (Heteronom)
vi s hnh thnh cc nhm t to nn cc b phn khc nhau ca c th (thng thng c th c chia
lm 3 phn: u, ngc v bng). Trn mi t c th ch yu phn u v ngc c mang cc i phn ph
phn t (c ngun gc t chi bn ca giun t). Cc phn ph c bin i ph hp vi cc chc nng
khc nhau: thnh cc b phn ca ming thu nhn thc n, cc phn ph ca c quan sinh dc, chn
vn ng (b, nhy, bi); nhiu loi (cn trng) cn pht trin thm (1 hoc 2 i) cnh bay.
Bc ngoi c th l lp v cuticun, cn c gi l b xng ngoi (va c tc dng bo v va l ni
bm cho cc c bn trong). Trong qu trnh pht trin, cc loi chn khp c hin tng lt xc (thay bng
lp v mi) tng kch thc c th.
H c pht trin v phn ho: hnh thnh nhng b ring bit m bo mt s hot ng a dng v
linh hot.
Xoang c th ng vt chn khp l hn hp gia xoang nguyn sinh (xoang gi) v xoang th sinh
(xoang tht), v th c gi l th xoang hn hp (mixocoelome) hay xoang huyt (haemocoelome, v c
cha mu) v tr thnh mt b phn ca h tun hon. V vy, khc vi giun t, h tun hon chn khp
l h tun hon h. Tham gia vo h tun hon cn c tim lng (c phn ho t mch lng ca giun t)
v h mch ti cc c quan. Chc nng ch yu ca h tun hon l vn chuyn cht dinh dng ti cc m
v cht thi n c quan bi tit.
Vic cung cp oxy do c quan h hp m nhim. Phn ln cc loi nc pht trin cc l mang (mt
s t c th h hp qua da); cc loi cn trao i kh qua h thng kh qun (cn trng, nhiu chn) hoc
qua ti phi (nhn, b cp).
Ngoi cc b phn chnh ca ng tiu ho (ming, hu, thc qun, rut, hu mn) cc tuyn tiu ho
cng pht trin: tuyn rut gia ( gip xc), tuyn nc bt ( nhn v cn trng). ng vt chn khp c
th s dng thc n rn (t cc ng, thc vt) hoc theo cch ht dch. Tu theo cch thu nhn thc n khc
nhau m phn ph ming c bin i ph hp. Chc nng bi tit do cc tuyn thc hin: tuyn ru hoc
tuyn hm gip xc, tuyn hng sam, tuyn Malpigi nhn v cn trng. Hai b phn chnh ca h thn
kinh l no v dy thn kinh bng. nhiu loi no phn ho thnh no trc, no gia v no sau. Cc
loi chn khp c i sng x hi (ong, kin, mi) no trc c th nm pht trin (l trung khu ca bn
nng).
Hu ht cc loi chn khp l ng vt n tnh v sinh sn bng cch trng. S pht trin hu phi
nhiu loi l mt qu trnh bin thi phc tp (tri qua dng u trng vi nhiu ln lt xc v giai on
nhng).
L nhm ng vt c s lng loi phong ph nht trong gii ng vt, ngnh chn khp gm 4 ngnh
ph: trng ba thu (ngnh ph Trilobitomorpha), c km (ngnh ph Chelicerata), c mang (ngnh ph
Branchiata) v c kh qun (ngnh ph Tracheata).
2.6.3.1. Mt s lp chnh v i din thng gp
Lp hnh nhn (Arachnida): gm cc ng vt chn khp sng trn cn, c th phn thnh u, ngc
v phn thn sau, c 4 i chn. i din l nhn nh (Heteropoda pressula), b cp (Pandinus dictador).
Lp gip xc (Crustacea): gm cc ng vt chn khp sng bin, nc ngt, c 2 i ru, hu ht
u c mu ph. i din l tm sng (Macrobrachium nipponense), cua ng (Somaniathelphusa sinensis).
Lp chn mi (Chilopoda): thn di, c nhiu chn b. C th trng thnh mi t mang chn c mt
cp chn. i din l rt (Scolopendra).
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Lp chn kp (Diplopoda): gm cc ng vt chn khp c thn di, c nhiu chn di chuyn. C

