Mc lc

Trang
Chng 1

Cc t s phn ly trong trng hp n

gen
Tm tt
Bi tp

Chng 2

Chng 3

Cc t s phn ly trong trng hp hai

hoc nhiu gen
Tm tt
Bi tp

Chng 4

Gim phn
Tm tt
Bi tp

Chng 5

Di truyn lin kt vi gii tnh

Tm tt
Bi tp

Chng 6

Lin kt gen
Tm tt
Bi tp

Chng 7

Di truyn hc ngi
Tm tt
Bi tp

Chng 8

t bin nhim sc th
Tm tt
Bi tp

Chng 9

Phn tch b bn
Tm tt
Bi tp

Chng 10

Di truyn hc vi khun
Tm tt
Bi tp

Chng 11

t bin phn t v m di truyn

Tm tt
Bi tp

3
8
8
11
17
17
21
23
24
26
30
31
34
41
42
46
50
51
55
61
61
65
70
71
74
80
81
86
88
89
91
95

Chng 12

Kim sot hot ng ca gen

Tm tt
Bi tp

Chng 13

Sinh hc phn t v thao tc vi ADN

Tm tt
Bi tp

Chng 14

Chng 15
Ph lc
Chng 1
Chng 2
Chng 3
Chng 4
Chng 5
Chng 6
Chng 7
Chng 8
Chng 9
Chng 10
Chng 11
Chng 12
Chng 13
Chng 14
Chng 15

Tn s alen v trng thi cn bng

Hardy-Weinberg
Tm tt
Bi tp
Tn s alen do t bin, di nhp v
chn lc
Tm tt
Bi tp
Nhng bi tp kh
Gii p bi tp
Li gii
Li gii
Li gii
Li gii
Li gii
Li gii
Li gii
Li gii
Li gii
Li gii
Li gii
Li gii
Li gii
Li gii
Li gii

95
99
103
103
108
111
112
114
117
117
119
126
131
139
141
148
153
157
164
167
171
175
178
182
186
190

Chng 1
cc t s phn li trong trng hp n gen
A. S c mt ca mt kiu hnh i con cho bit ch c mt
gen.
Nu c b v m u l ng hp t v mt alen, th hai b m cng
ch to ra c mt kiu giao t v cc con biu hin cng mt tnh trng
(ging ht nhau).
AA x AA

hoc

aa x aa

tt c AA

tt c aa

Nu b hoc m l ng hp t tri th cc con u nhn c t

nht mt alen tri, v do vy, tt c cc con u biu hin tnh trng
tri.
AA x aa

AA x Aa

tt c Aa

1/2AA : 1/2Aa

(tt c biu hin tnh trng tri)

trng tri)

(tt c biu hin tnh

V d: Drosophila, php lai gia rui c thn mu en vi rui c thn

mu xm sinh ra tt c rui con c thn mu xm. Hi kiu di truyn
mu thn ca cc con rui ny?
Chng ta thy b m khc nhau v kiu hnh, nhng li thy ch c
mt kiu hnh i con. V con ch biu hin tnh trng tri, ta hy
qui c T = mu xm, t = mu en.
Php lai s phi l:
TT x tt

tt c Tt
3

(xm)
B. S c mt ca hai kiu hnh i con cho bit ch c mt
gen v t nht mt trong hai b m trng thi d hp t.
Hai kiu hnh c th xut hin khi:
1. C b m u l d hp t:

Aa x Aa

3/4A- : 1/4aa
(tri)

(ln)

V d: Khi cho lai hai chut lng en, ngi ta thu c 16 chut con lng
en v 5 chut con lng nu. Hy vit s ca php lai trn.
V c b v m c cng mt kiu hnh m cc con sinh ra li c hai
kiu hnh nn c b v m phi trng thi d hp t v mu en tri
i vi mu nu. Ta qui c B = mu en v b = mu nu.
Bb

Bb

3/4B- : 1/4bb
(en)

(nu)

2. Mt trong hai b m l ng hp t ln v mt l d hp t:
Aa x aa

1/2Aa : 1/2aa
(tri)

(ln)

V d: Php lai gia mt con chut lng en vi mt con chut lng nu

sinh ra 8 chut lng en v 7 chut lng nu. Hy vit s php lai?
Chng ta thy i con xut hin hai kiu hnh vi s lng gn bng
nhau. Nh vy th mt trong hai b m phi trng thi ng hp t v
mt trng thi d hp t.
Bb x bb

1/2Bb : 1/2bb
Trong php lai ny chng ta khng th khng nh c alen no l
tri. xc nh c alen tri, chng ta cn c thm thng tin tng t
nh v d B-1.
4

c. T l phn li 3 : 1 v kiu hnh i con cho bit ch c mt

gen v c hai b m u trng thi d hp t.
V d: Xem li php lai v d B-1:

en x en

16 en : 5 nu
V i con c hai kiu hnh, nn c th ch do mt gen. c hai
kiu hnh, phi c mt hai alen. Nu mu en l tnh trng ln th
php lai s l:
bb x bb
Php lai ny sinh ra cc con ng tnh (mu en) . Kt qu ny
khng ph hp vi bi.
Nu mu en l tnh trng tri v t nht b hoc m ng hp t,
php lai by gi s l:
BB x Bb

tt c BTt c con ci sinh ra li c mu en. Kt qu ny cng khng ph hp.

Php lai phi l:
Bb x Bb

3/4 B- : 1/4 bb
(en)

(nu)

Nu ta thy i con xut hin hai kiu hnh v phn li theo t l 3

: 1 th c hai b m u l d hp t.
Ch : Mu cht ca lp lun ny l c hai b m c cng kiu hnh. Nu
c hai b m c cng kiu hnh v nhn c t s phn li 3:1 i con
th c th l do mt gen (xem mc 2-J v nhng ngoi l).
d. T l phn li kiu hnh 1:1 i con cho bit ch c mt gen
v l php lai d hp t x ng hp t.
Nu bn nhn c t s phn li 1:1 i con, hy kim tra li b m.
Nu b m c kiu hnh khc nhau th php lai ny c th l ph hp.
Tuy nhin, trong php lai ny bn khng th xc nh uc alen no l
tri m cn phi c thm thng tin.
5

V d: Cho giao phn cy c chua thn cao vi cy c chua thn thp.

Kt qu i con nhn c: 26 cy c chua thn cao v 28 cy c chua
thn thp. Hy xc nh kiu gen ca cc cy b m?
Chng ta thy c hai kiu hnh cc cy c chua con vi s lng gn
bng nhau v cc cy c chua b m cng c hai kiu hnh khc nhau.
Do vy, nu qui nh T = alen qui nh tnh trng thn cao v t = alen
qui nh tnh trng thn thp
Php lai l:
Tt x tt

1/2 Tt : 1/2 tt
E. Ba kiu hnh vi t l phn li 1:2:1 i con cho bit ch c
mt gen vi tnh tri khng hon ton (th d hp t c kiu hnh
khc vi c hai th ng hp t).
Trc khi bt u phn tch, bn hy quan st cc kiu hnh. C mt
kiu hnh dng nh c pha trn gia hai kiu kia? Nu ng nh vy, bn
c th c tnh tri khng hon ton. Nhng kh nng khc, bao gm t
nht l c hai gen, s c cp n chng 2. Ta hy xem xt trng
hp mt gen trc.
V d: Khi lai cy hoa mu vi cy hoa mu trng, tt c cc cy F 1
cho hoa mu hng. Cho F1 t th phn, F2 nhn c 11 cy hoa , 23
cy hoa hng v 12 cy hoa trng. Xc nh kiu di truyn v mu
sc ca cc cy hoa?
Mu hng l mu trung gian gia mu v mu trng. Hy cho rng
mu v mu trng l biu hin ca cc trng thi ng hp t . Nu
mt trong hai mu hoc trng l tri th cc cy F 1 s c hoa mu
hoc trng. Nhng thc t khng phi nh vy. V vy khng c mu
no l tri. Mu hng l biu hin ca th d hp t.
Gi C = alen qui nh mu sc ca hoa
Cr = alen qui nh mu
Cw= alen qui nh mu trng
Vy php lai th nht c th l:
CrCr x CwCw
() (trng)
CrCw
(hng)
Php lai th hai l:
CrCw x CrCw
6

(hng) (hng)
1/4 CrCr : 1/2 CrCw : 1/4 CwCw
Lai hai th d hp t bt k no cng s cho t l phn li 1:2:1 v kiu
gen hoc trong trng hp ny s l: 1/4 CrCr : 1/2 CrCw : 1/4 CwCw. Gn mt
na s cy con s c hoa mu hng. Chng ta kim tra li php lai v thy
23/46 l mu hng.
F. T l phn li kiu hnh 2:1 i con cho bit c mt gen v
gy cht mt trng thi ng hp t.
im li chng ta vn ch c hai kiu hnh v do vy ch cp ti
mt gen. ph hp vi iu kin ny th c b v m phi trng thi
d hp t, v nh vy s c cng mt kiu hnh. Trc tin hy xc nh trng
hp nu c hai b m ging nhau.
Xem xt php lai gia hai d hp t:
Aa x Aa
Php lai ny lun cho t l phn li 1/4AA : 1/2Aa : 1/4aa. Thng th
kiu gen AA v Aa s c cng mt kiu hnh v chng ta c t l phn li
3:1. Nhng trong trng hp ny mt kiu gen trng thi ng hp t b
cht. Nu gen gy cht cng c tnh nh mt kiu hnh, th thc ra s
c ba kiu hnh trong php lai ny. Bn c th ch m nhng c th
m bn nhn thy. i con, tnh c nhng c th b cht th 1/2 c
kiu gen Aa v 1/4 c kiu gen aa. Trong s nhng c th sng st th
2/3 c kiu gen Aa v 1/3 c kiu gen aa. Do vy ta thy c t l phn li
2:1.
V d: Drosophila, php lai gia hai rui cnh cong sinh ra 50 rui con
cnh cong v 23 rui con cnh thng. Hy gii thch kt qu thu c bng
s lai thch hp?
Ta thy c hai b m c cng kiu hnh v i con li c hai kiu
hnh. Kt qu ny cho php chng ta ngh ti mt gen v c hai b m
u l d hp t. Thng thng, chng ta ch i t l phn li 3:1 i
con. Cha cn s dng mt php thng k no chng ta cng c th thy
ngay t l 50/23 khc xa vi t l 3:1 m gn vi t l 2:1 hn. Cnh
cong phi l tnh trng tri v c hai b m u c cnh cong v cnh
cong chim u th i con. Do vy, nhng con rui cnh cong ng hp
t phi b cht trc khi sinh ra.
Qui c: Cy = cnh cong, cy = cnh thng. Ta c php lai:
Cycy x Cycy

1/4CyCy : 1/2Cycy : 1/4cycy

(cht)

(cong)

(thng)

G. Nu cc kiu hnh ging nhau li cho ra cc t l phn li khc

nhau trong cc php lai khc nhau, th kh nng c th c nhiu
alen.
V d: Php lai gia c mo mu v c mo mu bc, khi th sinh ra
ton c mo mu , khi th sinh ra 1/2 c mo mu , 1/2 c mo mu
bc v i khi 1/2 : 1/4 trng : 1/4 bc. Php lai gia hai c mo mu
cng sinh ra c khi ton mu , c khi 3/4 : 1/4 bc hoc 3/4
: 1/4 trng. Xc nh kiu di truyn ca cc tnh trng ny?
Trong mi trng hp, chng ta thy rng cc t s phn li u l kt
qu ca di truyn n gen ( 3 : 1; 1: 2: 1 v 1: 1 ). Chng ta c th d
dng gii thch mu v mu bc nu chng ta cho rng l tnh
trng tri i vi bc. Do vy php lai th nht c th l:
RR x rr hoc Rr x rr
Tng t php lai th hai c th l:
RR x R- hoc Rr x Rr
Nhng chng ta khng th gii thch c trng hp mu trng i
con ch vi hai alen. Bi v tt c cc t s ny u l cc t s phn li
trong trng hp n gen, nn chng ta phi d kin mt alen th ba to ra
mu trng khi n ng hp t v mu phi tri so vi mu trng. Nh
rng mt c th lng bi ch c hai alen.
lit k tt c cc kiu gen v kiu hnh c th c, chng ta hy qui
c:
R = v r = khng .
RR, Rrs, Rrw : ( l tri i vi tt c cc alen khc)
rsrs : bc
rwrw : trng
Vy rsrw : s c mu no? Ta cho rng mt trong hai alen s l tri. Trong
trng hp ny, cho rng s tri i vi w th rsrw = mu bc.
Hy tr li vi nhng kt qu trn. Theo nh trn rwrw = trng, do vy,
i con c mu trng, mi b m phi gp t nht mt alen w.
Php lai gia mu v mu bc phi l:
Rrw x rsrw

1/4 Rrs : 1/4 Rrw : 1/4 rsrw : 1/4 rwrw

()

()

(bc)

(trng)

H. Xc sut mt a con c mt kiu gen c th khng ph

thuc vo nhng a con sinh ra trc n.
S di truyn khng cn bit lch s. Mi mt giao t c sinh ra c
lp vi tt c cc giao t khc v do vy c cng mt xc sut xut hin.
Mi khi bn gieo ng xu, bn s nhn c 1/2 s ln xut hin mt nga,
khng cn n cc ln gieo trc c bao nhiu ln mt nga xut
hin.
V d: Mt cp v chng bnh thng c ba ngi con, tt c u b cht
do cn bnh Tay-Sach, mt bnh di truyn do gen ln qui nh. Hi xc
sut m a tr sinh ra tip theo ca cp v chng ny li b bnh?
sinh ra a tr b bnh, mi b m phi d hp v cp gen ny. Do
vy php lai s l:
Tt x Tt
Chng ta bit rng mi hp t c xc sut c kiu gen tt l 1/4. Do
vy xc sut a tr tip theo b bnh cng s l 1/4.
Tm tt
1. Vi mt gen, i con c th c cc t l phn li 3/4 : 1/4 ; 1/2 : 1/2 ;
1/4 :1/2 : 1/4 ; 2/3 : 1/3. Hy xem cc t s phn li i con, nu thy
mt trong cc t s phn li nh trn th ch c mt gen.
2. Hy vit s php lai vi mt gen. Nu bn nhn c cc t s phn
li nh trn th ngha l ch c mt gen.
3. Nu cc t s nhn c khng ph hp vi nhng t s n gen, bn
phi xem xt s di truyn ca cc tnh trng vi s tham gia ca nhiu
hn mt gen. (xem chng 2).
Bi tp
1. Hi chng mt xanh l mt bnh bm sinh rt him gp do sai st
trong qu trnh trao i cht ngi. Axit amin nht nh no
khng c rut non hp th v b bi tit ra ngoi lm cho da mt c
mu xanh.
a. Nu hai v chng bnh thng c mt ngi con b bnh. Hi kiu di
truyn ca cn bnh ny?
b. Hi xc sut a con th hai cng b bnh?
2. u H Lan, php lai gia mt cy u mu vng v mt cy u
mu xanh sinh ra 61 cy mu vng v 57 cy mu xanh. Hy lp s
ca php lai trn?
9

3. Drosophila, php lai gia hai rui cnh ngn sinh ra 43 rui con cnh
ngn v 25 rui con cnh bnh thng.
a. Hi kiu di truyn ca tnh trng cnh ngn?
b. Hi t l phn li i sau nu ta lai mt rui cnh ngn vi mt rui
cnh bnh thng?
c. Hi t l phn li i sau nu ta lai hai rui cnh bnh thng?
4. Nu lai hai ch en vi nhau, kt qu sinh ra 7 ch en v 2 ch nu.
a. Hi alen no l tri?
b. Alen no l ln?
c. Kiu gen ca ch b m?
5. Trong h thng nhm mu ngi, alen A v B tri hn alen O. Xc
nh t l kiu hnh i con nu mt ngi c nhm mu A kt hn
vi mt ngi c nhm mu B?
6. Lai hai con mo ui ngn, i con nhn c 2 con ct ui, 6 con
ui ngn v 3 con ui di. Hy gii thch kt qu thu c bng s
php lai?
7. Drosophila, khi giao phi hai rui mt sinh ra 140 rui mt
v 48 rui mt mu cam. Hy v s php lai v xc nh alen no
l tri?
8. Da vo cc php lai trong bng di y, xc nh kiu gen ca mi
c th:
Php lai

i con

trng-1 x trng-2

6 trng : 1
en

trng-1 x trng-3

5 trng

trng-1 x en-1

3 trng : 3
en

9. Drosophila, tnh trng mt (se+) l tri i vi mt nu (se). Hai

rui mt d hp t c lai vi nhau, mt trong s rui mt i
con li c lai tr li vi rui b m mt . Hi xc sut rui con
ca php lai tr li c mt mu nu l bao nhiu?
10. nga, php lai lp li gia hai nga palamino sinh ra 7 nga
chestnut, 11 nga palamino v 6 cremello.
a. Hy gii thch cc kt qu ny bng cch vit cc kiu gen ca cc
kiu hnh khc nhau?
b. Php lai no to ra t l phn trm nga palamino ln nht?
10

11. th, bn alen ca mt gen to nn mu sc ca b lng. Nhng alen

ny biu hin tnh trng tri theo th t sau y:
C (xm) > cch (chinchilla) > ch (Himalayan) > c (bch tng)
Cp alen cchch v cchc : cho mu xm sng.
Mt th mu xm lai vi cc con th khc v c i con nh trong bng
di y:
Php lai

i con

xm x chinchilla

6 xm, 5 xm sng

xm x xm sng

8 xm, 3 xm sng, 4
Himalayan

xm x bch tng

9 xm, 8 Himalayan

Xc nh cc kiu gen ca cc con th trn?

12. Tc ca ngi c 5 mu, tr mu l: Vng, nu sng, nu, nu
xm v en. Php lai gia nhng ngi c mu tc khc nhau cho kt
qu nh sau:
vng x vng
en

tt c vng

x en

vng x

tt c en
nu

tt c nu sng hoc 1/2

vng : 1/2 nu
nu x nu

tt c nu hoc 1/2 nu xm :

1/2 nu sng
hoc 1/2 nu : 1/4en : 1/4
vng
a. Xc nh cc kiu gen ca tt c 5 mu tc?
b. Nu mt ngi c tc mu nu sng kt hn vi mt ngi c tc mu
nu xm th con ca h s c mu tc no?
c. Nu mt ngi c tc mu nu sng kt hn vi mt ngi c tc mu
en th con ca h s c mu tc no?
13. Mt cy u tng c l mu xanh m th phn vi cy u tng
c l mu xanh nht. F1 nhn c s cy c l mu xanh m v s
cy c l mu xanh nht bng nhau. Nu cho hai cy c l mu xanh
nht F1 th phn vi nhau, F2 nhn c 29 cy l xanh m, 57 cy
l xanh nht. Hy gii thch kt qu thu c?
14. Hai ngi c nhm mu A kt hn, c th sinh ra nhng a con:
a. C nhm mu O khng?
b. C nhm mu B khng?
11

15. Hoa ca cy Chromus baffleus c th c mu , hng hoc trng.

T cc php lai di y, xc nh kiu gen ca cc cy b m:
Php lai

i con

-1 x hng

2/3 , 1/3 hng

-1 x trng

1/2 , 1/2 hng

-2 x hng

1/2 , 1/4 hng, 1/4

trng

-3 x hng

tt c

-3 x trng

tt c

Chng 2
Cc t s phn li
trong trng hp hai hoc nhiu gen
A. S c mt ca 4 kiu hnh i con cho thy c t nht l 2
gen.
Nhc li trng hp nu ch c 1 gen, th ch c 3 kiu gen (v v
vy, nhiu nht c 3 kiu hnh), l: AA, Aa, aa.
V d: u H Lan, trong php lai gia 2 th u ht vng trn, i
con thu c ht vng trn, vng nhn, xanh trn v xanh nhn. Hi
bn cht ca kiu di truyn ny?
Chng ta thy 4 kiu hnh v gi s c 2 gen. Thm ch khng c s lng cc c th i con, chng ta vn c th suy ra tnh trng ht vng,
trn phi tri hn tnh trng xanh nhn. Nu mt gen no l gen ln,
th c th l ng hp t v tnh trng , do vy, s ch xut hin
mt lp kiu hnh.
B. Kiu gen ca b m c th c xc nh da vo cc t s
n gen i con.
Cc gen khng lin kt s phn li mt cch c lp. Bng cch kim tra
li cc t s n gen 1: 0 ; 3: 1; 1: 1 v 1: 2: 1 bn c th xc nh c
kiu gen ca b m.
V d: Xem xt cc php lai di y v xc nh kiu gen ca b m
trong mi php lai:
Php lai

i con
vng,
trn

vng,
nhn

xanh,
Xanh,
trn
nhn
12

vng, trn x vng,

trn

45

15

16

vng, nhn x vng,

nhn

42

15

Xanh, trn x vng,

nhn

31

30

36

33

php lai th nht, da vo t s vng: xanh l 60 : 21, gn vi t s

3 : 1. Do vy mu vng l tnh trng tri v c hai b m u d hp t.
Php lai c th l: Yy Ss x Yy Ss.
php lai th hai, i con khng thy c ht trn. Do vy, ht nhn
i con c th l ng hp t ln hoc c th ht nhn l tnh trng
tri v t nht mt trong hai b m ng hp t. Do thiu 1 php lai,
chng ta khng th xc nh c kiu di truyn ca tnh trng ht nhn.
Tuy nhin chng ta cng c th kt lun c rng mu vng l tnh trng
tri (ta c t l 3 : 1) v c hai b m u d hp t (xem chng 1) . Da
vo php lai 1, chng ta c th kt lun php lai ny l: Yy ss x Yy ss.
php lai th ba, do thiu nhng php lai khc, chng ta thy 61
vng : 69 xanh v 67 trn :63 nhn. C hai u c t s 1 : 1 v do vy
cng c th kt lun rng nhng t s ny l kt qu ca php lai gia
mt d hp t v mt ng hp t ln. Nu ch vi php lai ny chng ta
khng th xc nh c tnh trng no l tri, nhng nu xem xt tt c
cc php lai th php lai ny s l: yy Ss x Yy ss.
C. T s phn li 9:3:3:1 i con cho bit c hai gen khng lin
kt v l php lai gia 2 th d hp t kp vi tnh tri ln
hon ton.
V d: Rui thun chng c thn en, cnh di lai vi rui thun chng c
thn xm, cnh ngn. Tt c rui F1 c thn xm, cnh di. Cho F1 t
phi v F2 nhn c:
88 thn xm, cnh di
32 thn en, cnh di
28 thn xm, cnh ngn
12 thn en, cnh ngn
Hi kiu di truyn ca cc tnh trng ny?
T kt qu ca F1 c th thy rng thn xm, cnh di phi l tnh
trng tri v kt qu ca F2 xc nhn li gi thit ny. Chng ta thy t l
3: 1 cho c xm : en v di : ngn. Tng s rui F 2 l 160 con. Mt t
l l tng 9:3:3:1 s l 90:30:30:10. Kt qu ca chng ta rt gn vi t l
ny. Do vy c th kt lun rng thn xm, cnh di l tnh trng tri v
F1 d hp t.
13

D. T s phn li 3:3:1:1 i con cho bit c 2 gen, mt gen d

hp t c hai b m, gen kia d hp mt bn b (hoc m) v
ng hp t ln bn kia.
V d: Rui mt nu, cnh di lai vi rui mt , cnh ngn. i con
nhn c:
51 cnh di, mt
53 cnh di, mt nu
18 cnh ngn, mt
16 cnh ngn, mt nu
Hi kiu gen ca rui b m?
Trc tin, hy kim tra mi tnh trng mt cch ring r. Ta thy c 104
cnh di : 34 cnh ngn v 69 mt : 69 mt nu. V tnh trng
di cnh thy c t l 3 : 1, do vy cnh di l tnh trng tri v c hai b
m u d hp t. V tnh trng mu mt ta thy c t l 1: 1, nn
khng th xc nh c alen no l tri. Xt ton b chng ta c th kt
lun rng mt trong hai b m ng hp t ln v cp alen ny v bn
kia l d hp t. Nu qui nh L = cnh di; l = cnh ngn; R = mt
v r = mt nu, th php lai l:
Llrr x LlRr
E. S c mt ca 4 nhm kiu hnh i con vi t l tng ng (1:1:1:1) cho bit c hai gen v l php lai d hp t kp
x ng hp t ln kp.
V d: Mt cy thn cao, ht mu xanh th phn vi mt cy thn thp
ht mu vng cho ra:
30 cy thn cao, ht mu xanh
28 cy thn cao, ht mu vng
32 cy thn thp, ht mu xanh
27 cy thn thp, ht mu vng
Hi kiu gen ca cc cy b m?
Chng ta thy c 4 kiu hnh i con, nh vy chng phi c hai gen.
Ta khng th xc nh c alen no l tri, nhng thy 58 cy thn cao :
59 cy thn thp v 62 cy ht mu xanh : 55 cy ht mu vng; Mi
cp tnh trng c t l phn li 1:1. Nu T = thn cao; t = thn thp; Y
= mu vng v y = mu xanh, th php lai l:
Tt yy x tt Yy
Ch rng t s 1:1:1:1 i hi mt bn b m l d hp t v bn kia
l ng hp t cho mi gen. Nhng php lai AaBb x aabb v Aabb x aaBb
u cho t s 1:1:1:1 .
14

F. Trong trng hp tri hon ton, s xut hin nhiu hn bn

kiu hnh i con cho thy c hn hai gen tham gia vo.
Trong trng hp hai gen, vi tnh tri hon ton ch cho ra bn kiu
hnh : tri- tri ; tri- ln ; ln- tri; ln- ln. S kiu hnh cng l 2 n,
trong n = s cp gen. Do vy trong trng hp ba gen, chng ta c tm
kiu hnh.
Ch : Nhng kt lun trn ch nghim ng trong cc trng hp
tnh tri hon ton. Nu chng ta c hai gen v tnh tri biu hin
khng hon ton, th s nhn c 32 kiu hnh t cc kiu gen sau
y: AABB, AABb, AAbb, AaBB, AaBb, Aabb, aaBB, aaBb, aabb (xem
chng 1 v tnh tri khng hon ton).
V d: u H Lan, tnh trng thn cao tri i vi tnh trng thn
thp, tnh trng ht mu vng tri i vi ht mu xanh v ht trn tri
i vi ht nhn. Nu mt cy d hp t v c ba gen t th phn. Hi cc
kiu hnh c th nhn c i con?
Cc kiu gen c th c:
T-Y-S- cao, vng, trn
T-Y-ss cao, vng, nhn
T-yyS- cao, xanh, trn
T-yyss cao, xanh, nhn
ttY-S-

thp, vng, trn

ttY-ss

thp, vng, nhn

ttyyS-

thp, xanh, trn

ttyyss

thp, xanh, nhn

G. Xc sut nhn c mt kiu gen hoc kiu hnh c th

bng tch ca cc xc sut ring.
V d: Xem xt php lai phn F:
TtYySs x TtYySs
Hi xc xut nhn c mt cy c:
1. thn cao, ht vng, trn?
2. thn thp, ht xanh, nhn?
3. thn cao, ht xanh, trn?
Bn c th lp bng Punnett v tnh theo bng, nhng hi di v y
l mt ma trn vi 64 tnh. Ta t php lai:
TtYySs x TtYySs
v xem xt tng gen.
15

Trong cu hi th nht, xc sut nhn c mi tnh trng tri l

3/4. Do vy xc sut nhn c c ba tnh trng tri l 3/4 x 3/4 x 3/4
= 27/64.
Trong cu hi th hai, xc sut nhn c mi tnh trng ln l 1/4.
Do vy xc sut nhn c c ba tnh trng ln l 1/4 x 1/4 x 1/4 =
1/64.
cu hi th ba, xc sut nhn tnh trng cy thp l 1/4 , ht
mu xanh l 1/4 v ht trn l 3/4. Do vy tch cc xc sut l 1/4 x 1/4 x
3/4 = 3/64.
H. Nguyn tc nhn xc sut c th p dng tnh cc t l
cho bt c php lai no, thm ch vi c cc php lai c cc alen
gy cht.
V d: rui, tnh trng cnh cong (Cy) l tri i vi cnh thng (cy), nhng rui cnh cong ng hp t b cht ngay trong trng. Thn mu en
(e) l tnh trng ln i vi thn mu xm (E). Hi i con s c kiu
hnh nh th no nu lai hai rui cng d hp v hai gen ny?
Trc tin vit s php lai:
Cycy;Ee x Cycy; Ee
Sau s dng nguyn tc nhn xc sut.
1/ 4 CyCy ..................................... cht
3/4E- ..................... 3/8 cnh cong, xm
1/2Cycy
1/4 ee ..................... 1/8 cnh cong, en
3/4E- ...................... 3/16 cnh thng, xm
1/4cycy
1/4 ee ...................... 1/16 cnh thng, en
Cn ch rng tng cc t l ny khng bng 1. V chng ta ch
tnh nhng con rui cn sng. Quy ng tt c v mu s = 16 (12/16
rui sng v 4/16 rui cht), sau xc nh t l rui ca php lai:
6 rui cnh cong, thn xm
2 rui cnh cong, thn en
3 rui cnh thng, thn xm
1 rui cnh thng, thn en.
I. Nu php lai gia hai c th c cng kiu hnh t bin sinh ra
i con kiu di, th kiu hnh t bin do hai gen khc
nhau tng tc gy ra.
16

V d: Drosophila, php lai gia hai rui thun chng mt mu c sinh

ra tt c rui con c mt mu (mt kiu di). Hy gii thch cc kt
qu thu c?
Nu cho rng kiu hnh t bin l tri v ch do mt gen qui nh,
th php lai c th l AA x AA, AA x Aa hoc Aa x Aa. Hai kh nng u
cho ra tt c con c kiu hnh t bin; kh nng th ba cho ra con kiu
di v c kiu hnh t bin. Khng c kh nng d on no ph hp vi
kt qu thu c. Do vy cc gen ny khng phi l gen tri.
Nu t bin l tnh trng ln th php lai s phi l aa x aa, tt c
con s c kiu hnh t bin. Khng c kh nng d on no ph hp vi
kt qu thu c, do vy phi c t nht l hn mt gen tham gia vo.
Nu mt trong hai gen ln c lp c th qui nh nn kiu hnh
t bin th php lai s l:
AAbb x aaBB
c c
AaBb
kiu di
J. S tng tc gia hai gen to ra t l phn li 9:3:3:1 v cc dng
bin i ca t l ny i con.
V d: Xt php lai gia cc c th d hp v hai gen A v B. Hi t l
phn li v kiu hnh i con?
Cc kiu hnh c th xut hin:
9/16 A-B3/16 A-bb
3/16 aaB1/16 aabb
Nu mi gen t bin c lp qui nh cng mt kiu hnh t
bin, th cc kiu gen A-bb, aaB- v aabb cng l mt th t bin, v
kt qu cho t l 9:7. Ch rng t l ny gn ging vi t l 1:1 trong
php lai phn tch, nhng trong php lai phn tch b m c kiu hnh
khc nhau.
Nu mt gen t bin c ch s biu hin ca gen kia, th tt c
nhng c th ng hp t ln v mt gen aabb v aaB- s cng l mt
th t bin v kt qu c t l 9:4:3. T l ny gn ging vi t l
1:2:1 trong tnh tri khng hon ton. Nhng trong tnh tri khng hon
ton, cc c th c dng trong php lai c th c kiu hnh trung gian.
Tt c nhng bin i khc ca t l 9:3:3:1 c th cho ra cc t l
9:6:1, 15:1, 12:3:1 v 13:3. Nu ta nhn c hai hoc ba kiu hnh
i con, u tin hy kim tra cc kt qu i vi mt gen. Nu kt qu
17

d on trong trng hp n gen l khng ph hp vi kt qu thu c th

phn tch s tng tc gia hai gen. Nu ta c hai gen, mi gen biu hin
tnh tri khng hon ton th s c t l phn li sau: 1AABB : 2AABb :
1AAbb : 2AaBB : 4AaBb : 2Aabb : 1aaBB : 2aaBb : 1aabb.
K. Lun lun phn tch t n gin n phc tp.
V d: ng, kiu gen A-C-R- qui nh ht c mu. Cc c th ng hp
t ln t nht v mt gen s khng c mu. Hy xem xt cc php lai di
y bao gm cc cy ht c mu v tt c c cng mt kiu gen. Da
vo kt qu thu c, hy tm kiu gen ca cy ht c mu.
B m

i con

c mu x aaccRR
c mu x aaCCrr
c mu x Aaccrr

1/ 2 c mu; 1/ 2 khng
mu
1/ 4 c mu; 3/ 4 khng
mu
1/ 2 c mu; 1/ 2 khng
mu

u tin lp danh sch cc kiu gen c th cho cy ht c mu:

AACCRR
AACCRr
AACcRR
AACcRr
AaCCRR
AaCCRr
AaCcRR
AaCcRr
Kiu gen u tin c th loi ra v tt c con u c mu, bt k
dng th nghim c kiu gen g.
AACCRr c th c loi ra v tt c con trong php lai th nht u c
mu.
AACcRR c th c loi ra v tt c con trong php lai th hai u c
mu.
AaCCRR c th c loi ra v tt c con trong php lai th ba u c
mu.
By gi chng ta ch cn:
AACcRr
18

AaCCRr
AaCcRR
AaCcRr
Th t php lai AACcRr x aaccRR v s dng nguyn tc nhn xc
sut: 1A- x 1/ 2 C- x 1R- = 1/2A-C-R- c mu. C th l kiu gen ny, vy
hy th n php lai th hai AACcRr x aaCCrr : 1A- x 1C- x 1/2R- = 1/2 c
mu. Khng ph hp vi kt qu, do vy kiu gen cn tm khng phi l
AACcRr, . By gi th vi php lai AaCCRr x aaccRR : 1/2A- x 1C- x 1R= 1/2A-C-R- c mu; kiu gen ny ph hp vi kt qu.
Th cho AaCcRR x aaccRR : 1/2A- x 1/2C- x 1R- = 1/4 c mu; khng
ph hp vi kt qu, do vy kiu gen cn tm khng phi l AaCcRR.
Th tip cho AaCcRr x aaccRR : 1/2A- x 1/2C- x 1R- = 1/4 c mu;
khng ph hp, kiu gen cn tm khng phi l AaCcRr.
Vy kiu gen cn tm phi l: AaCCRr. Kim tra li kiu gen ny vi
hai php lai khc.
AaCCRr x aaCCrr :
1/2A- x 1C- x 1/2R- = 1/4 c mu; ph hp.
AaCCRr x AAccrr :
1A- x 1C- x 1/2 R- = 1/2 c mu; ph hp.

Tm tt
1. C bn kiu hnh chng t c hai gen.
2. C nhiu hn bn kiu hnh trong trng hp tnh tri hon ton chng
t c nhiu hn hai gen.
3. Hai gen, mi gen u trng thi d hp t s cho t l phn li 9:3:3:1
i con hoc cc bin i ca t l ny.
4. Nguyn tc nhn xc sut c th c s dng d on cc t l
kiu hnh.
Bi tp
1. Xem xt cc php lai di y Drosophila. Da vo kt qu thu c,
hy cho bit alen no l tri v kiu gen ca b m.
B m

i con
19

nu nu
di
ng
n

di
ng
n

Mt nu,cnh di x mt
nu,cnh di

78

24

Mt ,cnh di x mt
,cnh ngn

30

27

98

95

Mt ,cnh ngn x mt
,cnh di

80

87

Mt ,cnh di x mt
,cnh di

45

16

13
9

51

Mt ,cnh ngn x mt
nu,cnh di

48

42

46

45

2. Xem xt php lai di y giun trn:

ngn, mp x di, thon

47 ngn, mp; 45 di, mp;

53 ngn, thon; 56 di, thon.
Hi kiu gen ca giun b m?
3. Php lai gia chim nhn mu ta vi chim nhn mu trng cho ra
cc con c mu ta. Nu F1 ch c mt mu th F2 li xut hin
cc mu sau y: ta, trng, , xanh da tri v nu. C th kt
lun th no v kiu di truyn ny?
4. Hai ngi b ic kt hn sinh c ba ngi con, tt c u nghe c
bnh thng. Nu hai trong s con ca h kt hn th t l b ic
i chu s l bao nhiu?
5. Drosophila, gen tri qui nh khng c lng (H) cho ra nhng con
rui khng c lng cng ng thi cng l gen gy cht khi trng thi
ng hp t. Mt gen tri khc (S) l gen t ch gen H, s c mt ca
gen S lm cho kiu hnh c lng. Gen S khng biu hin khi t hp vi
hh. Nu hai rui u d hp t c hai gen cho giao phi, hi t l
phn li kiu hnh i con?
6. ngi, mt xanh, bnh bch tng, PKU v ung huyt do bn gen
ln c lp nhau quy nh. Mt ngi n ng bnh thng d hp t
c bn gen kt hn vi mt ngi n b mt xanh, bnh ng huyt
v hp t v gen bch tng v PKU. Hi xc sut a con u
ca h:
a. Biu hin tt c tnh trng tri?
20

b. Mt xanh, bnh PKU v ng huyt?

c. Mt xanh, bnh bch tng v ng huyt?
7. Xem xt hai php lai di y:
Php

Php lai B

lai A
P
AaBb

AaBb x

F1
1 ln

15 tri :

AaBb x AaBb
9:4:3

a. Hi t l kiu hnh F 1 nu P trong php lai A c thay bng php

lai phn tch?
b. Hi t l kiu hnh F 1 nu P trong php lai B c thay bng php
lai phn tch?
8. Trong php lai sau y:
AaBbccDd x AabbCCDd
Hi xc sut nhn c mt c th:
a. Biu hin tt c tnh trng tri?
b. Biu hin tt c tnh trng ln?
c. Kiu hnh ging b?
d. Kiu gen ging b?
e. D hp t v tt c cc gen?
9. Nhiu php lai gia ch en lng ngn v ch en lng di sinh ra
ch con theo t l sau:
36 ch en, lng ngn
39 ch en, lng di
11 ch bch tng, lng ngn
13 ch bch tng, lng di
Hy gii thch cc kt qu ny?
10. Nhiu ngn (mt tnh trng t gp dn n thm ngn tay, ngn
chn) v li un cong l nhng tnh trng tri c lp ngi. Mt ngi
n ng c nhiu ngn v v ng ta th khng. C hai v chng u li
cong nhng b m h u khng. Hi xc sut mt a con ca h
biu hin:
a. Bnh thng, li thng?
21

b. Nhiu ngn, li cong?