th trng thnh mi t mang chn c 2 cp chn. i din l cun chiu (Polydesmus).
Lp cn trng (insecta = Hexapoda): gm cc loi chn khp sng nc ngt v trn cn, c mt
i ru. C th phn ha thnh u, ngc v bng, c 3 i chn. Gm 2 phn lp:
+ Phn lp khng cnh (Apterygota) gm cc cn trng khng cnh. i din l b ui bt (b bt)
Lepisma.
+ Phn lp c cnh (Pterygota). Phn lp ny c chia thnh 2 nhm l nhm bin thi khng hon
ton (Cnh ngoi- Exopterygota), vng i khng qua giai on nhng, giai on su non ging vi con
trng thnh, cnh pht trin pha bn ngoi; i din l chun chun (Anax). Nhm bin thi hon ton
(Cnh trong-Endopterygota), vng i qua giai on nhng, cc giai on su non khng ging vi con
trng thnh, cnh pht trin bn trong. i din l cc loi bm (Pieris).
Lp cn trng c khong 850.000 loi, thuc trn 30 b khc nhau.
2.6.3.2. ngha y hc v kinh t ca ngnh chn khp
Nhiu loi c li (cho cc sn phm nh mt ong, t tm), nhng cng khng t loi c hi nh ph hoi
cy trng, c bit l vai tr vt trung gian truyn bnh (vector) cho ngi v ng vt khc. in hnh l
bnh st rt (malaria) do mui ci Anopheles gy ra (hnh 4.22), bnh Lyme, bnh st chm ni Rocky do ve
gy ra....
Hnh 4.22. Mui Anopheles sp.
2.6.4. Ngnh thn mm (Mollusca)

Vi gn 130.000 loi (trong khong 35.000 loi ho thch), thn mm l ngnh ln th hai trong gii
ng vt (sau chn khp). Thn mm sng trn cn hoc di nc, a dng v t chc c th cng nh cu
to c quan. C nhiu ngha kinh t v thc tin: ngun cung cp thc phm (trai, c, mc, s,...), cho vt
liu qu (x c, ngc trai),... Tuy nhin, mt s chng cng gy hi, ph hi cy trng hoc l vt truyn
bnh (c tai, sn trn).
Ngnh ny c 4 lp chnh:
Lp song kinh (Amphineura);
Lp chn bng (Gastropoda) c i din l: c nhi (Pila polita), bo ng (Haliotis diversicolor), c
sn (Achatina fulica).
Lp 2 mnh v (Bivalvia) vi i din l: trai sng (Sinanodonta jourdyi), s huyt (Arca granosa),
hn (Corbicula lamarkiana), trai ngc (Pteria martensi),...
Lp chn u (Cephalopoda) c i din l: mc (Sepia, Loligo); bch tuc (Octopus), duc bin
30/09/2009
Page 30 of 36
( Architeuthis); c anh v (Nautilus pompilius).