11. Hy xem xt cc php lai di y gia mt dng nhn mu ta
thun chng v ba dng nhn trng thun chng.
Php lai 1

Php lai 2

Php lai 3

ta x
trng-1

ta x
trng- 2

ta x
trng-3

F1

18 ta

17 ta

18 ta

F2

95 ta

79 ta

31 trng

36 trng

54 ta
32 trng

28

19
17 xanh da
tri
6 nu

a. Hi kiu gen ca mi con nhn trng?

b. Hi t l phn li kiu hnh nhn c nu F1 ca php lai 2 c lai vi
nhn trng-2?
c. Hi t l phn li kiu hnh nhn c nu F1 ca php lai 3 c lai vi
nhn trng-3?
12. Trong mt php lai gia hai cy hoa trng thun chng, F 1 ton
cy hoa trng. Nu cho F1 t th phn, F2 thu c 104 cy hoa
trng v 24 cy hoa .
a. Hy gii thch kt qu ny bng vit s lai v cc kiu gen ca
tt c cc c th?
b. Hi kt qu nhn c nu nh F1 c lai tr li vi cy b v cy m?
13. chut, bt c c th no c kiu gen cc u b bch tng. Hai cp
alen khc A/a v B/b cng vi C cho ra mu xm, mu en, mu nu
vng v mu scla. Bn dng chut bch tng thun chng khc nhau
c lai vi nhng dng chut xm thun chng (AABBCC). F1 c ton
chut xm. F1 ca mi php lai li c t phi v F2 thu c kt qu
sau:
Dng bch
tng
1

xm

en

44

nu
vn
g
16

scl bch tng

a
0

20
22

31

48

15

21

145

43

46

15

82

Hi kiu gen ca mi dng bch tng?

14. Mt rui c mt mu c thun chng lai vi rui mt nu thun
chng, sinh ra ton rui mt (F 1). Cho F1 lai vi nhau, F2 sinh ra 92
rui mt , 28 rui mt nu, 31 rui mt c v 9 rui mt
trng.
Hy gii thch kt qu thu c?
15. Mt ngi nui ch tin hnh nhiu php lai lp li v thu c cc
kt qu di y:
Php lai

i con

vng x vng

6 vng : 1 en :
1 nu

vng x en

4 vng : 3 en :
1 nu

Hy gii thch cc kt qu ny bng cch vit cc kiu gen ca c ba

con ch?
16. Cho cc kiu gen to nn cc kiu hnh sau:
A-B- : mu
A-bb : mu mn
aaB- : mu ta
aabb : mu trng
Mt gen ln th ba cc gy cht tt c cc c th ng hp t
v mu mn, nhng khng nh hng n cc kiu gen khc. Thng thng C- khng biu hin kiu hnh. Nu hai c th u d hp t v
mi gen em lai vi nhau. Hi t l kiu hnh nhn c i con?

Chng 3
Gim phn

23

A. Gim phn phn chia cc nhim sc th ca b m v sinh ra

cc t bo con mang mt na s nhim sc th c trong t bo
ban u.
V d: t bo soma ca rui gim c 8 nhim sc th (4 cp). Hi
giao t c bao nhiu nhim sc th?
Mt na ca 8 l 4 nhim sc th. Nhng 4 nhim sc th ny phi
th hin c c 4 kiu khc nhau ca nhim sc th b m. S di
y s tm tt khi nim ny. Nhim sc th ca b l hnh c k gch
v nhim sc th ca m khng c k gch. Trong trng hp ny mi
nhim sc th mang 1 gen v c k hiu t A n D.

Sau gim phn cc giao t phi nhn c mt bn copy ca mi nhim

sc th; v d:

Ch rng c nhiu t hp khc nhau ca cc nhim sc th b m.

(Xem KN3-B). Gim phn gii thch s phn ly ca cc alen.
B. S cc t hp nhim sc th khc nhau giao t l 2 n, trong
n = s cp nhim sc th khc nhau (s nhim sc th n
bi).
V d: Mt sinh vt c 3 cp nhim sc th khc nhau l AA', BB', CC'.
Trong nhim sc th ca b c nh du phy v ca m th
khng. Hi sinh vt ny c th to ra bao nhiu kiu giao t?
Mi giao t phi nhn mt nhim sc th A, mt B v mt C. Khi n =
3, chng ta c 8 kiu giao t khc nhau. C th suy ra theo kinh nghim :

ABC
24

ABC'
AB'C
AB'C'
A'BC
A'BC'
A'B'C
A'B'C'
C. Nhng sinh vt c s l b nhim sc th (v d cc th tam
bi) thng bt th.
V d: Mt sinh vt c ba bn sao ca mt nhim sc th A, A' v A''. Hi
cc loi giao t n c th to ra?
Nu nh c ba nhim sc th c gn trn cng mt thoi v sc, mt
t bo s c hai nhim sc th v mt t bo s ch c mt nhim sc
th. Nh vy kh nng giao t c th l: AA' v A''; AA'' v A'; hoc A v
A'A''. S th tinh ca mt giao t c mt nhim sc th (th mt) vi mt
giao t cng l th mt to ra hp t c hai nhim sc th (th hai, hoc
trong trng hp ny l th lng bi). S th tinh ca mt th hai vi mt
th hai to ra hp t c bn bn copy ca nhim sc th v s th tinh
ca mt th hai vi th mt to ra mt th ba. Cng vi s tng ln v s lng nhim sc th, xc sut to ra mt hp t c s lng nhim sc
th chnh xc l rt nh. Hu ht hp t s c mt b nhim sc th
khng cn bng, hoc tha nhim sc th hoc s lung nhim sc
th khng bnh thng. Nhng hp t nh vy thng khng sng c hoc
may mn sng c th cng b d tt bt thng. Trong nhiu trng hp
giao t c s nhim sc th t hn bnh thng s khng sng st c.
C. Mt t bo lng bi, trc khi bc vo gim phn, c s phn t
ADN cng nh s nhim sc th gp i.
Nhim sc th c nhn i l mt iu kin cn thit cho gim
phn. Ngay trc gim phn, mi nhim sc th c hai nhim sc t
(phn t ADN) nh vi mt tm ng. Mt t bo lng bi thc s s c
mt phn t ADN trong mt nhim sc t.
V d: Drosophila c 4 cp nhim sc th. Hi c bao nhiu phn t
ADN trong mt t bo cui gim phn I? Cui gim phn II ?
V cc nhim sc th cc giai on khc nhau. Mt t bo lng bi s
c 8 nhim sc th. Nh rng, mi nhim sc t c mt phn t ADN.

ADN

25

Nhim sc t
Trong pha S, nhim sc t (ADN) c nhn i.
ADN
Tm ng

Nhim sc
t
Do vy, ngay trc gim phn c 16 phn t ADN (8 nhim sc th x
2 si ADN trong mt nhim sc th). Sau gim phn I mi t bo ging
nh sau:
ADN
Tm ng

Nhim sc
t
C 4 nhim sc th trng thi kp. Do vy, sau gim phn I c 4
nhim sc th trng thi kp x 2 si ADN/1 nhim sc th = 8 ADN.
Gim phn II tch cc nhim sc t ra:

ADN

Nhim sc
t

Do vy, t bo n bi to thnh c 4 nhim sc th n, v th c 4

si ADN.
Tm tt
1.Gim phn lm s lng nhim sc th gim i mt na.
2. S lung cc loi giao t khc nhau l 2 n, trong n = s nhim sc
th n bi.

26

Bi tp
1. Ch c 78 nhim sc th.
a. Hi ch c bao nhiu cp nhim sc th?
b. Giao t c bao nhiu nhim sc th?
2. Mt sinh vt c b nhim sc th lng bi l 12. K hiu nhng nhim
sc th ny l: Aa, Bb, Cc, Dd, Ee, Ff.
a. C th xut hin bao nhiu t hp nhim sc th khc nhau giao
t?
b. Xc sut giao t nhn ton b nhim sc th k hiu bng ch
in hoa?
3. Ngi n b c th to ra bao nhiu t hp nhim sc th khc nhau?
4. Ngi c 23 cp nhim sc th. Nu s ADN trng c th hin l c,
hi s ADN :
a. Mt t bo sau gim phn I ?
b. Mt t bo lng bi mi kt thc phn bo nguyn nhim?
c. Mt t bo ngay trc khi bt u gim phn?
5. Mt cy c 4 cp nhim sc th k hiu l: AA, BB, CC v DD. Nu
cy ny t th phn, hi b nhim sc th no trong s di y s c tm thy r ca cy con?
a. ABCD

d. AABBCC

g. AABBDD

b. BCD

e. CCDD

h. AAAABBBBCCCC

c. ABC

f. AABBCCDD

6. Mt sinh vt c 3 cp nhim sc th. Hy k hiu nhim sc th ca

b l P, Q, R v nhim sc th ca m l P',Q', R'. Hi xc sut
giao t ca mt c th c kiu gen PP'QQ'RR' s:
a. Mang tt c nhim sc th gc ca b?
b. Mang tt c nhim sc th gc ca m?
c. Mang 2 nhim sc th gc ca b v 1 nhim sc th gc ca m?
7. La m c 2n = 42 v ng c 2n = 20 nhim sc th. Hy gii thch ti
sao con lai gia ng v la m li lun lun bt dc?

27

8. la mch, b nhim sc th n bi = 7. Bn c th tm c bao

nhiu nhim sc th :
a. Nhn ng phn (nhn sinh dng)?
b. Mt t bo l?
c. Ni nh?
9. C bao nhiu tinh trng c to thnh t:
a. 60 tinh bo s cp?
b. 60 tinh bo th cp?
c. 60 tinh t?
10. ngi, s c bao nhiu trng c to thnh t:
a. 60 non bo s cp?
b. 60 non bo th cp?
11. Mt c th c 8 cp nhim sc th. Nu khng c trao i cho
xy ra, kh nng s c bao nhiu t hp nhim sc th khc nhau giao
t?

28

Chng 4
Di truyn lin kt vi gii tnh
A. hu ht ng vt, gii tnh c xc nh bi loi tinh trng
th tinh vi trng.
nhiu ng vt, gii c l XY v gii ci l XX. Phn bo gim
nhim gii ci to ra tt c cc giao t c mt nhim sc th X. gii
c, mc d nhim sc th X v Y khc nhau, nhng chng kt cp trong
gim phn v nh vo cng mt thoi v sc. Kt qu l mt na s giao
t mang nhim sc th X v mt na mang nhim sc th Y. Nu mt
giao t mang nhim sc th X th tinh vi mt trng th hp t s l XX v
do vy sinh ra gii ci. Nu mt giao t mang nhim sc th Y th tinh vi
trng, kt qu hp t c XY v sinh ra gii c.
B. B truyn nhim sc th X cho con gi v nhim sc th Y
cho con trai.
V gii ci l XX, n phi nhn mt nhim sc th X t mi b m. V
gii c l XY, n phi nhn X t m v nhim sc th Y t b. Trong trng hp cc tnh trng lin kt vi Y s c truyn trc tip t b cho con
trai.
Ch : mt s ng vt, nh nhiu loi chim, bm v.v..., gii c l gii
ng giao t (chng c hai nhim sc th gii tnh ging nhau).
trnh nhm ln, chng c k hiu l ZZ. Gii ci l gii d giao t, v do
vy c hai nhim sc th gii tnh khc nhau k hiu l ZW. Trong trng hp ny, cc tnh trng ln biu hin gii ci F 1 nhiu hn gii
c F1.
V d: Mt chim hong yn trng mang kiu hnh ca mt gen ln lin
kt vi gii tnh c lai vi mt chim hong yn mi mang alen tri. i
con, gii no s biu hin tnh trng tri, gii no s biu hin tnh trng
ln?
Nh rng chim, gii ci l ZW v gii c l ZZ. Hy t ZA = alen tri
v Za = alen ln. Khi gii c biu hin tnh trng tri, kiu gen phi l
ZAZA, gii ci l ZAW. By gi php lai l:
ZAW x ZaZa

ZAZa

ZaW

(c,tri) (ci,ln)

29

C. S khc nhau v kiu hnh gia gii c v gii ci i con

cho thy c s di truyn lin kt vi gii tnh ( lin kt X hoc lin
kt Y).
V d: Php lai gia rui ci mt trng vi rui c mt . i con
tt c rui ci c mt v tt c rui c c mt trng. Hi kiu di
truyn ny?
Chng ta thy c s khc v mu mt gia hai gii, cho thy y l
mt tnh trng lin kt vi gii tnh. Nu mu mt c xc nh bi mt
gen trn nhim sc th Y th tt c rui c s c mt . iu ny
khng ng vi kt qu php lai. Do vy, mu mt phi c qui nh bi
gen trn nhim sc th X. V gii ci c hai nhim sc th X v v tt
c gii c c mt , cho nn mu phi tri i vi trng. V cng
v tt c gii c c cng kiu hnh cho nn m phi ng hp t.
Nu ta gi X+ = v Xw = trng, php lai s l:
XwXw x X+Y
(trng) ()
X+Xw
()

XwY
(trng)

D. M d hp t v mt tnh trng lin kt X phi sinh ra hai dng

con c vi t l tng ng.
V d: Rui ci mt lai vi rui c cng mt . i con tt c
rui ci c mt ; Rui c c mt na mt , mt na mt trng.
Hi kiu di truyn ca cc con rui ny v kiu gen ca cc rui b m?
y l s di truyn lin kt vi gii tnh v c s khc bit v kiu
hnh gia hai gii tnh i con. V tt c rui ci c mt , mu
phi l tnh trng tri, v rui c phi l XRY. Khi mt na s rui c c
mt trng, rui ci m phi d hp t. Chng ta c th vit s php
lai nh sau:
Qui c: XR = , Xr = trng, ta c php lai:
XRXr x XRY
() ()
XRXr XRXR XRY XrY
() () () (trng)
E. Nu tt c cc con ci ca th h con cng biu hin tnh trng
ca b, th tnh trng c di truyn nh l mt tnh trng tri
lin kt X .

30

V d: Gi s mt sinh vt no , mt con c lng en c lai vi

mt con ci lng vng. i con tt c con c c lng vng v tt c
con ci c lng en. Hi kiu di truyn ny v kiu gen ca b m?
y l kiu di truyn lin kt vi gii tnh v chng ta thy c s khc
bit v kiu hnh gia con c v con ci i sau. V con c ch
c mt nhim sc th X, m tt c con ci u c mu en, nn mu
en phi l tnh trng tri. Nu mu en l tri th con ci phi ng
hp t, do vy chng ta thy ch c mt kiu hnh con c. Hy t
XB = en v Xb = nu.
By gi chng ta c th vit s php lai nh sau:

XbXb x XBY

XBXb XBXb XbY XbY

(en) (en) (nu) (nu)
F. T l phn li 3:3:1:1 i con cho thy c hai gen, mt gen
trn nhim sc th thng v mt gen lin kt vi gii tnh.
V d: Da vo cc php lai Drosophila sau y, hy gii thch c s di
truyn ca mi tnh trng v xc nh kiu gen ca tt c cc c th?
P : mt trng, thn en

mt , thn xm

F1 : tt c mt , thn xm
tt c mt trng, thn xm
F2 : 27 mt , thn xm
24 mt trng, thn xm
9 mt , thn en
7 mt trng, thn en
(Khng c s khc bit gia v F2)
Chng ta thy c bn kiu hnh, do vy chng phi c t nht l hai
gen (xem chng 2). Chng ta cng thy khng c s khc bit v mu
sc thn gia c v ci. Do vy chng ta c th kt lun rng tnh
trng mu sc thn do gen nm trn nhim sc th thng qui nh.
Lu rng nu mu sc thn l tnh trng lin kt X th gii c phi
c thn en. V tt c rui F 1 c thn xm, nn thn xm phi tri i
vi thn en.
31

Chng ta thy c s khc bit v mu mt lin quan n gii tnh

F1, c th cho thy tnh trng mu mt lin kt vi nhim sc th X. V
rui ci F1 c mt , nn phi l tnh trng tri, v do vy rui ci
mt trng th h P phi ng hp t v alen ln.
F2, t l phn li gia xm v en l 51 : 16, rt gn vi t l phn
li 3:1 cho mt gen trn nhim sc th thng (xem chng 1). T l phn
li gia v trng l 36:31, rt gn vi t l phn li 1:1 ca mt tnh
trng lin kt vi gii tnh.
Hy t: + = thn xm, dk = thn en, XR = , Xr = trng.
Ta c th vit s sau:
F1:

+/dk XRXr x +/dk XrY

Php lai sinh ra:

1/2 XrXr (hoc XrY) ............... 3/8 xm, trng

3/4 +/1/2 XRXr (hoc XRY)............... 3/8 xm,
1/2 XrXr (hoc XrY) ................ 1/8 en, trng
1/4 dk/dk
1/2 XRXr (hoc XRY).........1/8 en,
Ch : Nu php lai c t theo cch khc:
mt , thn xm x mt trng, thn en
chng ta s nhn c nhng kt qu khc nhau. Tt c rui F 1 ca php
lai c mt , thn xm. F2 chng ta thy:

3/4 mt , thn xm
1/4 mt , thn en

3/8 mt , thn
xm
3/8 mt trng, thn
xm
1/8 mt , thn
en
1/8 mt trng, thn
en

32

Bn c th t li cc php lai xui v lai ngc t thuyt phc v

t l phn li ny. Lu rng trong trng hp ny, chng ta khng bit s di
truyn lin kt vi gii tnh cho n khi nhn c t l phn li F2. Bt
c s khc bit v kiu hnh no gia ging c v ging ci thng ch
ra rng tnh trng lin kt vi gii tnh.
G. Mt gen gy cht lin kt X s lm gim s lng con c i
con.
V d: Rui ci c cnh b khuyt lai vi rui c c cnh bnh thng.
i con nhn c:
35 rui , cnh khuyt
39 rui , cnh bnh thng
33 rui , cnh bnh thng
Hi kiu di truyn ca kiu hnh cnh khuyt?
C th nhn thy ch thy rui c c cnh bnh thng, nhng rui ci
c c cnh bnh thng c cnh khuyt. S khc bit ny gi n tnh
trng lin kt vi gii tnh. V chng ta thy c hai kiu rui ci, cnh
khuyt phi l tnh trng tri. Nu cnh khuyt l tnh trng ln th tt
c rui ci phi c cnh bnh thng. Tt c rui ci nhn mt nhim sc
th X bnh thng t b. Mt s rui ci c cnh bnh thng, cho thy rui
ci th h P l d hp t. T l phn li 2: 1 cho thy rng mt s
rui c b cht. y phi l nhng con rui c nhn nhim sc th
X mang alen cnh khuyt. Cnh khuyt phi l tnh trng tri, nhng gy
cht trng thi ng hp t hoc bn hp t.
Nu XN = cnh khuyt v Xn = cnh thng, chng ta c th t s
php lai:
XNXn x XnY

XNXn

XnXn

XNY

XnY

(khuyt) (bnh thng) (cht) (bnh thng)

Nhng con rui c kiu gen XNY s cht.
H. ng vt c v, s xut hin cc kiu hnh khc nhau cc
t bo khc nhau, hoc cc vng khc nhau trn cng mt con
vt, cho thy tnh trng lin kt vi nhim sc th X .
Thuyt Lyon cho rng, ng vt c v tt c nhim sc th X th
hai trong mt cp thng khng hot ng. Thuyt ny cho rng bt c
t bo no ca con ci cng ch c mt trong hai nhim sc th X hot
ng. Nu s hot ng ny l ngu nhin, chng ta s thy nhng vng
m mt nhim sc th hot ng v nhng vng th tng
ng ca n hot ng. Ngha l chng ta thng gp th khm nhng
33

con ci d hp t. Con c th khm him gp, n phi c kiu gen XXY

(xem chng 6 v 7).
V d: Lai mt con mo c mu vng v mt con mo ci mu en.
mo con, tt c mo c mu en v tt c mo ci c lng vng en
vng vng. Hy gii thch cc kt qu ny?
Kiu hnh khc nhau gia cc gii chng t c s di truyn lin kt vi
gii tnh. Con ci phi ng hp t v tnh trng mu en v tt c con
c c mu en. Cc m en v m vng con ci c th c gii
thch l mt trong hai nhim sc th X con ci ngu nhin khng hot
ng.
Hy t XB = en v XO = vng. S php lai c th l:
XBXB

x XOY

(en) (vng)
XBXO
(en/vng)

XBY
(en)

Nu mu en l tnh trng tri, tt c con F1 s c mu en v kt

qu ny khng ng.
Tm tt
1. Chng ta pht hin s di truyn lin kt vi gii tnh bng cch quan st
s khc bit v kiu hnh gia gii c v gii ci.
2. M d hp t s sinh ra cc con c c t l phn li v kiu hnh l
1:1.
3. B truyn nhim sc th X cho con gi v nhim sc th Y cho con
trai.
4. Mt gen trn nhim sc th thng v mt gen lin kt vi nhim sc
th X s cho t l phn li 3:3:1:1 i con.
5. Gen gy cht lin kt vi gii tnh s lm gim s con c sinh ra.
Bi tp
1. Bnh mu kh ng l do s ri lon ca mt gen ln lin kt vi gii
tnh. Mt ngi ph n bnh thng c cha b bnh mu kh ng kt
hn vi mt ngi n ng bnh thng. H c mt con trai b bnh mu
kh ng. Ngi chng trch v khng chung thu. Nu bn l thm
phn, bn s phn x nh th no, ti sao?
2. nhng ngi Hippy (trc y ngi ta cho rng b tuyt chng), tc
qun l tnh trng lin kt vi gii tnh, tri i vi tnh trng tc thng.
34

Bn hy lp ra mt chng trnh sinh gii tnh ca nhng a

con c th c xc nh ch bng cch xem xt hnh dng ca tc?
3. Mt rui mt , cnh ngn c lai vi rui mt nu, cnh di.
Tt c rui F1 c mt , cnh di. Rui F1 c ni phi sinh ra:

75 cnh di, mt

39 cnh di, mt

23 cnh
mt

37 cnh di, mt
nu

ngn,

14
cnh
mt

ngn,

10
cnh
mt nu

ngn,

Hi kiu di truyn ca nhng gen ny v kiu gen ca rui b m?