2.6.5. Ngnh da gai (Echinodermata)
Vi khong 6.000 loi cn sng (tt c u bin) v trn 20.000 loi ho thch, ng vt da gai (gm
cc lp hu bin, sao bin, ui rn, cu gai v hi sm) thuc nhm ng vt u tin c ming th sinh
(Deuteros-tomia). T chc v cu to c th a dng. Mt s loi c khai thc lm thc phm (hi sm,
cu gai).
Ngnh ny c 5 lp chnh: lp cu gai (Echinoidea); lp sao bin (Asteroidea); lp hi sm
(Holothuroidea); lp ui rn (Ophiuroidea) v lp hu bin (Crinoidea).
2.6.6. Ngnh ng vt c dy sng (Chordata)
Phn ngnh cxng sng (Vertebrata)
ng vt dy sng, ngnh ng vt cui cng ca gii ng vt, bao gm nhng loi gn gi v quen
thuc vi chng ta: c, ch, nhi, b st, chim, th (c con ngi cng thuc ngnh ng vt ny). a dng
v cu to, phong ph trong cch sng, hn 40.000 loi thuc ngnh ng vt ny hin l ch nhn trn khp
cc lc a v i dng.
Ngoi mt s c im xut hin mt vi ngnh ng vt khc (c th phn t v i xng 2 bn,
th xoang v ming th sinh) c th nu 3 c im c trng nht cho nhm ng vt ny nh sau:
- Hoc tn ti trong sut cuc sng (i vi ng vt bc thp) hay ti thiu trong giai on pht trin
phi ( ng vt bc cao); cu to c trng nht ca ngnh l dy sng (Chorda dorsalis). Dy sng c
ngun gc ni b, chy dc lng, cu trc mm do nhng vng chc, c coi l tr ct ca b xng trong.
cc ng vt bc cao, dy sng c ct ho dn dn cc mc khc nhau to thnh ct sng.
- Thn kinh trung ng gm mt ng thn kinh (m lng ng l xoang thn kinh) chy dc lng, nm
pha trn dy sng, c ngun gc ngoi b. V pha trc, ng thn kinh phnh to v pht trin thnh no b
(c bc trong hp s bng sn hay bng xng).
- hu ht cc ng vt dy sng (ti thiu trong giai on pht trin phi) c hnh thnh cc khe mang;
qua , khoang hu (phn u ca ng tiu ho) c thng vi bn ngoi. i vi cc ng vt nc, khe
mang tn ti sut i.
ng vt dy sng c phn thnh 3 phn ngnh: c bao (Tunicata), u sng (Cephalochordata) v
c xng sng (Verebrata). Trong , nhng ng vt quen thuc s c ni ti thuc phn ngnh th 3.
Khi qut mt s c im chnh (c xem l nhng c im thch nghi v tin ho) ca nhng ng
vt thuc phn ngnh Vertebrata:
- B xng trong: Khc vi cc ng vt dy sng bc thp, ng vt c xng sng c b xng trong
bng sn hay xng, to thnh mt b khung vng chc nng v bo v c th (hp s bo v no b,
khung xng sn che ch ni tng). Ngoi ra, b xng cn l ni bm cc c, tu theo mc pht trin
v tin ho ca ng vt m b xng c ct ho (ho xng) dn dn. Trong khi dy sng (khng phn
t, mm do) ch xut hin trong thi k pht trin phi, giai on u trng hoc nhng ng vt bc
thp th nhng ng vt bc cao c thay th bng ct sng (phn t v ho xng). Cc b phn
khc ca b xng cng c ct ho, nh m tnh cht c hc ca b xng c tng cng.
H thn kinh - cm gic: Hot ng sng phong ph v a dng ca ng vt c xng sng l kt qu
ca s pht trin ngy cng hon thin v mang tnh cht tin ho thch nghi ca h thng thn kinh v cc
c quan cm gic. ng vt bc cao, h thn kinh gm 2 h thng chnh: h thn kinh trung ng v h
thn kinh ngoi bin. H thn kinh trung ng gm no b v tu sng. No b t ch l mt phn phnh
n gin ca u ng thn kinh dn dn c phn ho thnh 5 b phn chnh: no trc (vi s pht trin
ca hai bn cu no), no trung gian, no gia, hnh tu v tiu no; cng vi s xut hin cc trung khu
iu ho hot ng ca c th. Tu sng thc hin 2 chc nng quan trng: dn truyn cc xung ng n
30/09/2009
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no v l trung khu phn x. T no b v tu sng xut pht tng cp dy thn kinh n cc th quan
v gic quan ca c th, cc dy thn kinh ny tp hp thnh thn kinh ngoi bin.
Ngoi ra, cc gic quan cng rt pht trin: c quan ng bn (nhn bit cc xung ng v p lc ca
nc, chuyn ho cho cc loi sng trong mi trng nc); c quan thnh gic, th gic, khu gic,... ngy
cng pht trin v hon thin nhng ng vt bc cao.
H tun hon kn gm tim v h mch. Tim c chia thnh cc ngn (tm tht, tm nh); h mch
gm 3 loi: ng mch (dn mu t tim n m v cc c quan), tnh mch (dn mu v tim) v mao mch
(mt h thng mch mu nh b, thnh mch mng, nm trong cc m, ni lin ng mch v tnh mch, vi
chc nng c bn l thc hin s trao i kh, cht dinh dng v cc sn phm trao i cht khc gia mu
v cc m). Cc loi ng vt nc tim c 2 ngn vi mt vng tun hon. Cc loi cn (chim, th) tim
c 4 ngn (2 tm tht, 2 tm nh) v 2 vng tun hon (vng tun hon nh mu qua phi, vng tun hon
ln mu i nui c th). Trong khi , ch nhi, b st c mt h tun hon trung gian: tim c 3 ngn (1
tm tht v 2 tm nh), 2 vng tun hon khng tch bit r rng nn mu b pha trn.
Ngoi h tun hon mu (cha hng cu), cc ng vt cn c mt h tun hon khc gi l h bch
huyt (cha bch cu). y l mt h tun hon h (ch gm mao mch v tnh mch, khng c ng mch,
dng dn cc cht dch v tim). Chc nng ch yu ca h bch huyt l iu chnh nng protein gia
dch m v h mao mch, ngn cn v tiu dit cc vt l (chng hn vi khun) xm nhp vo c th.
Ngoi mt s c im c bn (v b xng, h thn kinh - cm gic, h tun hon), cc h c quan
khc ca ng vt c xng sng (h hp, tiu ho, bi tit, sinh dc,...) cng th hin nhng c im v
cu to v chc nng cao hn, hon thin hn so vi cc ng vt khc.
y l mt phn ngnh ln ca ngnh dy sng (Chordata), cc ng vt c xng sng (Vertebrata)
c phn thnh 8 lp: Agnatha, Placodermi, Chondrichthyes, Osteichthyes, Amphibia, Reptilia, Aves,
Mammalia. c cp y ch l nhng ng vt quen thuc ca 5 lp cui.
(1) Lp c xng (Osteichthyes)
B xng c ct ho cc mc khc nhau, da c nhiu tuyn nhy v ph vy, vy bi pht trin,
th bng mang, thng c bong bng hi, tim c 2 ngn vi 1 vng tun hon; khong 21.000 loi.
Cc loi thng gp: c chp, c m, c thu,...
(2) Lp lng c (Amphibia)
Sng nc hoc trn cn, bin nhit, b xng bng xng, da trn v m; giai on trng thnh th
bng phi v da, trong khi giai on u trng (nng nc, sng nc) th bng mang; tim 3 ngn vi 2 vng
tun hon. Khong 2.500 loi. Thng gp: ch, nhi, cc...
(3) Lp b st (Reptilia)
Sng trn cn, bin nhit, b xng c ct ho ton thn; thn ph vy sng hay nhng tm xng b
(sn phm ca biu b); da kh; h hp bng phi; tim 3 ngn vi 2 vng tun hon. Khong 6.000 loi.
Thng gp: rn, ra, c su, thn ln, tc k,...
B st cng l lp c nhiu dng ho thch khng l: thn ln sm (di 20m, nng 30 tn), thn ln bay,
khng long (di 15m, cao 6m).
4) Lp chim (Aves)
B xng c ct ho hon ton, xp v c nhiu khoang kh; da mng, khng cha cc tuyn (tr
tuyn phao cu); hm khng c rng, m v vut chn bng sng; tim 4 ngn, mu nng, thn nhit tng
i n nh (40 - 42oC). Chim l nhm ng vt duy nht c cu to thch nghi vi hot ng bay (chi trc
bin i thnh cnh); phn b rng, s lng loi nhiu (khong 8.600 loi) l c tr ca nhiu mm bnh
nh cc loi virus, nm.
Thng gp: b cu, chim s, g, vt,...
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5) Lp c v (Mammalia)
Gm cc ng vt c t chc c th cao nht, a dng, phn b rng; c th ph lng mao; v da c
nhiu tuyn (m hi, tuyn sa, tuyn thm); nui con bng sa; h thn kinh trung ng v tun hon pht
trin hon chnh, mu nng, ng nhit. Hn 4.000 loi c chia thnh 3 phn lp.
Phn lp nguyn th (Prototheria): th m vt, th lng chim.
Phn lp th thp (Metatheria): kanguru, gu ti, chut ti.
Phn lp th cao (Eutheria) hay th c nhau (Placentalia): gm cc ng vt cn li (tru, b, kh,
vn, c voi,...).
Con ngi (Homo sapiens) d c habilis (kho lo) hay sapiens (khn ngoan) n u, ng v mt
phn loi hc vn c xp chung cng b vi h hng nh kh - b linh trng hay kh hu (Primates).
CU HI LNG GI
V minh ha cu trc ca Bacteriophage.
K tn mt s bnh do virus gy ra.
V minh ha cc hnh dng thng gp ca t bo vi khun.
Trnh by s khc bit c bn ca thnh t bo vi khun Gram (-) v Gram (+).
Trnh by vai tr v c ch ca cc bc trong phng php nhum Gram.
Trnh by c ch chuyn ng ca amip.
Trnh by s khc bit ca nm men v nm mc.
Trnh by cc ng dng quan trng ca gii nm trong ngnh cng nghip thc phm v ngnh
dc.
9. K tn cc dng bnh thng gp do vi nm gy ra.
10. K tn mt s vector thng gp ca ngnh chn khp.
1.
2.
3.
4.
5.
6.
7.
8.
TI LIU THAM KHO
30/09/2009
Page 33 of 36
Ting Vit
1. B mn thc vt hc, i hc Dc H Ni (1995); Sinh hc x hi sinh vt v tnh a dng
ca s sng, Trung tm th vin thng tin, i hc Dc H Ni.
2. Hong c C (1999), Sinh hc i cng, tp 1, NXB i hc Quc gia H Ni.
3. H Hunh Thu Dng (2002), Sinh hc phn t, NXB Gio dc.
4. Nguyn Nh Hin, Trnh Xun Hu (1997), T bo hc, NXB i hc Quc gia H Ni.
5. Phm Thnh H (2004), Di truyn hc, NXB Gio dc.
6. L nh Lng - Phn C Nhn (2003), C s di truyn hc, NXB Gio dc.
7. Nguyn Ngha Thn, V Trung Tng, L V Khi, Nguyn Vn nh (1995), Sinh hc x hi
sinh vt v tnh a dng sinh hc, i hc Khoa hc
T nhin.
8.
Trng i hc Y H ni, B mn Y sinh hc Di truyn (2002), Cc nguyn l sinh hc,