4. Drosophila, mt rui lng ngn c lai vi rui lng di. i con
c 42 rui lng di, 40 rui lng ngn v 43 rui lng di.
a. Hi kiu di truyn ca tnh trng lng ngn?
b. Hi t l phn li kiu hnh i con nu bn lai hai rui lng di?
5. Mt mo lng khoang en vng c lai vi mt mo lng vng. i
con nhn c:
: 3 vng, 3 khoang en vng
: 2 en, 2 vng
Hy gii thch nhng kt qu ny?
6. Php lai gia mt chim hong yn mu vng nht vi mt chim
mu xanh sinh ra tt c chim c mu xanh v tt c chim c mu
vng. Hy gii thch cc kt qu ny?
7. Hy xem xt cc php lai di y chim hong yn:
B m

i con

vng x vng

tt c vng

vng x xanh

tt c xanh

xanh x vng

tt c
vng
tt c
xanh

35

Hy gii thch nhng kt qu ny bng cch xc nh alen no l tri

v tnh trng mu sc c di truyn nh th no?
8. Xem xt php lai di y gia rui gim thn vng v thn xm
thun chng:
Php lai
xm x
vng

F1

F2

tt c xm

95 xm

tt c xm

40
vng

45 xm
Da vo nhng kt qu trn, cho bit F1 v F2 nu bn lai mt rui
ci vng thun chng v mt rui c xm thun chng?
9. Mt ngi n ng da c vy sng. ng ta ly mt ngi n b bnh thng. H c bn c con gi, tt c u c da c vy sng v ba cu con
trai da bnh thng. Cc con trai h u ly v da bnh thng v cc
chu ni u da bnh thng. Mt trong s cc c con gi ca h ly
chng c da bnh thng v sinh ra nm chu ngoi, trong hai chu
gi da c vy, mt chu gi da bnh thng, mt chu trai da c vy v
mt chu trai da bnh thng.
a. Bnh da c vy sng c di truyn nh th no?
b. Da vo cu tr li phn (a), hi xc sut a chu sinh ra tip
theo ca cp v chng trn s b da c vy?
10. Drosophila, thn mu vng l mt tnh trng do gen ln lin kt vi
gii tnh qui nh. Cnh ct l mt tnh trng do gen ln nm trn
nhim sc th thng qui nh. Mt rui ci thn vng ng hp t c lai vi mt rui c cnh ct ng hp t.
a.Hi t l phn li kiu hnh F1?
b.Hi t l phn li kiu hnh F2?
c.Gi s php lai u tin c o li: ci cnh ct x c vng. Hi t
l phn li kiu hnh F2?
11. Drosophila, mt rui cI c mt mu cam c lai vi mt rui c
cng c mt mu cam sinh ra tt c rui F 1 c mt c hai gii.
Rui F1 li c cho lai vi nhau v sinh ra:

245 mt mu

119 mt mu
36


75 mt mu cam

201 mt mu
cam

Hy gii thch cc kt qu ny?

12. Drosophila, cnh khuyt l tnh trng tri lin kt vi gii tnh, n l
mt gen gy cht bn hp t; mt mu son l tnh trng ln lin
kt vi gii tnh; thn en l tnh trng ln trn nhim sc th thng.
Mt rui ci cnh khuyt, mt , thn xm c lai vi rui c cnh
bnh thng, mt , thn en sinh ra 1/3 rui ci cnh khuyt, mt
, thn en; 1/3 rui c cnh bnh thng, mt , thn xm v
1/3 rui c cnh bnh thng,mt son, thn xm.
a. Hi kiu gen ca mi rui b m?
b. Hi t l phn li kiu hnh nu lai rui c F 1 vi rui ci F1 cnh
bnh thng? vi rui ci F1 cnh khuyt?
13. Mt rui c mt trng c lai vi rui ci mt nu. Tt c rui F 1 c
mt kiu di. Cho F1 ni phi. Kt qu thu c:

mt 450

mt
230

mt nu 145

mt trng
305
mt nu
68

Hy gii thch cc kt qu ny?

14. enzyme 6- PGD gm c hai tiu n v. c rui gim v ngi, mt
gen lin kt vi nhim sc th X to ra protein ny. Hai alen, 6- PGDA v
6-PGDB , u c c hai loi. C bao nhiu dng 6- PGD khc nhau c
th tm thy con ci d hp t :
a. rui gim?
b. ngi trng thnh?

37

Chng 5
Lin kt gen
A. T l phn ly i lai khc vi t l mong i i vi hai gen
phn li c lp cho thy cc gen lin kt.
V d: Xt php lai

AaBb X aabb

42 A- B- : 39 aabb : 10 A-bb :9 aaBHy xc nh quy lut di truyn v gii thch kt qu ca php lai
trn.
y l php lai phn tch v t l mong i l 1: 1: 1: 1. Khi xt t
l phn ly ca tng cp gen ta c:
A- : aa = 52: 48 = 1: 1
B- : bb = 51: 49 = 1: 1
Nu 2 gen phn li c lp th t l cc t hp hai tnh trng i
con l 1:1:1:1. Kt qu trn cho thy cc gen A v B khng tun theo quy
lut ny. Ta c t l A-B- = aabb = 40%. iu chng t A v B nm
trn cng mt nhim sc th nhng lin kt khng hon ton, v nu hai
gen lin kt hon ton s phi c A-B- = aabb = 50%. Tn s ti t hp RF
(recombination frequency) bng:
S th ti t hp / tng s c th x 100 = 19/100 x 100 = 19%
V d: Xt php lai

AaBb X AaBb

66 A-B-: 16 aabb: 9 A-bb: 9 aaB-

Ta thy F1 khng thu c t l 9: 3: 3: 1. C th ng hp ln v hai

cp gen c t l sai khc nhiu so vi mong i. Nh vy lin kt gen c
38

th xy ra, nhng kh xc nh RF v cc nhm ti t hp v lin kt cha r

rng.
B. Cc lp kiu hnh c tn s cao nht l nhng lp ging b m,
khng phi do ti t hp to thnh.
Ti t hp l s kin him gp. Hu ht cc nhim sc th khng
trao i cho.
V d: Xt php lai trong v d 1 phn A. Cc alen c sp xp th
d hp t th h b m nh th no?
Chng ta lp lun rng A lin kt vi B. D hp t c th l AB/ab
hoc Ab/aB. V kiu gen A-B- v aabb F1 c tn s cao nht nn chng
phi c to ra t giao t mang cc nhim sc th khng c trao i
cho ( ti t hp). V vy, cc alen b m d hp t phi trng thi
ng (cis), tc l AB/ab.
C. Lin kt gen khng lm thay i s lp kiu hnh mong i m
ch lm thay i tn s ca mi kiu hnh.
V d: Xt li v d1 phn A. Chng ta vn thy c 4 kiu hnh. Tn
s ca mi kiu hnh xut hin ph thuc vo khong cch gia hai gen
lin kt. Tng t, nu c ba gen lin kt, trong php lai phn tch ta vn
thu c 8 kiu hnh.
D. Trong mt php lai phn tch , vic c hai lp kiu hnh c
tn s ln bng nhau v hai lp kiu hnh c tn s nh bng nhau
cho bit trong c gen lin kt.
V d: Xem li v d 1 phn A. Chng ta suy lun l lin kt gen v
t l cc lp kiu hnh b thay i. Kim tra tip kt qu th thy rng
A-B- v aabb c tn s bng nhau, A-bb v aaB- c tn s bng nhau nhng
nh hn. Nu cc gen khng lin kt, chng ta c t l mong i l
1:1:1:1.
E. Cc lp kiu hnh tng h lun xut hin vi tn s bng nhau.
Ti t hp l kt qu ca s trao i cho cc on nhim sc th tng ng. Cc s kin to ra mt lp c kiu hnh ti t hp cng ng
thi to ra mt lp khc tng ng. Nu chng ta v hai nhim sc th tng ng v biu din s trao i cho nh sau:
A

B
a

th cc nhim sc th thu c s l:
A

Nu khng c ti t hp th ch to ra hai loi giao t l:

39

F. Trong cc php lai lin quan n ba gen du d hp t, cc lp

kiu hnh to thnh t trao i cho kp lun c tn s nh
nht
V d: Xt php lai sau: AbC//aBc X abc//abc.
Cc lp kiu hnh no c tn s nh nht?
Ti t hp l mt s kin rt him. Tn s hai s kin rt him
cng xy ra trong qu trnh gim phn xp x bng tch tn s cu mi
s kin ring l. V vy trao i cho kp cng l mt s kin him.
V nhim sc th c trao i cho kp:
A

b
a

C
B

Cc giao t l kt qu ca trao i cho kp ny l: ABC v abc, v

hai lp kiu hnh ABC v abc s tn s nh nht.
Trong mt vi trng hp vi ba gen lin kt chng ta s ch thy 6 hoc
7 lp kiu hnh. Nu ch c 6 kiu hnh v hai lp xut hin l tng h,
th cc lp khng xut hin l nhng lp kiu hnh do trao i cho
kp.
V d: AbC/aBc X abc/abc to ra:
AbC: 40
aBc: 38
ABc: 6
abC: 5
aBC: 5
ABC: 3
Abc: 3
Tn s trao i cho kp quan st c l bao nhiu?
Nhng kt qu ny cn phi xem xt k, v chng ta thy s lng gn
nh tng ng 5 lp kiu hnh khc nhau.
Cc lp abC v aBC mc d bng nhau nhng l nhng lp khng tng
h nhau. Lp abc khng xut hin. V vy chng ta c th sp xp li
theo nhm nh sau:
AbC v aBc: 78
ABc v abC: 11
aBC v Abc: 8
40

ABC v abc: 3
Cc lp cui cng c tn s nh nht v v vy l cc lp do trao
i cho kp. Do vy tn s trao i cho kp l 3 x 100/100% = 3%.
Lu rng cc lp tng h phi c nhm li vi nhau.
G. Trao i cho kp lm thay i s lin kt ca gen nm gia.
V d: Mt th d hp t v ba gen lin kt nhau. A, B, C c lai vi mt c
th ng hp t ln v to ra: Abc v aBC: 125, ABC v abc: 35, ABc v
abC: 30, AbC v aBc: 10. Cc gen th d hp t sp xp nh th no v
trt t cc gen?
Cc lp c tn s cao nht l Abc v aBC, phi l nhng kiu hnh
c to ra t cc nhim sc th khng c trao i cho. Do vy b m
c kiu gen Abc/aBC. Lu rng cc ch ci biu hin cc gen ny khng
biu din trt t gen m ch cho thy gen tri A lin kt vi hai gen ln b
v c.
Chng ta s dng cc lp c tn s nh nht ( c trao i cho kp )
xc nh trt t cc gen. Cc lp ny l AbC v aBc. So snh cc lp
ny vi cc lp khng c ti t hp, ging b m: Ab v aB vn lin kt vi
nhau ging nh kiu gen ca b m; gen C c trao i gen A v
C cng cng nm trn mt nhim sc th. Do vy C phi nm gia. Chng
minh kt lun ny bng cch v mt trao i cho kp:
A

s to ra cc giao t ACb v acB.

H. lp bn , u tin phi xc nh trt t cc gen, sau
tnh khong cch gia gen gia v cc gen hai u.
V d: Xt 3 gen lin kt ng: +/b, +/lg, +/v. Mt php lai phn tch
gia th d hp t v 3 gen v th ng hp t ln to ra th h con nh
sau:
165 +vlg
125 b++
64

b+lg

56

+v+

37 ++lg
33 bv+
11 +++
9

bvlg

Tng s: 500 c th.

41

Xc nh cu trc di truyn ca th d hp t; xc nh trt t gen

v khong cch bn gia cc gen; tnh tn s trao i cho kp.
Chng ta tin hnh theo cc bc sau:
1. Xc nh nhng th khng phi do trao i cho c tn s
cao nht. Trong trng hp ny l +vlg v b++. V vy c th d hp t
ny l +vlg/b++.
2. Xc nh trt t bng vic so snh cc lp do trao i cho kp
to ra vi cc lp ging b m. Trong php lai ny +++ v bvlg c tn s
nh nht. V v v lg nm cng nhau nh kiu gen b m, ch c b b
trao i, vy b phi nm gia. Chng ta v li kiu gen ca th d hp
t v+lg/+b+:
v

3. Tnh RF gia v v b bng vic s dng tt c cc t hp cha b

v v ( v +, +): (37 + 33 + 11 + 9)/500 x 100 = 18 n v bn .
4. Tnh RF gia b v lg bng vic s dng cc t hp ng thi
cha b v lg: (64 + 56 + 11 + 9)/500 x 100 = 28 n v bn .
Vy ta c th v bn nh sau:
v

18

28

lg

5. Tnh h s trng lp CC (coefficient of coincidence)

CC = Trao i cho kp thc t / Trao i cho kp l
thuyt
v d trn, tn s trao i cho kp l thuyt = (0,28)(0,18) =
0,05. Vy s c th c trao i cho kp theo l thuyt = 0,05 x 500 =
25. S c th c trao i cho kp thc t = 20.
Vy CC = 20/25 = 0,8
I. d on cc tn s kiu hnh khi bit khong cch gia cc
gen, trc ht ta tnh tn s trao i cho kp theo l thuyt,
sau ly khong cch bit tr bt i.
V d: Xt ba gen lin kt theo trt t sau:

A_____30____B__20___C

Nu mt th d hp t v 3 gen AbC/aBc c lai vi abc/abc th t l

cc kiu hnh theo l thuyt l bao nhiu? Gi s rng tn s ca cc c
th c trao i cho kp l tch cc tn s trao i cho n ( khng
c nhiu ).
V y l php lai phn tch, nn tn s ca cc giao t s bng tn
s ca cc kiu hnh.
1. Tnh tn s ca trao i cho kp.

42

Cc lp ny s l ABC v abc. Vy tn s trao i cho kp = 0,3 x 0,2

= 0,06.V ti t hp l tng h nn 1/2 x 0,06 l tn s ca mi lp ABC v
abc, v bng 0,03.
2. Tnh tn s trao i cho n gia A v B.
Ta c tn s trao i cho gia A v B = 0,3, tn s ny bng tng
tn s cc trao i cho n v tn s trao i cho kp, v vy:
Tn s trao i cho n - tn s trao i cho kp = tng tn s
ca cc trao i cho n. Vy tn s trao i cho n gia A v B
= 0,3- 0,06 = 0,24. Tn s ca mi lp ABc v abC s bng 0,12.
3. Tnh tn s ca trao i cho n gia B v C:
0,2 - 0.06 = 0,14. Tn s mi lp Abc v aBC s bng 0,07.
4. Tnh cc c th to ra do lin kt hon ton bng cch ly 1 tr
i tt c cc c th c ti t hp
1 - ( 0,24 + 0,14 + 0,06 ) = 1 - 0,44 = 0,56. Tn s mi lp AbC v
aBc s l 0,28.
Trong mt s trng hp, vic xy ra mt trao i cho s c ch hoc
khuch i mt trao i cho th hai bn cnh. Hin tng ny c gi l
nhiu (I - interference) v c tnh: I = 1 - CC. Trong trng hp trn, ta gi
s rng I = 0,02, hy tnh cc tn s mong mun ( theo l thuyt ).
tnh ton, trc ht tn s trao i cho kp theo l thuyt phi
c bin i.
V I = 1 - CC, vy CC = 0,8.
CC = Tn s trao i cho kp thc t / Tn s trao i cho kp l
thuyt
Ngha l : 0,8 = tn s trao i cho kp thc t/ 0,06
Suy ra tn s trao i cho kp thc t = 0,8 x 0,06 = 0,048.
V vy tn s cc lp trao i cho kp bng 0,048 v tn s trao i
cho n gia A v B l
0,3 - 0.048 = 0,252
Tn s trao i cho n gia B v C l
0,2 - 0,048 = 0,152
V tn s cc lp khng c trao i cho l 0,548.
J. S c mt ca 4 lp kiu hnh c tn s ln bng nhau v 4 lp
c tn s nh bng nhau cho thy c hai gen lin kt v mt gen
phn li c lp.
V d: Mt rui ci d hp t v 3 gen: +/sc, +/ec, +/vg c lai vi rui c
ng hp t ln. Tt c cc t bin u ln. Th h sau thu c:
43

sc ec vg 233
+++

239

sc ec + 241
+ + vg 231
sc + vg 12
sc + +

14

+ ec +

14

+ ec vg

16

Tng s 1000
Gii thch v v s lai ph hp.
Chng ta bit rng nu 3 gen khng lin kt, khi lai phn tch s
cho t l 1: 1: 1: 1: 1: 1: 1: 1. V php lai trn khng cho t l ny nn
t nht c hai gen lin kt. Bn lp c t l cao l kt qu ca s phn ly
c lp. Nu tt c ba gen lin kt vi nhau, chng ta ch c hai lp kiu
hnh vi t l cao. xc nh gen no lin kt vi gen no, ta kim tra
hai gen mt. Hy xem xt ec v vg trong cc lp t l cao:
ec vg 233
+ + 239
ec + 241
+ vg 231
T l ny tng ng t l: 1: 1: 1: 1. Do vy ec v vg khng lin kt,
chng phn li c lp.
Xt sc v ec:
sc ec = 233+ 241 = 474
+ + = 239+ 231 = 470
+ ec = 14 + 16

= 30

sc + = 14 + 12

= 26

Ta thy ch c hai t hp lp kiu hnh c t l cao. V vy sc v ec

lin kt khng hon ton. C 4 lp kiu hnh c tn s thp l cc th ti
t hp gia sc v ec.
Vy tn s trao i cho gia sc v ec l:
( 12+14 + 16 + 14)/ 1000 x 100% = 5, 6% = 5, 6 n v bn
K. rui gim, t l 5: 1: 1: 1 cho thy hai gen lin kt cch xa
nhau trn 50 n v bn .
V d: Hai rui gim d hp t v hai gen quy nh tnh trng ln lin kt
nhau l mt nu v thn en lai vi nhau thu c:
44

52 thn xm, mt
9 thn xm, mt nu
10 thn en, mt
9 thn en, mt nu
Gii thch kt qu trn v lp bn di truyn.
Quy c +: mt , br: mt nu, +: thn xm, b: thn en
Php lai trn s l: br b/+ + X br b / + +
Chng ta thy hai gen lin kt nhau nn kt qu ca php lai khc vi
t l 9: 3:3:1. Lu rng php lai ny khng phi l php lai phn tch.
Do vy s kh tnh RF. Nh rng rui gim c khng c trao i cho.
V th giao t ca con c ch c hai loi: br b v ++, cn con ci to ra
4 loi giao t: + +, + b, br + v br b. Nu hai gen ny cch xa nhau trn
50 n v bn , chng c xem nh l chng phn li c lp nhau v
c 4 loi giao t c to ra vi tn s bng nhau:
++

+b

br +

br b

++

+ +/+ +

+ b/+ +

br +/+ +

br b/+ +

Br b

+ +/br b

+ b/br b

br +/br b

br b/br b

T l thu c l:
5 + +/ + -

thn xm, mt

1 + b/br b

thn en, mt

1 br + / br b thn xm, mt nu
1 br b/ br b

thn en, mt nu

Ch : Nu hai gen lin kt kiu i (hai alen ln nm trn hai nhim

sc th tng ng khc nhau, v d Ab/aB), t l thu c l: 2 kiu di:
1 nu: 1 en. Vit s lai t khng nh iu ny l ng.