NXB Y hc.
9. V Vn V, V Thanh Tm, Hong Minh Tn (1999), Sinh l hc thc vt, NXB Gio dc.
10. W. D. Phillips, T.J. Chilton (1999), Sinh hc, tp 1, 2, NXB Gio dc.
Ting Anh
11. David H. Cormack (1997), Essential histology, Lippincott Williams & Wilkins.
12. Edward D. Frohlich (1997), Rypins - Basic science review, Lippincott Williams & Wilkins.
13. Michael T. Madigan, John M. Martinko, Jack Parker (1995), Biology of Microorganisms,
Prentice Hall International, Inc.
Chu trch nhim xut bn:

Ch tch HQT kim Tng Gim c NG TRN I
30/09/2009
Page 34 of 36
Ph Tng Gim c kim Tng bin tp NGUYN QU THAO

Chu trch nhim ni dung:
Ch tch HQT kim Gim c Cng ty CP Sch H - DN
TRN NHT TN
Bin tp ni dung v sa bn in:
HONG TH QUY
Thit k m thut v trnh by ba:
INH XUN DNG
Thit k sch v ch bn:
HUYN TRANG
30/09/2009
Page 35 of 36
30/09/2009
Page 36 of 36
SINH HC I CNG
M s: 7K787Y8 DAI
In 1300 bn, (Q: 79 ), kh 19 27 cm, In ti Cng ty C Phn In Phc Yn

a ch: ng Trn Ph, th x Phc Yn, Vnh Phc
S KKH xut bn: 922 2008/CXB/7 1873/GD.
In xong v np lu chiu thng 11 nm 2008.
30/09/2009

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Page 1 of 4

Bn quyn thuc B Y t (V Khoa hc v o to)

922 2008/CXB/7 1873/GD

u trnh t biu hin (expressed sequence tags)

Nguyn phn (mitosis)

Phn ng chui trng hp (polymerase chain reaction)

Protein lin kt si n (single strand binding protein)

Virus khm thuc l (tobaco mosaic virus)