L. Khong cch trn bn c th cng k tip, do vy ta c th

xc nh trt t ca nhiu gen.
V d: Cho khong cch gia cc gen nh sau, lp bn cc gen :
45

OR

RA

13

RG

MR

GA

OG

MG

12

GN

10

ON

18

Ta bt u vi cc gen c khong cch ln, chng phi nm hai pha

u ca nhim sc th. Sau sp xp cc khong cch gia cc gen
to ra cc bn gi ln nhau.
O______18_______N
M____12_____G___10____N
Ta c R G = 5. Nu R nm bn phi G th M R = 17. iu khng
ng. Vy R phi bn tri G.
M

R 5

10

Nu O nm bn phi N th O R c khong cch rt ln, nhng iu

ny khng ng. V vy O nm bn tri N. V O R = 3 v O G = 8
nn O phi nm gia M v R.
M

O 3R 5 G

10

V M R = 7, vy M O phi l 4. A c th nm c hai pha ca G.

Nu A bn phi G th R A = 13. Bn hon chnh nh sau:
M 4 O 3 R 5

A2 N

Tm tt
1.Lin kt gen lun c pht hin bi s sai khc so vi t l mong i
(nh lut Menen ).
2.Tn s trao i cho c tnh bng cng thc:
s c th do ti t hp x 100%/ tng s c th i con.
3. Cc lp kiu hnh do trao i cho kp to ra lun c tn s nh nht.
4. Lun phi nhm cc lp kiu hnh tng h vi nhau.
5. Khong cch bn gia cc gen nm hai u c tnh bng tng
cc khong cch c chia bi cc gen nm bn trong hai gen .
6. T l 5: 1: 1: 1 Drosophila cho thy hai gen trong nhm lin kt cch
xa nhau 50 n v bn .
7.Trong cc php lai lin quan n ba hoc nhiu gen, trong c mt
gen khng lin kt s to ra 4 lp kiu hnh c tn s ln v 4 lp kiu
hnh c tn s nh vi t l bng nhau.

46

Bi tp
1. chut, gen ng mu (khng m) V l tri so vi gen m v; gen
lng ngn S tri hn gen lng di s. Lai chut ci d hp t v c hai
gen trn vi con c ng hp t ln, thu c:
75 lng m, di
66 ng mu, ngn
10 ng mu, di
7 lng m, ngn
a. Hai gen trn c lin kt vi nhau khng ?
b. S sp xp cc gen trn nhim sc th chut ci nh th no?
c. Nu cc gen lin kt, xc nh khong cch gia cc gen.
2. mt c th gi nh, cc alen trn nhim sc th thng l cao (T),
v v nhn (R) tri so vi thp (t) v trn (r) tng ng. Cc cy m d hp
t cao, v nhn c lai vi cy b thp, trn v thu c th h con nh
sau:
cao, nhn

105

cao, trn

236

thp, nhn

273

thp, trn

110

a. Hai gen ny c lin kt vi nhau khng? Ti sao?

b. cc c th ci d hp t, cc alen quy nh tnh trng cao, trn c sp xp nh th no? Gii thch.
c. Tnh tn s trao i cho gia hai gen ?
3. V bn di truyn da vo cc tn s ti t hp sau:
c b: 9

d e: 10

a b: 3

a f: 7

c f: 13

f d: 8

b d: 12
a e: 25

4. Xt php lai sau gia c th c ba gen lin kt DeF/dEf X def/def.

Xc nh trt t gia cc gen khi thu c kt qu php lai nh sau:
DeF

76

def

30

dEf

82

DEF

24

Def

15

DEf

1
47

dEF

18

deF

Tng s 250 c th.

5. Trong mt c th gi nh, con ci thn b, lng trng, thng c lai
vi con c thn mnh, lng en, qun to ra F 1 thn mnh, lng
trng, thng. Cho con ci F 1 giao phi vi con c thn b, lng en,
qun thu c i con:
thn mnh, lng trng, thng

169

thn mnh, lng en, thng

19

thn mnh, lng en, qun

301

thn b, lng trng, qun

21

thn mnh, lng trng, qun

thn b, lng en, qun

172

thn b, lng en, thng

thn b, lng trng, thng

304

Tng s

1000

a. Lp bn di truyn xc nh trt t cc gen v khong cch

gia chng.
b. Tnh h s trng lp CC.
6. rui gim, tnh trng mt hnh bu dc (b), thm (cd) v thn
en (e) l cc alen ln lin kt vi nhau. Nu con ci mt bu dc,
thm c lai vi con c thn en ng hp t, ta thu c tt c rui
con F1 u c kiu di. Cho cc con ci F 1 d hp t lai vi cc con c
mt bu dc, thm thn en, thu c th h sau gm 1000 con
t l nh sau:
bu dc, thm

440

thn en

443

bu dc, thn en
thm

32
34

en, thm
bu dc

23
25

bu dc, thm, en
kiu di

a. Xc nh s sp xp ca cc gen trn nhim sc th ca con ci F1.

b. V bn di truyn ca ba gen trn.

48

7. Lai rui gim ci mt nu, cnh b vi rui c kiu di. Rui con F 1
u c kiu di. Giao phi gia cc rui con F1 thu c:
110 kiu di
17 cnh b
13 mt nu
20 cnh b, mt nu.
Gii thch kt qu.
8. c tr, mt con ci dng trn, nht, c mu c lai phn tch vi con
c dng dt, khng nht, khng mu v to ra th h con nh sau:
trn, khng mu, nht

145

trn, khng mu, khng nht

trn, c mu, nht
trn, c mu, khng nht
dt, c mu, nht
dt, c mu, khng nht

38
138
33
40
133

dt, khng mu, nht

35

dt, khng mu, khng nht

148

Tng s:

710

a. Xc nh nhng gen no lin kt vi nhau?

b. Xc nh kiu gen ca c th d hp t, ch ra cc alen no c th
nm trn cng mt nhim sc th.
c. Tnh khong cch gia cc gen lin kt.
9. ra Nijna Turtle, tnh n ho (A) l tri so vi khng n ho (a), chm
chp (B) l tri so vi nhanh nhn (b), in di (C) l tri so vi khng
in (c). Mt ra n ho, nhanh nhn, bnh thng c lai vi ra khng
n ho, chm chp v b in. Cc con F1 c lai vi ra khng n ho
v nhanh nhn bnh thng, thu c:
n ho, chm chp, in:

192

n ho, chm chp, bnh thng:

208

n ho nhanh nhn, in:

780

n ho, nhanh nhn, bnh thng:

810

Khng n ho, chm chp, bnh thng: 790

Khng n ho, chm chp, in:

820

Khng n ho nhanh nhn, in:

195

Khng n ho, nhanh nhn, bnh thng:

205
49

Tng s:

4000

Hy gii thch kt qu trn v v bn nhim sc th.

10.Hai gen Q v R lin kt nhau v cch nhau 15 n v bn . Nu
mt con ci d hp t c em lai phn tch, hy tnh tn s ca cc
kiu hnh th h con.
11. Xt c th c ba cp gen ln lin kt vi nhau: fz, b, stw cng nm
trn nhim sc th s 2 rui gim. Khong cch gia 3 gen nh sau:
fz

15

b 7 stw

Nu lai con ci + b stw/fz + + vi con c fz b stw/fz b stw. Cho CC

=0,6. Hy tnh tn s cc lp kiu hnh?
12. Cc gen a v b lin kt nhau v cch nhau 10 n v bn . Cc
gen c v d lin kt nhau v cch nhau 5 n v bn trn nhim
sc th khc vi nhim sc th c cha a v b. Mt c th ng hp t
v cc alen tri c lai vi c th ng hp t v tt c cc gen ln. Cc
c th F1 c lai phn tch. Lp kiu hnh no s c tn s nh nht?

50

Chng 6
di truyn hc ngi
Nhiu khi nim trong chng ny c chng 1, 2 v 4. Chng ta
nn c cc chng ny trc khi tip tc. Cc ph h ngi thng khng
cha thng tin xc nh m hnh di truyn mt cch r rng.
A. Cc c th biu hin tnh trng tri thng l th d hp t v
alen t bin.
Tt c cc t bin u l nhng s kin him. Xc sut mt c
th c c 2 alen t bin tri l bnh phng tn s c mt alen t
bin tri. Hn na, vic c c 2 alen t bin tri l qu kh khn v s
lm c th ng hp kh sng st c.
B. Nu mt tnh trng ngt qung mt th h, n c th l tnh
trng ln.
V d: Trong ph h sau, quy lut di truyn c th c l g?

II

III
IV
Chng ta thy rng tnh trng b ngt qung qua cc th h, do vy
n c th c xc nh bi gen ln nm trn nhim sc th thng. Nu
n lin kt vi X, con trai th h III c th mang tnh trng nhng thc
t th khng phi.
51

C. Mt tnh trng ln lin kt X c th xut hin 1/ 2 s con trai

ca ngi m bnh thng.
V d: T ph h sau y, xc nh quy lut di truyn ca tnh trng:

I
II
Tnh trng trn l ln v c hai b m u khng biu hin tnh
trng . N c th lin kt vi nhim sc th X v khong 1/ 2 s con
trai biu hin tnh trng ny. Nu gen quy nh tnh trng l gen ln
nm trn nhim sc th thng th chng ta d tnh c khong 1/4 s
con biu hin tnh trng. iu c th l ng ph h trn. V vy,
tnh trng ny c th l gen ln nm trn nhim sc th thng hoc lin
kt X.
D. Mt tnh trng tri do gen trn nhim sc th thng quy nh
c th xut hin 1/ 2 s con c hai gii ca mt ngi bu hin
tnh trng .
V d: Xc nh quy lut di truyn t ph h sau:

I
II
III
Tnh trng ny c th c xc nh bi gen ln nm trn nhim sc
th thng v ngi m th h I c th trng thi d hp t. Cng c th
l do gen tri nm trn nhim sc th thng v 1/ 2 s con c biu hin
tnh trng . Kh nng sau d xy ra hn v t bin l nhng s kin
him. Nu n l gen ln, kh nng mt th ng hp ln giao phi
vi mt th d hp t l rt him.
E. Tnh trng tri lin kt X c th xut hin tt c con gi ca
ngi cha biu hin tnh trng ln .
V d: Xc nh quy lut di truyn ca tnh trng da vo ph h sau:

I
II
III

52

Lu rng tt c con gi ca ngi b b bnh u biu hin tnh trng

, do vy, n c th l gen tri lin kt X. N cng c th l do gen ln
trn nhim sc th thng (aa x Aa), hoc tri trn nhim sc th thng
(Aa x aa).
F. Tnh trng lin kt Y s ch xut hin con trai ca ngi b
biu hin tnh trng .
C th c lun lun mang nhim sc th Y v truyn li cho th
h con gii c, do vy bt k tnh trng no do gen nm trn Y cng
ch xut hin con c.
V d: Da vo cc ph h sau, xc nh quy lut di truyn c th c:

I
II
I III
II
III
O
IV
Chng ta thy ch c cc con trai mang tnh trng mi th h .
Mt tnh trng tri do gen trn nhim sc th thng c th biu hin nh
trn, nhng thng thng th mt s em gi cng biu hin tnh trng. Vy
gen quy nh trng ny chc chn nm trn nhim sc th Y.
G. Trong cc ph h, ta u coi rng nhng ngi bnh thng l
ng hp t tr khi c bng chng ngc li.
Cc t bin thng l s kin him, v khng chc chn rng mt
ngi bnh thng li l d hp t khi ta khng c thng tin chc chn.
V d: Tn s con b bnh ca cp v chng th h II l bao nhiu?

I
II
III

53

u tin th suy lun ra quy lut di truyn. V hai c th bnh thng

sinh ra con biu hin bnh, tnh trng phi l ln. N khng th lin
kt X, v nu lin kt X th ngi n ng th h I phi mang tnh
trng v con gi cng biu hin bnh. V vy tnh trng phi
do gen ln nm trn nhim sc th thng v c th II-3 phi l ng hp
t. V tnh trng ny t gp nn khng chc chn rng ngi v i II l
d hp t. Nu chng ta gi s ngi v th h II l ng hp t tri th,
tn s biu hin bnh i con l bng 0 (aa x AA)
H. tnh tn s ca mt kiu gen no , ta s dng phng
php nhn xc sut.
V d: Mt ngi ph n b bnh bch tng c hai ngi con gi bnh thng, mi ngi ly mt ngi chng bnh thng v sinh ra con bnh thng.
Tnh tn s sinh ra a tr b bnh bch tng nu hai a chu kt
hn.
Hy v ph h v xc nh kiu gen. Gi s tt c nam gii th h I
v II l ng hp t th c hai con gi phi c kiu gen d hp t, Aa, v
tt c nhng ngi chng u c th c kiu gen AA. V vy, mi a tr
th h III u c 50% kh nng l d hp t. Kh nng c hai a tr III
u d hp t l 1/ 2. 1/ 2= 1/ 4.

I
II
III
IV

Nu hai dng d hp t giao phi vi nhau, c 1/ 4 s c th con c

kiu gen ng hp ln. V vy kh nng c bnh bch tng th h IV l
1/ 4. 1/ 4= 1/ 16.
I. Tn s ca mt hoc nhiu bnh bng tng xc sut ca mi
bnh.
V d: Gen quy nh bnh bch tng v PKU khng lin kt (nm trn 2
nhim sc th thng khc nhau) v l tnh trng ln ngi. Nu mt ngi ph n v mt ngi n ng u d hp t v c hai tnh trng trn
kt hn, hy tnh tn s a con u tin ca h mc bnh bch
tng v PKU.
54

u tin, ta tnh tn s ca ring mi tnh trng. Gi gen quy nh

bnh PKU: p, bnh thng: P, bch tng: a, bnh thng: A. S lai l:
PpAa x PpAa. Mi tnh trng c tn s xut hin bnh i con l 1/ 4.
i vi c 2 tnh trng, chng ta cng cc kh nng li =1/ 4 + 1/ 4 = 1/ 2.
J. Vi n con, tn s s con c kiu hnh ny v t con c kiu
hnh kia l (n!/(s! t!)) (p)s (q)t, trong p v q l tn s xut hin
mi kiu hnh tng ng.
V d: ngi kh nng un li l tnh trng tri. Nu 2 b m d hp t c
5 ngi con, tn s 3 ngi con trong s c kh nng un li l bao
nhiu?
Trc ht, xc nh tn s ca mi kiu hnh :
Tn s ngi un li = 3/ 4 = p. Tn s ngi khng un li = 1/ 4 = q.
V 3 ngi c kh nng un li nn ta c 2 ngi khng un li, tc l s
=3, t = 2. Ch rng s + t = n v p + q = 1. Thay s vo cng thc trn:
P = (5!/(3! 2!)) (3/4)3 (1/4)2 = 0,264, hoc khong 1/4 .
Nu t 2 tnh trng tr ln, cng thc trn tr thnh:
P = (n!/(s! t! u!)) (p)s (q)t (r)u
V d: Nu 2 ngi d hp t v tnh trng bch tng c 5 con th xc sut
h c 2 con trai bnh thng, 2 con gi bnh thng v 1 con trai bch
tng l bao nhiu?
Trc ht ta tnh tn s ca mi bnh:
Con trai bnh thng = (3/ 4) (1/ 2) = 3/ 8 = p.
Con gi bnh thng = (3/ 4) (1/ 2) = 3/ 8 = q.
Con trai bch tng

= (1/4) (1/2) = 1/8 = r.

Con gi bch tng

= (1/4) (1/2) = 1/8 = m.

Vy: P = (5!/ (2! 2! 1! 0!)) (3/8)2 (3/8)2 (1/8)1 (1/8)0 = 0,074

K. nhng th t bo lai chut ngi, mt kiu hnh c th s
ch xut hin trong tt c cc dng c mt nhim sc th ngi
nht nh.
V d: Mt enzim ca ngi no ch c mt dng B. Cc nhim sc
th ngi c mt cc dng A, B &C nh di y. Hy xc nh gen quy
nh enzim ny nm trn nhim sc th no?
Nhim sc th ngi
Dng

55

Nu gen quy nh tnh trng nm trn nhim sc th no th nhim

sc th phi c mt cc dng (+) (dng tnh). Cc nhim sc th
1, 2, 5, 6 c mt dng B. Nu gen ang xt nm trn nhim sc th 1
th enzim ny phi c mt c 3 dng. Nhim sc th duy nht ch c
dng B l nhim sc th s 6. V vy gen ny nm trn nhim sc th
s 6.
Tm tt
1. phn tch ph h:
a. Ta gi s mt quy lut di truyn c th v suy ra kiu gen ca c
th c biu hin tnh trng .
b. So snh i con theo d on v tn s ca chng vi cc kt qu
mong i theo l thuyt.
c. Loi tr m hnh di truyn khng ng.
2. cc th lai t bo ngi, ta tm mt nhim sc th m l duy nht,
ch c mt dng c biu hin enzim .

Bi tp
1. PKU l mt d tt bm sinh v s chuyn ho axit amin phenylalanine.
Ph h sau y l ca mt gia nh b nh hng bi bnh ny:

I
II
III
1

56

a. Gii thch quy lut di truyn chi phi PKU?

b. Nhng ngi no trong ph h trn l d hp t v PKU ?
c. Tnh xc sut III-2 l d hp t.
d. Nu III-3 & III-4 kt hn, tnh tn s a con u tin s b PKU?
2. Ph h di y m t s di truyn tnh trng lingus lanulatis. Xc
nh cc quy lut di truyn khng chi phi tnh trng ny.

I
II
3. Xc nh cc quy lut di truyn c th chi phi mi ph h sau?

4. Bn chn lc c 3 dng t bo lai chut-ngi v phn tch xem chng

c nhng nhim sc th no ca ngi. Sau bn phn tch mi
dng v s c mt hoc vng mt nhng enzim c trng ca ngi
( +: c mt nhim sc th ngi hoc hot tnh enzim). Da trn kt
qu di y, hy ch ra nhim sc th c gen quy nh mi enzim.
Nhim sc th ngi
Dng

10

12

13

22
+

57

Enzim
Dng

E
X

5. V ph h m t r nht cc kiu di truyn sau:

a. Gen ln nm trn nhim sc th thng.
b. Gen tri lin kt X.
6. Xt ph h sau:

Nhm mu A
Nhm mu B
Nhm mu AB
Nhm mu O

Tnh tn s a tr ca cp v chng 2 a chu u tin c

nhm mu O?

7. Vi mi ph h sau, xc nh xem quy lut di truyn no l khng th

p dng c i vi tnh trng ang xt:

58

8. B ca mt ngi n ng cht v bnh Huntington, mt bnh do gen

tri nm trn nhim sc th thng quy nh. Triu chng ca bnh
ch biu hin khi tui trung nin. Trong gia nh ca ngi n ng
ny khng c ai b bnh.
a. Tnh xc sut ngi n ng mc bnh.
b. Tnh tn s a con ca cp v chng ny b mc bnh.
9. ngi, s thiu rng hm l do gen tri nm trn nhim sc th thng
quy nh. Nu hai b m d hp t sinh 5 a tr, hy tnh tn s :
a. Tt c 5 a tr u khng mc bnh.
b. S c 3 a bnh thng v 2 a b bnh.
c. Hai a tr u tin s bnh thng v 3 a tr cui cng b bnh.
10. X nang l mt bnh di truyn bi gen ln. Nu hai ngi bnh thng
nhng d hp t ly nhau, tnh tn s :
a. Mt trong ba a tr s b bnh.
b. Trong bn a tr, th t s l: con trai bnh thng, con gi b bnh,
con trai b bnh, con gi bnh thng.
11. Mt ngi n ng bnh thng (1) nhng c ng ni b bch tng ly
mt ngi ph n bnh thng (2) nhng c m b bch tng. Tnh tn s
a tr th hai bnh thng.
12. ngi, kh nng chuyn ho phenylthiocarbamide (PTC) c xc
nh bi gen tri. Gi s mt ngi c kiu gen d hp t kt hn vi mt
ngi khng c kh nng chuyn ho.
a. Tnh tn s trong 4 a con ca h c hai a c kh nng
chuyn ho PTC?
b. Tnh tn s trong 6 a con ca h c 3 a con c kh nng
chuyn ho PTC.

59

13. ngi, thiu rng hm l do gen tri nm trn nhim sc th thng,

bnh bch tng v bnh Tay-sach l do gen ln nm trn nhim sc
th thng. Nu mt ngi n ng c rng hm v d hp t v bnh
bch tng v bnh Tay-sach ly mt ngi ph n d hp t v c 3 gen
trn, tnh tn s a con u tin ca h :
a. C rng hm, b bnh Tay-sach v bnh bch tng.
b. Thiu rng hm hoc c bnh Tay- sach
14. Ba dng t bo lai gia chut v ngi c hoc thiu mt s nhim
sc th ngi theo bng sau:
Dng t bo

nhim sc th ngi
1

11

13

17

a. Nu mt gen m ho mt enzim no nm trn nhim sc th s

11, nhng dng no l dng tnh v enzim ?
b. Nhng dng no c th dng tnh nu gen nm trn nhim sc th
s 5?
15. Bnh sc t tng phn trn da ngi l mt hin tng him c, trong
melanine khng c chuyn ho bi t bo sc t, gy ra nhng
dng t bo sc t dng xoy trn da. Mt ngi ph n b bnh ly mt
ngi n ng bnh thng . C ta c 3 a con gi bnh thng, 2 a
b bnh v 2 con trai bnh thng. Ngoi ra c ta c 3 ln xy thai m
thai u l nam gii b d tt. Hy gii thch nhng kt qu trn.

Chng 7
60

t bin nhim sc th
A. S thiu vng hai hoc nhiu kiu hnh mong i cho thy
th d hp t chuyn on lin quan n cc gen lin kt.
Hu ht cc giao t to thnh do trao i cho nm trong mt o on
khng sng st c v lc nhim sc th khng hon ton tng ng.
Nhim sc th bnh
thng
Chuyn on cn tm

trao i cho gia C v

D

Nhim sc th
bnh thng
o on
Lp on
v thiu
on
Lp on
v thiu
on

Chuyn on xa tm

trao i cho gia C v

D

Nhim sc th
bnh thng
o on
on khng
tm
Nhim sc
th hai tm

V khng phi mi kiu giao t u c to ra, nn mt s lp kiu

hnh nht nh s b thiu.
V d: Mt cy d hp t ABCDE/ abcde th phn vi mt cy ng hp t
ln abcde/ abcde. Th h sau thu c cc dng sau:
ABCDE
abcde
61

Abcde
aBCDE
ABCDe
abcdE
Kt qu trn th hin s bt thng no? hy gii thch?
Ta thy c 5 cp gen d hp t, v vy ta c th nhn c 25=32 t
hp kiu hnh khc nhau (xem chng 2), nhng ta ch thu c 6 kiu
hnh. Chng ta khng thy c trao i cho 3 gen B, C, D . Ba gen ny
c th lin kt nhau qu cht ch, v vy khng c s ti t hp gia
chng. Tuy nhin iu l khng chc chn. Cc gen B, C v D c th
bn trong mt o on, tc l d hp t trong b m d hp t. S ti
t hp xy ra bn trong o on ny to nn nhng hp t khng th tn
ti c.
B. Nu mt kiu hnh ln xut hin khng nh mong i, c th
l do mt on.
Mt t bin mt on mt vng trn mt trong hai nhim sc th
tng ng dn n ch c mt bn sao ca mt gen no , tng t
nh trng hp gen lin kt vi X khng c on tng ng trn Y.

a
V d: Mt rui ci en thm ng hp t lai vi rui c di ng hp t
c chiu x bi tia X. Trong s rui con xut hin mt con rui en
thm. Hy gii thch kt qu.
S lai (e: en thm, e+: kiu di)
ee x e+e+

e+ e
(tt c l kiu di)
Nu nhim sc th con c mang t bin mt on vng gen m
ho tnh trng en thm, F1 c kiu gen:
e
//////////
V khng c alen kiu di t alen en thm nn kiu hnh ca rui F 1
ny l en thm. Chng ta c th khng nh t bin mt an bng
vic lai rui F1 en thm vi rui F1 kiu di:
62

//////

1 e/e : 1 e/ : 1 +/e : 1 +/
1 en thm : 1 kiu di
Kt qu ny ng nh kt qu php lai phn tch.
C. Cc mt on lun gy cht khi trng thi ng hp t.
V d: Xt li v d phn 7-B. Chng ta gi s lai rui c v ci F 1 cng
c kiu hnh en thm trn. Kt qu s nh th no ?
Ta c s lai:
e

/////

/////

: 2

:1

/////
3 (en)

/////
/////

1 (cht)

V 1/ 4 s con b mt on e c 2 nhim sc th nn chng khng

th sng c. S con cn sng s l 3/ 4 v tt c chng u c mu en
thm.
D. Mt on c th c s dng nh v cc gen nhng vng
c th.
Nu mt gen cn xc nh nm trong mt mt on no th mt
lng nht nh cc enzim s b mt i (khong 50%) so vi bnh thng.
V d: Ta cn xc nh v tr ca 1 gen Drosophila, v bit rng gen
nm nhim sc th s 2. Chng ta c 5 dng rui vi nhng mt
on nhng vng khc nhau trn nhim sc th s 2 nh sau:
Nhim sc th bnh thng:

___10___20___30___40___50___60

n v bn

A
B

//////////
//////////////////////////
63

////////////////////////

////////////////////

////////////////

Ta lai mi dng vi rui gim kiu di v xc nh hm lng enzim

mi c th F1 t mi php lai. Kt qu thu c c trnh by di y.
Hy xc nh vng m gen nh v.
Dng em lai

T l enzim (%)

100

48

49

97

101

Chng ta thy dng B v C hot tnh enzim ch t 1/ 2. V vy gen

m ho enzim ny phi nm vng lin quan n vng b mt on c 2
dng ny, tc l vng 25-35 n v bn k t tri.
E. S gim s lng con lai c th gi n s chuyn on tng
h.
Vi nhng chuyn on tng h, ch mt vi kiu phn ly nhim
sc th c kh nng to ra cc giao t hoc hp t c cp nhim sc
th tng ng.

(a)phn ly xen k

nhim sc th
bnh thng
nhim sc th
chuyn on

(b) phn ly lin

k(1)

thiu ht lp
on
thiu ht lp
on

(c) phn ly lin

k(2)

thiu ht lp
on

64

thiu ht lp
on

V d: Xt cc php lai sau ng, sau gii thch kt qu:

P

Dng A x dng A

B xB

AxB

F1

513

510

515

F2

531

514

341

Chng ta thy rng s c th F 2 t php lai A x B thu c t hn cc

php lai khc. iu c th lin quan n mt s bt thng no .
Mt cch gii thch l, mt trong nhng dng l ng hp t v mt
chuyn on tng h. Chuyn on, khi trng thi d hp t, s to ra
nhng giao t hoc con kh sng st.
F. Mt t gy nhim sc th c th c x l nh mt t bin
v v vy c th lp bn .
V d: Di y l bn 3 gen ln lin kt X Drosophila:
|

33, 0

36, 1

43, 0

v: mt son, m: cnh ngn, s: thn en

Rui c kiu di c chiu tia X v lai vi rui ci mt son, cnh
ngn, thn en. Trong s rui con, mt rui ci (mt , cnh di) c
khi phc. Khi cho rui ci ny giao phi vi rui c v m s, kt qu thu c:
Rui ci

Rui c

73 v m s

70 v m s

80 ++ s

4++s

Hy gii thch kt qu thu c bng cch lp bn di truyn.

65

Vic s dng tia X c th to nn nhng bin i nhim sc th.

Thc t, con ci thn en xut hin khi chng ta ngh rng tt c F1
u l kiu di cho thy chng ta c mt t bin mt on (xem KN 7B). Mt on phi kt thc gia m v s. Nu mt on cha c gen m
(cnh ngn), chng ta phi c rui ci thn en, cnh ngn.
Chng ta c th v s lai vi cc nhim sc th ca php lai tr li:
v

////

v
x

Cu hi y l on mt cch gen m l bao nhiu ? Chng ta xt

nhng con c t php lai phn tch trn. Ch l chng ta ch c 1/ 2
s rui c so vi rui ci. Nhng con c nhn c nhim sc th X b
mt on b cht. Nhng rui c mu en phi l kt qu t s ti
t hp gia mt phn cui on mt v gen m nh minh ho di y:
v

s
/////

Trao i cho sinh ra cc nhim sc th sau:

+

/////

V vy tn s trao i cho RF = s rui c en/tng s rui c x

100%
= 4/74 x 100% = 5, 4%
Nhim sc th ca con ci F1 c v li nh sau:
v

5,4
+

/////
+

G. Nhng kiu hnh bt thng c th l kt qu ca hin tng d

bi.
Cc gen ca nhng c th bnh thng l rt cn bng. S nhim
sc th nhiu hn (nh 2n +1) hoc t hn (nh 2n -1) bnh thng lun
lm thay i kiu hnh. Cc th mt (2n -1) s ging nh t bin mt
on, cc alen ln s c biu hin.
V d: Qu ca c c dc bnh thng c dng to, trn vi t l di/ rng
=2. Lai gia nhng cy bnh thng vi nhau thu c qu thng dt hn
bnh thng (t l chiu di /rng nm gia 3 v 4). Gii thch s xut
hin kiu hnh qu dt.

66

Hnh dng qu hu ht b chi phi bi nhiu gen. Kiu hnh dt gi

n kiu d bi no . Phn tch nhim sc th s cho thy c nhng
nhim sc th tha.
H. Nu hai loi khc nhau cho ra con lai hu th, hy xt n trng hp a bi ha.
V d: 2 loi bng c i A v B, mi loi c 13 cp nhim sc th.
Php lai gia A v B i khi thu c nhng con lai hu th. Gii thch kt
qu.
Thng thng nhng con lai khc loi l bt th v cc nhim sc th
ca nhng loi khc nhau khng th bt cp trong qu trnh giao phi.
Nu mi b nhim sc th n bi bt ngun t mi ngun gc c gp
bi ln, b nhim sc th a bi s kt cp c v gim phn xy ra bnh
thng.
Gi A l b n bi t loi A; B l b n bi t loi B. Con lai bt th
in hnh c mt b A v mt b B. Nu mi b n bi ny c gp
i, con lai s c b nhim sc th l AABB. Cc nhim sc th trong b
A v B s kt cp c v gim phn c th din ra bnh thng. Con lai
hu th c th c 13 x 2A +13 x 2B = 52 nhim sc th hoc 26 cp
nhim sc th.
Tm tt
1. Cc o on lm gim s lp kiu hnh.
2. Mt on cho php cc t bin ln c th xut hin F1.
3. Nhng t gy nhim sc th c th c x l nh mt gen khc, do
vy c th lp bn di truyn.
4. Cc d bi th lun lun lm thay i kiu hnh.
5. Nhng chuyn on tng h thng lin quan vi tnh bn bt th.

Bi tp
1. Cy Q c 14 nhim sc th v cy R c 20 nhim sc th. Cc
nhim sc th con lai QR hu th s phi l bao nhiu ?
2. Rng mu nu ngi do gen ln lin kt X quy nh. Mt ngi ph n
rng trng c b m rng nu kt hn vi ngi n ng rng nu.
Cp v chng ny c con trai rng nu, chm pht trin v tr tu v
tinh hon km pht trin. Hy gii thch v chng minh rng c s
khng phn ly nhim sc th b m.
67

3. C 5 gen ln a, b, c, d v e c v tr nh sau trn nhim sc th s 2

rui gim:
a

b
10

15

25

30

40

Mt rui ci ng hp t v c 5 gen c lai vi rui c kiu di

ng hp t, thu c tt c F1 u kiu di. Khi mt rui ci F1 c lai
vi rui c ng hp t ln, kt qu i con nh sau:
abcde

80

+++++

94

a++++

+bcde

ab+++

++cde

abcd+

++++e

Tng s

200

a. Kt qu thu c c g bt thng?
b. Gii thch hin tng .
c. V bn nhim sc th ca rui ci F1.
4. Cc bng (16) nhim sc th tuyn nc bt ca rui gim c th
hin di y cng 5 mt on, k hiu t A n E.
Bn c 6 alen ln: p, q, r, s, t v u nm trn vng ny nhng th t
ca cc gen cha bit. Bn to ra cc dng rui trong mi t bin
mt on l d hp t vi mi alen. Trong bng sau, ch ci biu din
mt kiu hnh t bin v du (+) biu din kiu hnh di. Hy xc
nh bng no tng ng vi gen no, tc l xc nh trt t ca gen.
1

Mt on:
A
B
C
D
E

///////////////////////// ////////
////////////////////////////////////
//////////////////////////////////////
//////////////////////
///////////////////////////////////
68

Mt on

Kiu hnh tng ng vi

p

5. Mt cy ng c kiu gen Sh/Sh c lai vi cy c kiu gen sh/sh. Cc

cy F1 bn bt th v c kiu hnh Sh. Cc cy ng F1 c lai vi cy
sh/sh, to ra:
820 cy bn bt th, Sh
800 cy hu th, sh
195 cy bn bt th, sh
185 cy bnh thng, Sh
a. Hin tng di truyn no lin quan n kt qu trn?
b. V bn nhim sc th ph hp ca cy F1.
6. Mt th t bi vi n = 3 (A, B, C) c lai vi th lng bi (n = 3) c cng
cc nhim sc th ging nh th t bi (A, B, C).
a. C bao nhiu nhim sc th v bao nhiu b nhim sc th c
con lai.
b. Bn cho rng con lai l bt th hay hu th ?
7. Thn vng rui gim l do gen ln lin kt X quy nh. Mt rui
c thn xm c chiu x v lai vi rui ci thn vng. th h con
c mt rui ci thn vng.
a. Ti sao li thu c kt qu trn, hy gii thch.
b. Xc nh t l i con nu lai rui ci thn vng F 1 vi rui c
thn xm bnh thng khng b chiu x ?
8. Loi thc vt A c 2n = 22 v loi B c 2n = 12. Ngi ta tm c mt
con lai hu th gia hai loi. Con lai ny c bao nhiu nhim sc th ?
9. Mt ngi ph n c kh nng nhn bnh thng c cha m b m mu
ly mt ngi n ng b m mu. H c mt con gi b m mu v mc
hi chng Turner (XO). b hoc m c gi xy ra s khng phn
ly nhim sc th?
10. Gii thch c ch to thnh c th nam gii XYY.

69

11. Drosophila, 2 gen ln a v b nm v tr 16, 5 v 35, 5 trn nhim

sc th s 3. Rui ci kiu di vi kiu gen ++/ab c lai vi rui c
ab/ab v cho th h con:
++

500

ab

450

+b

26

a+

24

Tng s

1000

a. Kt qu trn c iu g bt thng? Gii thch.

b. V bn nhim sc th s 3 rui ci ++/ab.
12. C 4 dng Drosophila (a, b, c, d) c phn lp nhng vng a l
khc nhau. So snh cc mu bng nhim sc th s 3 v nhn c
kt qu nh sau (mi s tng ng vi mt bng nht nh).
a

1 2 6 5 4 3 7 8 9 10

1 2 3 4 5 6 7 8 9 10

1 2 6 5 8 7 9 4 3 10

1 2 6 5 8 7 3 4 9 10

Nu c l dng gc, cc dng khc c to ra nh th no ?