Hnh 1.1. Cu trc t bo Prokaryota

2.2. Mng t bo cht

Vai tr ca bao nhy l gip bo v t bo khi cc tc ng bn ngoi (nh s kh hn hoc s tn

2.6. Lng (roi) v khun mao

Hnh 1.2. Cu trc t bo Eukaryota

3.1. Mng sinh cht

Hnh 1.3. S th nghim lai t bo chut v t bo ngi

t phospholipid v pha mnh v y u khng phn cc ra xa. Bi hin tng ny nn cc phn t

Dng lipid th ba l glycolipid. l cc lipid lin kt vi oligosaccarid.

protein phn b tng i ng u, nhng khi c mt tc ng no y ca mi trng (nhit , pH,

Hnh 1.6. M hnh li ni cht c ht (theo I. Livingstone

3.2.1.2. Li ni cht trn

Hnh 1.7. S lp rp hai tiu n v to nn ribosom

V mt cu to, ribosom c kch thc khong 20 - 35nm. c cu to t hai tiu n v ln (60S) v

Da vo kch thc, ngi ta chia cc ti to nn phc h Golgi thnh 3 loi:

Hnh 1.8. Phc h Golgi (theo David H. Cormack)

Vai tr ca th Golgi l thu gp, ch bin, ng gi v phn pht cc sn phm ca li ni cht. Sn

chuyn dn ln mt trans. Trong qu trnh vn chuyn , sn phm s c lp rp vi nhau, c bin

Hnh 1.9. S vn chuyn sn phm t li ni cht c ht n th Golgi

Hnh 1.10. S to thnh v hot ng ca lysosom

ca lysosom b gii phng tc ng ln cc ph nang v gy bnh vim phi.

Phn ln t bo ch c mt nhn. Th nhng cng c nhng trng hp m mt t bo c th c 2, 3

Hnh 1.13. Nhn t bo

3.3.1. Mng nhn

3.4. Mt s thnh phn khc trong t bo Eukaryota

Hnh 1.14. B khung t bo

Hnh 1.15. Vi si actin

Ngoi vai tr to nn b khung, cc vi si cn tham gia vo qu trnh vn ng t bo. Cc dng vn

Ngoi vic kt hp vi vi si to nn b khung t bo, vi ng cn tham gia vo nhiu vai tr khc

Hnh 1.17. Trung t

3.4.3. Lng v roi

Hnh 1.18. Cu to ca lng v roi

4. S VN CHUYN CHT QUA MNG

Hnh 1.19. S vn chuyn nc qua mng

C th thy rng tr trng hp ng trng, cn li cc mi trng nhc trng v u trng u c

4.2. Vn chuyn cc cht khng tiu ph nng lng

Hnh 1.20. nh hng ca Gradient nng n tc vn chuyn cht qua mng

A. Qua cc knh Protein, B. Thm thu trc tip

4.3. Vn chuyn cc cht c km theo tiu ph nng lng

Knh Na+/K+: Cc t bo ng vt c kh nng duy tr nng ni bo ca Na+ rt thp, cn K+ rt

Hnh 1.21. C ch hot ng ca knh Na+/K+

l nhp bo v xut bo. y l mt hnh thc vn chuyn ch ng. Trong , vt cht s c vn

Hin tng nhp bo l s hnh thnh cc bng ni bo do s lm vo v tch ra ca mt phn mng c

Hnh 1.22. Qu trnh nhp bo - th th

Nhp bo th th l dng nhp bo, trong to thnh cc bng nhp bo do s lm vo v tch ra ca

Thnh phn c a ra ngoi nh bng xut bo cng kh a dng. c th l cc cht cn b, cc

Hnh 1.23. M hnh cu to v hot ng ca xynap

S TRAO I CHT V NNG LNG

Trnh by c nh hng ca cc nhn t nhit , pH, nng enzym/c cht, cc cht c ch ln

Trnh by c khi nim v cc giai on chnh ca qu trnh quang hp.

1. NNG LNG SINH HC

1.1. S bin i nng lng t do

s bin i nng lng t do

nng glyceraldehyd 3 -phosphat

[DHAP]: nng dihydroxyaceton phosphat

1.2. Nng lng hot ho