13. rui gim, gen ln quy nh mu vng nm gn u ca nhim
sc th X. Mt rui gim c thn xm b chiu x v lai vi rui gim
ci thn vng. th h con c mt con c thn xm. Hy gii thch
v s to thnh rui c xm v lm th no kim tra gi thuyt
?
14. Hi chng Down ngi l kt qu ca s tha nhim sc th s 21.
Lun c 3 nhim sc th s 21 ngi b Down. Bnh ny cng c th l
kt qu t s chuyn on mt phn nhim sc th 21 n nhim sc
th 15. Gi s mt ngi n ng bnh thng l d hp t v chuyn
on trn.
a. Ngi n ng to ra nhng loi giao t no?
b. Tnh tn s ngi n ng c th sinh ra mt a con mc
hi chng Down.
15. rui gim, c 3 gen lin kt X: notch, dwarf v facet to ra cc kiu
hnh cnh x, thn ngn, mt dng que. Xt php lai sau:
Php lai 1:

ci cnh x x c thn ngn

300 ci cnh x
300 c kiu di
70

300 ci kiu di
Php lai 2:

F1 ci cnh x x c thn ngn

300 ci cnh x

300 c thn ngn

300 ci thn ngn

34 c kiu di

33 ci kiu di
33 ci cnh x, thn ngn.
Php lai 3: ci thn ngn, cnh x t php lai 2 x c mt que

500 ci cnh x, mt que

500 ci kiu di
500 c thn ngn
Php lai 4:

ci kiu di t php lai 3 x c kiu di

500 ci kiu di

220 c thn ngn

220 c mt que

30 c thn ngn, mt que

30 c kiu di
a. C t bin no xy ra khng hay kt qu trn l do cc mt on?
b. Lp bn nhim sc th thch hp t nhng s liu trn.

Chng 8
Phn tch b bn
A. Trong php phn tch bo t ngu nhin, bn khong cch
gia cc gen lin kt c tnh bng s cc th ti t hp chia cho
tng s cc bo t nhn vi 100.
V d: nm men, cc php lai thu c kt qu nh di y. Hy xc
nh trt t cc gen v khong cch gia chng.
a+b- X a-b+

a+c- X a-c+

b+c- X b-c+
71

a+ b -

490

a+c-

a-b+ 500
a+b+
a-b-

a-c+

425

417

a+c+

a-c- 73

1000

1000

b+cb-c+

425

425

85

b+c+

b-c-

70

80
1000

mi php lai, hai lp cui cng l nhng th ti t hp. V vy:

Khong cch

a- b = 10/1000 x 100 = 1 n v bn
a - c = 158/1000 x 100 = 15, 8 n v bn
b - c = 150/1000 x 100 = 15 n v bn

Do trt t cc gen l:
a 1 b

15

B. Kiu bn c to ra do s trao i cho gia mt gen v tm

ng
i vi hai gen khng lin kt, s phn b ngu nhin trong qu trnh
gim phn to ra cc kiu hnh khc b m v nhng kiu hnh ging
b m.
Nu ta c php lai
nhim sc th, ta c

ab

++, do cch phn ly khc nhau ca cc

b
a

b
sinh ra 2 ab : 2 ++, l cc

kiu hai ging b m

+

72

+
a

+
sinh ra 2 a+ : 2 b+, l

cc kiu hai khng ging b m

+

nhn c 4 loi bo t khc nhau phi c s trao i cho gia

mt gen v tm ng ca n.
a

b
a

b
sinh ra kiu bn: 1

aa: 1 +b: 1 a+: 1 ++

+

V d: Trong php lai: leu+ his- x leu- his+ di y, hy phn bit cc dng
bo t ging v khc b m (hoc b bn) v cho thy dng bo t no
lin quan n s ti t hp gia gen v tm ng.
I

II

Leu+ his-

III

leu+ his+

leu+ his-

Leu+ his- leu+ his+

leu+ his+

Leu- his+ leu- his-

leu- his-

Leu- his+

leu- his+

leu- his-

Cc lp I v II cha hai dng bo t v cc bo t lp I l ging b

m. V vy lp I thuc kiu hai ging b m (parental ditype -PD) v lp II
l kiu hai khc b m (non-parental ditype-NPD). Lp III cha 4 kiu bo
t, do vy n l kiu bn (tetratype - TT), l kt qu t s ti t hp gia
mt gen v tm ng.
C. Nu tn s PD > > NPD th 2 gen lin kt.
Tham kho hnh v sau:
Vi hai gen khng lin kt, do cch phn ly ca cc nhim sc th:
a

73

+
hoc
+

b
b
to ra

to ra

2 ab

2 a+

2 ++

2 +b

PD

NPD

Vi hai gen lin kt, s phn ly s sinh ra:

Khng c trao i cho
cho
a

C trao i

b
a

+
+
2 ab

2 a+

2 ++
PD

2 +b
>>

NPD

Nu 2 gen khng lin kt, th PD = NPD do s phn b ngu nhin

k gia.
Nu 2 gen lin kt, NPD ch c th to ra t trao i cho kp 4 si
v rt t khi xy ra.

74

V d: Xt cc php lai sau Clamydomonas. i vi mi php lai, xc

nh xem 2 gen c lin kt khng ?
PD

NPD

TT

Tng s

ac+ X + arg

51

49

ac+ X +pab

88

12

62

32

100
100
ac+ X + pf
100
pab+ X +pf

64

32

100
pf + X + thi

29

68

100
So snh s lng PD v NPN mi php lai. Nu PD > > NPD th 2 gen
lin kt nhau. Do gen ac, pab, pf v thi lin kt nhau, arg khng lin
kt vi cc gen ny.
D. Tn s ti t hp gia hai gen lin kt c tnh bng [(1/2TT)+
NPD] / Tng s x 100.
V d: Xt li php lai KN C v tnh khong cch gia cc gen.
Khong cch

ac pab = [1/2(12) +0]/100 = 6 n v bn .

ac pf = [1/2(32) +6]/100 = 22 n v bn .
pab pf = [1/2. 32 +4]/100 = 20 n v bn .
pf thi = (1/2. 68 +3]/100 = 37 n v bn

E. Trong cc b 4 xp hng, s phn ly ca ln phn bo th 2 c pht hin bi s phn tch ca cc lp bo t.

Nu php lai u tin l a X +, cc th phn tch ca ln phn chia
u tin l:
a

hoc +

Nu hai alen ging nhau khng lin k nhau th s phn ly ca ln

phn chia th 2 to ra:
75

V d: Trong php lai : a + X + b Neurospora thy xut hin cc kiu

bo t sau. Hy phn bit kiu phn ly xy ra i vi mi gen:
I

II

III

IV

VI

a+

ab

a+

++

a+

ab

+b

a+

a+

ab

+b

++

a+

++

+b

ab

++

ab

+b

+b

+b

++

ab

++

Hy tm nhng ct cc cp alen thay th nhau khng lin k

nhau. Cc lp I, IV, V, VI th hin s phn ly ln phn chia th 2 i vi
a. Cc lp I, II, IV, V, VI th hin s phn ly ln phn chia th 2 vi b.
F. Trong cc b bn xp hng, khong cch t gen n tm ng
c tnh bng (1/2 t l phn tch ln phn chia th 2)/tng
s x 100.
c c s phn ly ln phn chia th 2, phi s trao i cho
gia gen v tm ng.
Khng c trao i cho
a

a
sinh ra

C trao i cho

76

sinh ra
+

V d: Neurospora, 3 gen: b, c, d c lai theo cc t hp tng cp. Da

trn kt qu di y, hy tnh khong cch gia tm ng v gen:
b+

+c

b+

c+

b+

++

++

+c

bc

bc

+c

bc

b+

70

26

++

+c

+d

cd c+
+d
++

c+

cd

cd

c+

cd

c+

cd

c+

+d

++

+d

++

+d

++

++

+d ++

c+

+d

++

+d

c+

cd

cd

15

42

30

Hy tm s phn ly ln phn chia th 2. Chng ta khng thy c s

phn ly i vi b, do khong cch t b ti tm ng bng 0, hay b lin
kt v tm ng. c 2 php lai, chng ta thy c 26 bo t c phn
ly ln phn chia th 2 i vi c. V vy khong cch gia c v tm
ng bng 1/2(26)/100 x 100 = 13 n v bn .
i vi d, chng ta thy c 41 bo t nh vy. Do khong cch t d ti
tm ng bng 1/2(41)/100 x 100 = 20,5 n v bn .
Tm tt

77

1. Cc khong cch bn trong php phn tch bo t ngu nhin c

tnh khi bo t th lng bi.
2. Pht hin s lin kt khi thy PD >> NPD.
3. Khong cch gia cc gen c tnh bng (1/2TT + NPD)/ Tng s bo
t x 100.
4. Khong cch gia gen v tm ng c tnh bng (1/ 2 s t hp
phn ly ln phn chia th 2)/Tng s bo t x 100.

Bi tp
1. Cc bo t ngu nhin sau y nhn c t php lai a++ X +bc. Hy
lp bn 3 gen .
abc 52

a++ 102

ab+ 34

+b+ 8

a+c 6

++c 28

+bc 112

+++ 58

Tng s: 400
2. Bn phn lp c mt loi nm mi v nhn c dng khuyt dng nicv dng khuyt dng thi-. Lai hai chng ny v nhn c 500 bo t.
Nui bo t mi trng ti thiu. C 35 bo t mc c. Tnh khong
cch gia 2 gen.
3. Xt cc php lai v cc b 4 xp hng sau y. V bn ca 3 gen
ny. Tnh khong cch gia cc gen v khong cch t gen ti tm ng.
I II

III

IV

VI

VII

a+

ab

a+

a+

a+

a+

a+ab

+b

ab

++

+b

++

+b ++

ab

++

ab

a+

++
ab

a+ X +b:

+b

++

++

+b

+b

+b

32

30

10

18

II

III

IV

VI

ab

Tng s =100

VII
78

a+ X +c: a+

ac

a+

a+

a+

a+

ac

a+

ac

+c

ac

++

+c

++

+c

++

ac

++

ac

a+

++

+c

++

++

+c

+c

+c

ac

69

12

Tng s =100.

4. Nu gen q nm cch tm ng 12 n v bn , th trong php lai

q X +, t l % bo t th hin s phn ly ln phn chia th 2 l bao
nhiu ?
5. Xt php lai nm to ra cc bo t nang sau
PD

NPD

TT

a+ x +b

48

52

a+ x +c

90

a+ x +d

51

49

b+ x +d

60

10

30

a. Cc gen no lin kt nhau ?

b. Tnh khong cch gia cc gen lin kt.
6. Nu cc gen a v b lin kt, cch nhau 2 n v bn th c bao
nhiu kiu b 4 trong 100 bo t c to ra t php lai a+ x +b ?
7. Neurospora, php lai ad trp x ++ to ra cc b 4 xp hng sau:
I

II

ad trp
+trp

III
ad+

ad trp
++
++
126

IV

ad trp

ad+

ad+

VI

ad trp
+trp

ad trp

++

+ trp

+ trp

ad +

ad trp

+ trp

++

++

++

+ trp

18

30

VII
ad+

+trp

++

ad +

ad trp

ad +
12

a. Xc nh khong cch gia cc gen v tm ng.

b. Nu cc gen lin kt, tnh khong cch gia cc gen v lp bn .

79

8. C 3 tnh trng khc nhau: x, y, z trong mt php lai Neurospora. T

php lai x++ X +yz, cc b bn xp hng sau c to ra. Da trn s
liu , hy xy dng bn ca 3 gen v ch ra v tr tm ng:
I

II

x++

III

IV

x++

x++

VI
x++

VII
x++

VIII
x++

x+z

x+z
x++

+yz

xyz

+yz x++

+++

+y+

+yz +yz

+yz

x+z

+yz

+yz

+yz

+y+
+yz

+++

xyz

x+z
+y+

+y+ x++
x++

+yz

9. nm men, khong cch t gen a n b l 12 v t a n c l 28.

Tnh PD, NPD, TT nu trong mt php lai ta c:
a. a+ x +b, TT/NPD = 4.
b. a+ x +c, TT/NPD = 6.
10. Mt php lai gia hai gen sinh ra s PD bng TT, sTT = 12 ln NPD.
Tnh khong cch gia cc gen.

80

Chng 9
Di truyn hc vi khun
A. Tn s ng bin np v ng ti np cao cho thy 2 gen rt
gn nhau.
V d: Trong mt thc nghim bin np, x+y+z+ c s dng l th cho v
x-y-z- l th nhn, thu c 250 th bin np. Sau nui cy bn sao
xc nh xem c y+ v/ hoc z+ khng. Kiu gen ca cc th bin np
thu c nh di y. Bn c th kt lun g v v tr tng ng ca cc gen?
x+y-z-

53

x+y-z+

173

x+y+z-

x+y+z+

17

Chng ta thy rng 76% khun lc c x+z+. Kt qu ny cho thy x v

z gn nhau. x+ v y+ xut hin cng nhau vi t l 10% cho thy y cch xa
x, v c th khng lin kt vi nhau.
81

 tnh tn s ti t hp gia x v z, chng ta xem s ln x+ v z+

khng cng xut hin ( l x+ y- x- v x+ y+ z -).
RF = (53 + 7)/240 = 25 n v bn .
B. cc gen khng lin kt ng bin np v ng ti np cn c
2 trao i cho.
Xt hai gen a v b xa nhau, tt nht l hai gen trn hai phn t
ADN ring bit:
a+

b+

cc th bin np l a+b+, c hai an ADN ny cng phi xm nhp

vo th nhn. Qu trnh ny cn hai t gy, tng ng vi mt trao
i cho kp. V vy iu l rt him.
a+

b+

th cho

a-

b-

th nhn

Lu rng nu a v b gn nhau v trn cng mt mch ADN, th ch

mt t gy v trao i cho l .
a+
a- b-

b+

th cho
th nhn

V d: Xt cc tn s ng bin np sau:
A v B:

73%

A v C:

7%

B v C:

6%.

Nhng gen no khng lin kt vi nhau?

Chng ta thy gen C xut hin cng vi A hoc B vi t l thp. Nh vy
c th gen C khng lin kt vi A v B.
C. xc nh trnh t cc gen trong qu trnh bin np hoc ti
np, hy tm cc trt t gen cho php phn on v cc trao i cho
kp.
V d: Khi a+b+c- l th cho v a-b-c+ l th nhn, ta thy lp kiu hnh c
tn s thp nht c kiu gen a+b+c+. Hy xc nh trt t ca cc gen.
Gi s trt t gen l abc, khi chng ta c th v s php lai nh
sau:
82

___ a+____ b+ ___ c- ____

___ a- ____ b- ___ c+ ____
Mt trao i cho kp s to ra a+ b- c-, lp ny khng phi c tn s t
nht. V vy b khng th gia.
Gi s c gia:
____a+ _____ c_ _____b+ _____
____a_ _____ c+ _____ b_ _____
Trao i cho kp s to ra a+b+ c+, lp ny c t l nh nht, chng
t rng c gia.
D. Trong qu trnh tip hp, mt gen c tn s truyn gen cao
phi nm gn vi im khi u qu trnh truyn gen.
V d: Php lai gia cc t bo his+, leu+, thr+, pro+, mn cm vi str v
cc t bo his-, leu-, thr-, pro-, khng str, ko di 25 pht. Khi ngng tip
hp, cc kiu gen c ti t hp c xc nh. Kt qu thu c bng di
y. Hy xc nh cc gen u tin v trt t c th ca cc gen.
Kiu gen

S khun lc

his+

leu+

12

thr+

27

pro+

Chng ta thy rng cc t bo thr+ nhiu nht. Tn s t gy ng

tip hp tng vi s tng thi gian tip hp. V vy cc gen cng xa
im khi u truyn gen s xut hin vi tn s cng nh. Chng ta c
th xp xp cc gen mt cch tng i da trn tn s ca cc kiu gen
pht hin c. Trt t l thr leu pro his. V khng thy c his+ v v
chng ta dng qu trnh tip hp 25 pht, nn gen his phi nm sau 25
pht trn bn thi gian.
E. Cc gen ca virut thng c xc nh trt t bng cch tng t
nh vic lp bn di truyn eukaryote.
V d: E. coli c gy nhim mt cch ln ln vi hai loi phage T4: r+ m+
tu+ v r m tu. i con xut hin nh sau:
+++

1243

++ tu

322

r m+

284

r m tu

1158

+m+

175
83

+ m tu

54

r++

58

r + tu

160

Tng s

3454

Tnh khong cch gia cc gen v v bn di truyn.

Xt cp r-m, hy xc nh cc lp m trong r+m+ khng cng xut
hin.
RF r-m = (175 + 54 + 58 + 160)/3454 x 100 = 13 n v bn .
Tng t, RF r-tu = (322 + 284 + 54 + 58) x 100/3454 = 21 n v bn
.
RF m-tu = (322 + 284 + 175 + 160)/3454 x 100 = 27 n v bn
.
T kt qu trn ta c bn di truyn l:
___ m ____ 13 ___ r _________ 21 ___ tu __
kim tra li, hy xc nh cc trao i cho kp to thnh cc
th ti t hp r+m tu v r m+tu+. Ta thy rng ch c cc alen r b thay
i do s sp xp li t b m, v vy r phi nm gia.
F. Thiu s b tr nghi l hai t bin thuc mt gen.
Xt hai trng hp sau:

I:

II:
gen 1

gen 1

gen 2

Trong trng hp I, c hai phn t ADN cng to ra mt protein khng c

chc nng. T bo khng c protein bnh thng cho thy rng n c kiu
hnh t bin. Trong trng hp II, ADN pha trn c gen 2 bnh thng v
ADN bn di c gen 1 bnh thng. T bo ny s c c protein 1 v protein
2 bnh thng v s xut hin kiu hnh bnh thng. S sinh trng c
ca t bo, hoc chnh l s b tr cho nhau, cho thy rng cc t bin
ny thuc nhng n v chc nng khc nhau, hay thuc cc gen khc
nhau.
V d: Xt s liu v hin tng b tr sau. Bao nhiu gen c lin quan
n kiu hnh t bin v t bin no thuc v gen no ? (+: b tr,
v -: khuyt dng).

1
2

+
84

Hy tm cc t hp trong khng c s b tr. Cc lp ny s cha

cc t bin thuc cng mt gen. 1 v 4 khng b tr v v vy n thuc
mt gen (gen A). Bng lp lun tng t, 2 v 3 thuc mt gen khc (gen
B). t bin 5 b tr vi tt c cc t bin khc v v vy n phi thuc
mt gen khc vi gen cha 1, 4 hoc 2, 3 (gen C).
G. Nu mt t bin khng b tr cho cc t bin hai gen
khc nhau th t bin mi ny chc chn l mt mt on.
V d: Xt mt t bin mi s 6 cng vi cc t bin khi nim F. N
cho nhng kt qu b tr sau. Bn cht ca t bin s 6 l g?
6

Chng ta thy t bin s 6 khng b tr cho 1 v 4, cho php ta

ngh rng t bin s 6 thuc gen A. Nhng t bin 6 cng khng b tr
vi t bin 5, l t bin ta bit thuc mt gen khc. t bin 6
nh hng n c gen cha t bin 1 v 4 v c gen cha t bin 5.
iu cho thy t bin ny nm gen khc.
Mt cch gii thch cho kt qu ny l c mt mt on bao trm
nhng vng c cc t bin 1 v 4 v vng c t bin 5. Hoc t bin
6 c th l mt t bin kp vi mi t bin ca n nm trong mt gen.
H. Nu hai mt on c chung mt phn b mt (ngha l chng
trm ln nhau) th chng khng th cho ra cc th ti t hp kiu
di.
Xt cc kh nng sau:
a. Hai mt on khng trm ln nhau:

S ti t hp s to ra:
ADN mang hai mt on:
v ADN bnh thng:

________/////////___/////////
______________

b. Hai mt on trm ln nhau:

85

________///////_______
_________////////_____
Ti t hp s to ra cc ADN u c on b mt.
V d: Cc chng b mt on di y c lai vi nhau v chn ra cc th
ti t hp kiu di. Kt qu nh sau (+: c cc th ti t hp kiu di, khng c cc th ti t hp kiu di). Hy xc nh trt t ca cc mt
on.
1
1

Chng ta thy rng chng 3 khng cho ra th ti t hp kiu di no.

N phi mang mt on ko di t nht n phn on b mt chng
1 v 2. Cc mt on chng 1 v 2 cho ra cc th ti t hp kiu di, v
vy n khng trm ln nhau.
Vy bn c th l:
1. ____//////______________
2. ________________/////////
3.

////////////////////////////

I. Nu mt t bin im cho ra cc th ti t hp khi lai vi mt

mt on, th t bin im nm ngoi vng b mt on.
Xt php lai sau gia mt mt on v mt t bin im:
Trng hp 1: im t bin bn ngoi on mt
______/////////
______________ * __
S ti t hp s to ra mt AND mang t bin kp:
_______////////_____ * _
v mt AND bnh thng:
Trng hp 2: t bin nm bn trong on mt
_____///////////_________
_____ * ______________
S ti t hp s khng xy ra trong vng c mt on. Do vy tt c cc
th ti t hp vn l t bin.
86

V d: Cc th t bin im 1, 2, 3 c lai vi cc chng c mt on a

v b; ngi ta chn c mt s th ti t hp kiu di. Da trn kt qu di
y, hy xc nh v tr ca mi t bin (+ = th ti t hp kiu di, = th ti t hp t bin).
1

- +

3
-

+
+

t bin 1 khng sinh ra kiu di vi mt on b, v vy n phi nm

bn trong vng tng ng vi b.
Lp lun tng t, ta c t bin 2 nm trong vng tng ng vi mt
on a. V t bin 3 cho cc dng di vi c hai mt on, nn n phi
nm ngoi c hai t bin mt on.
J. Cc t bin tn cng ca operon hoc cc t bin vng
kim sot c th cho ra nhng kt qu m tnh gi trong php th
b tr.
Nhng t bin tn cng ca operon lm mt chc nng ca cc gen
xa trong mt operon.
V d: Xt mt operon gm 3 gen theo th t A, C, B. Mt th lng bi mt
phn c kiu gen a- ( tn cng ca operon) b+ c+/ a+b-c+ c thit k.
Kiu hnh ca chng ny l g?
t bin cc a- lm cho phn t ADN th nht c biu hin thnh ab-c-. Chng th 2 c kiu hnh b-. Th lng bi mt phn s c kiu gen
b-/b-, vy n s l th t bin (xem KN 9-F).
K. Cc mt on thng khng phc hi li c.
Nu l mt t bin nhm ngha, th baz b bin i c th thay
i thnh baz bnh thng. Trng hp ny s khng xy ra nu mt s
baz cng b mt.
V d: C 5 th t bin trp- k hiu t 1 n 5 c nui trn mi trng
ti thiu. Mt s khun lc xut hin chng 2 v 5. Bn cht ca mi
t bin ny l g ?
Chc chn mt s t khun lc biu hin s phc hi, do vy cc
chng t bin 2 v 5 phi l nhng t bin im. Cc chng t
bin 1, 3 v 4 c th l nhng mt on.
L. C th tch nhng th hi bin chu c ch nh ti t hp.
Nhng th hi bin gm hai loi: nhng th hi bin tht v nhng
th hi bin do cc t bin v tr th 2 c ch n hoc b tr cho t
bin ban u.

87

1. Th t bin
bin tht:

_____*_____ b t bin tip, s sinh ra th hi

2. Th t bin ____*______ b t bin tip sinh ra t bin

kp: ____*___*___ c kiu hnh bnh thng.
Khi mt trong hai chng ny c lai vi kiu di s cho cc kt qu khc
nhau.
1.

nhim sc th ca th hi bin tht

nhim sc th kiu di
Ti t hp, sinh ra cc con kiu di:

2.

nhim sc th ca th hi bin chu c

ch
nhim sc th kiu di
Ti t hp, to ra hai dng t bin:
*

C hai th ti t hp trng hp 2 vn l dng t bin. Lu rng

tnh hung ny cng s xy ra, nu t bin th 2 nm trong gen gc
hoc nm trong gen khc hon ton cch bit. t bin c cng s c
khi phc trong c hai trng hp .
V d: Ba th hi bin ca leu- l r1, r2, r3 c khi phc bnh thng. Mi
dng ny c lai vi kiu di v kim tra i con. Th hi bin 1 v 3 khng
cho ra th h con kiu di. Bn cht ca mi th hi bin ?
V cc th hi bin 1 v 3 khng to ra cc th ti t hp kiu di, nn
chng phi cha mt t bin c ch v tr th hai. Th hi bin 2 l
th hi bin tht.

Tm tt
1. Trong bin np ( v ti np), trt t gen c th xc nh c bng vic
xem xt cc trao i cho kp.
2. Cc s kin cn 2 trao i cho ng thi rt him xy ra.
3. Trong qu trnh tip hp, chng ta c th xc nh trnh t cc gen
bng vic tnh tn s xut hin ca cc gen nh du khc nhau.

88

4. S thiu hin tng b xung gi n t bin cc hoc cc t

bin bn trong cng mt gen.
5. Mt on khng phc hi c v khng cho ra cc th ti t hp kiu
di khi c lai vi cc t bin im.
6. C th xc nh trnh t on mt v t bin im da vo s
xut hin i con kiu di.

Bi tp
1. a. Trong mt thc nghim bin np, th cho l ara+nic+gal+ v th
nhn l ara-nic-gal-. Cc th bin np ara c chn v sau kim tra
tip. Kt qu cho thy 40% l ara+gal+ v 5% l ara+nic+. Xc nh trt
t c th ca cc gen.
b. Bn c th ch cn lm thm mt thc nghim bin np xc
nh trt t gen. Bn phi s dng cng th cho v th nhn, nhng c
th thay i qu trnh chn lc cc th bin np u tin. Bn phi
lm g v kt qu nh th no khi xc nh trt t gen m bn cn?

2. ADN t mt chng a+b+c+ c s dng chuyn sang chng a-b-c-.

S lng tng kiu gen xut hin di y. Hy xc nh v tr tng i
ca cc gen.
Kiu gen

S lng

a+b-c-

312

a-b+c-

320

a-b-c+

308

a+b-c+

16

a+b+c+

11

a+b+c-

140

a-b+c+

21
89

3. Hai chng Hfr 1 v 2 u l a+b+c+d+e+ c lai vi mt chng F l a-b-cd-e-. Cc php lai b dng tng giai on cch nhau 5 pht v cc
kiu gen ca cc th ti t hp c xc nh. Kt qu nh sau:
a

d
Tn s

Hfr 1 x F

b
e
c

10 20

30 40 50 60

70 80 90 10
0
Thi gian
(pht)
a
d

Tn s

Hfr 2 x F-

b
e
c

10 20

30 40 50 60

70 80 90 10
0
Thi gian
(pht)

Hy v mt bn cha tt c cc gen v c hai nhn t F. Th hin F

trong cc nh v hng truyn gen bng mi tn. Trn bn chun, gen
b nm cch im khi u truyn gen 20 pht.

0/9
0
20

90

4. 8 t bin trp- c lp nhau c phn lp. Tin hnh php th b tr

vi cc t bin theo kiu gp i. Da trn kt qu thu c di y,
bn c th xc nh c bao nhiu gen v t bin no thuc gen
no? (+ = b tr, - = khng b tr).
1
1

3
4
5
6
7
8

5. Cc chng Hfr sau y truyn cc gen theo trt t sau:

Chng

Trt t truyn gen (gen u tin bn phi)

ACTER

UIRET

MBACT

ABMUI

Hy xc nh trt t cc gen trn nhim sc th.

6. Bn phn lp c hai th t bin rII l rII-1 v rII-2 phage T4. RII-1
khng cho th hi bin kiu di v t bin rII-2 cho 4 th hi bin l
2rva, 2rvb, 2rvc, 2rvd. Mi th hi bin c lai vi kiu di cho kt qu
sau (+ = th ti t hp hi bin kiu di):
rev a

rev b

rev c

rev d

a. Bn cht ca t bin rII-1 v rII-2?

b. Bn cht ca mi th hi bin?
7. Trong mt th nghim ti np, th cho l d+e+f+, th nhn l d-e-f-; tin
hnh chn lc theo f+. Kt qu thu c nh sau, hy xc nh trt t gen:
91

Kiu gen

S th ti np

d+e+f+

25

d+e-f+

d-e+f+

80

d-e-f+

256

8. Operon trp E. coli c trt t sau: p E C D B A. Bn tch c 3 th t

bin trp- c lp nhau v to ra cc th lng bi tng phn cho mi t
bin vi p+ E+D+C-B+A+; khng th lng bi tng phn no c th tng
hp c tryptophan. Ba v tr khc nhau c th c ca cc t bin
ny l g?
9. C 9 chng his-, ba trong s c nhng mt on v 6 chng c
nhng t bin im. Cy 9 chng trn mi trng ti thiu v m
s khun lc mc (+ = mc c).
1

Phage c nhim vo mt trong 9 chng v sau c s dng ti

np mi chng mt on. Cc th ti np t mi php lai c cy trn
mi trng ti thiu v m cc khun lc mc (+ = th ti np mc c). Da trn kt qu sau, xc nh:
a. Chng no l chng t bin mt on.
b. V tr tng ng ca mi t bin mt on vi mt on khc ?
c. V tr tng ng ca 6 t bin im.
Chng
Mt on

10. Bn phn lp c mt s t bin im (k hiu t a n g) vng

rII ca phage T4. Cc php lai ghp i cho t l phn trm cc vit
tan kiu di nh sau:
Php lai

%kiu di

Php lai

% kiu

di
axb

0. 5

axc

0. 3

axd

0. 8

axf

0. 3
92

bxe

0. 4

bxd

1. 3

bxf

0. 2

cxd

0. 5

cxg

0. 8

exg

1. 0

bxg

1. 6

gxd

0. 3

Bn cng lai mi t bin im vi mi t bin mt on v tnh

cc th ti t hp kiu di. Kt qu thu c di y (+ = th ti t hp
kiu di, - = th ti t hp t bin).
t bin im
Mt on

a. V bn cc t bin im, tnh khong cch.

b. Xc nh v tr ca cc mt on tng ng vi cc t bin im.
c. Cc th t bin a v e b tr vi nhau v e c bit l thuc rIIB.
Cc t bin no rIIA v t bin no thuc rIIB?
d. Nhng t hp ca cc mt on no s to ra cc th ti t hp kiu
di?
11. C 3 th t bin lac- (k hiu t a n c). Bn thc hin cc th
nghim ti np v thu c s khun lc kiu di nh sau:
Th cho

Th nhn

lac a- pro+

lac b- pro-

S khun lc kiu di
93

lac b- pro+

lac a- pro-

27

lac a- pro+

lac c- pro-

15

lac c- pro+

lac a- pro-

64

lac b- pro+

lac c- pro-

10

lac c- pro+

lac b- pro-

98

Xc nh trt t cc t bin lac so vi gen pro.

93

12. Th t bin phage 1 gy nhim vo chng vi khun B v D, th

t bin 2 c gy nhim vo A v D. Ch phage kiu di mi sinh trng c chng C. C hai chng phage t bin c s dng ly
nhim ng thi vo D. Phage th h con c pha long 10-7 v cy
vo D, thu c 200 vt tan. Phage th h con c pha long 10-6 v cy
vo chng C thu c 30 vt tan. S ti t hp gia th t bin 1 v 2
l g ?

Chng 10
t bin phn t v m di truyn
A. Khi xc nh trnh t baz, lun phi vit mi b ba quy nh
mi axit amin.
V d: Mt chui polypeptit c trnh t axit amin nh sau:
ala-pro-ser-lys-gly-gly-trp
mARN c th c nhng trnh t nh th no?

94

Hy vit tt c cc b ba c th c. Ch X c th l mt trong s bn
loi baz.
ala -

pro -

ser - lys - gly - gly - trp

GCX - CCX - UCX - AAA - GGX - GGX- UGG

AGU - AAG
AGC
Ch rng ch c mt b ba duy nht quy nh trp v lys c 2 b ba.
Cc axit amin khc c t nht l 4 b ba. arg, ser v leu c th c 6 b ba.
B. Lun phi tm nhng thay i ti thiu, sinh ra t
bin.
t bin l s kin him v thng ch c mt baz b bin i bng
cch thm vo hay mt i.
V d: Mt protein c gly v tr xc nh. Ngi ta tch c 4 t bin,
v, thay vo ch gly h thu c ala, arg, trp v leu. Nhng thay i no
c th xy ra vi cc b ba v b ba no c th c s dng m ho
gly?
Chng ta tm tt kt qu thu c trn y nh sau:
gly
ala

arg

trp

leu

Hy vit cc b ba c th tham gia m ho cc axit amin :

GGX
GCX

CGX

UGG

CUX

AGG

UUA

AGA

UUG

Ta hy phn tch t n gin n phc tp. B ba m ho trp phi l

UGG. C l c s bin i n gin t GGG sang UGG. V vy b ba
quy nh gly c l l GGG. Hy xt ala: S thay i baz th hai c th
cho ra ala. V gly l GGG, ala s phi l GCG. Nu arg l AGG th s bin
i t G thnh A v tr baz th nht s cho ra AGG. B ba m ho arg
khng th l AGA v nh vy phi cn thay i hai baz t GGG. Arg
cng c th do CGG nhng khng phi l CG(U, A, C) m ho. Nh vy arg s
do AGG hoc CGG m ho. Vi leu c kh khn hn v cc b ba m ho
leu khc vi gly t nht hai baz. Do vy b ba m ho leu c th l CUG
hoc UUG. Hoc, leu c th l kt qu ca t bin dch khung.
C. Nu ch c mt axit amin b bin i th c th l t
bin nhm ngha.
V d: Mt t bin ca chui peptit mc A c trnh t axit amin nh
sau:
95

bnh thng: ala-pro-ser-lys-gly-gly-trp

t bin:

ala-pro-ser-glu-gly-gly-trp

Bn cht ca t bin ny l g?
Chng ta thy ch c 1 axit amin b bin i: lys thnh glu. Cch gii
thch n gin nht l b ba m ho lys b t bin. Liu c phi
l kt qu do thay i 1 baz? Cc b ba m ho lys l AAA hoc AAG;
glu l GAA hoc GAG. S thay i baz th nht t A thnh G c th
sinh ra t bin trn.
D. Nu protein c hn mt axit amin b bin i th c th
l kt qu ca t bin dch khung (do thm hoc mt mt
baz).
V d: Hy xt mt protein bnh thng v mt protein t bin di y:
bnh thng: ala-pro-ser-lys-gly-gly-trp
t bin:

ala-pro-ile-lys-gly-arg-leu

Xc nh trnh t baz cho mi protein?

Hy xt axit amin u tin b bin i (ser thnh ile), v cng lu
rng c tt c 3 axit amin b bin i.
ser: UCX

ile: AUU

AGU

AUC

AGC

AUA

Ch bng cch thay i 1 baz, chng ta khng th chuyn cc b

ba quy nh ser thnh b ba quy nh ile. Thm baz A vo trc b ba
UCX ca ser ta c AUCX, v c AUC m ho cho ile. Do vy cc trnh t
baz s l:
bnh thng: GCX CCX UCA AAG GGC GGC UGC
t bin:

GCX CCX AUCAAA GGG CGG CUG C

Kim tra trnh t m ho, chng ta thy cc axit amin cn li th t

bin ng vi trnh t trn.
E. Mt protein c trnh t axit amin bnh thng nhng b ct
ngn c th l do mt trong s cc b ba b t bin thnh
b ba kt thc (b ba stop).
V d: Chui peptit mc A v th t bin ca n c trnh t sau:
bnh thng: ala-pro-ser-lys-gly-gly-trp
t bin:

ala-pro-ser

Bn cht ca t bin ny l g?
Trnh t axit amin dng li ser cho thy rng b ba quy nh lys
b t bin thnh b ba kt thc. Lys c m ho bi AAA hoc AAG.
96

Baz A u tin b bin i thnh U s sinh ra UAA hoc UAG, u l

cc b ba kt thc.
F. Mt protein c trnh t axit amin di hn bnh thng l do t
bin xy ra b ba stop.
V d: Hy xt hai on trnh t sau y u C ca hai protein:
bnh thng: ser-thr-lys-leu
t bin

: ser-thr-lys-leu-trp-tyr-gln

iu g xy ra?
mARN bnh thng phi c b ba stop (UGA hoc UAG) sau b ba quy
nh leu. Nu baz th hai hoc th ba ca du hiu kt thc ny thay
i (v d, A thnh G) s sinh ra b ba m ho cho trp. Kh nng khc l
AG c thm vo sau G ca b ba kt thc UGA.
G. t bin b ba i m (anticodon) trn tARN c th lm thay
i trnh t axit amin.
tARN b t bin c th cng vn s mang cng loi axit amin nh tARN
ban u, song n c th xen axit amin vo b ba khc.
V d: Mt t bin c tm thy b ba i m ca tARN nh sau:
bnh thng: 5 GUA 3
t bin

: 5 UUA 3

Axit amin no s c xen vo v n c xen vo u?

Trc ht chng ta phi xc nh xem tARN ny s mang axit amin no.
Trnh t 5 GUA 3 b tr vi 3 CAU 5 hoc 5 UAC 3 - b ba quy nh
tyrosine. By gi, hy vit trnh t b tr cho b ba i m mi: 5 UUA
3. B ba mi s l 5 UAA 3 - mt b ba stop. Tyrosine s c xen vo
im kt thc ny v protein s di hn bnh thng.
Tm tt
1. Hy vit cc b ba c th c ri tm nhng b ba ch sai khc nhau
mt baz.
2. Ch thay i mt axit amin thng l kt qu ca t bin nhm
ngha, cn s thay i nhiu axit amin thng l do t bin dch
khung.
3. Cc protein c trnh t axit amin ngn hn bnh thng thng l do
t bin kt thc sm.
4. Cc protein c trnh t axit amin di hn bnh thng thng l kt qu
t bin b ba stop.

97

Bi tp
1. Hemoglobin ca t bo hng cu hnh lim c glu b thay th bng
val. B ba ca n c th b bin i nh th no?
2. Trong nhng protein t bin di hn bnh thng, ngi ta tm thy
nhng axit amin u tin c thm vo gm gln, lys, glu, ser, leu, tyr,
arg, trp, gly hoc cys. Ti sao li ch l nhng axit amin ?
3. Hy xem xt trnh t ca protein bnh thng:
gly-leu-arg-gln-cys-ile-phe
v ca cc protein t bin:
a. gly-leu-arg-gln
b. gly-leu-arg-arg-cys-ile-phe
c. gly-phe-lys-thr-met-gln-ile
Bn cht ca mi t bin trn y l g v trnh t ca mARN
bnh thng?
4. Cc polymer c trnh t cc baz lp li c s dng trong mt h
thng tng hp protein in vitro. Hy tin on trnh t axit amin do cc
polymer c trnh t lp li di y to ra:
a. (UC)
b. (UUC)
c. (CAA)
d. (AUA)
e. (AUG)
5. Nu m di truyn c c theo kiu trm nhau v khi ribxm dch
chuyn mt baz l chuyn sang mt b ba mi th s c bao nhiu axit
amin b bin i do mt t bin ng hon n l (ch thay th
mt baz) ?
6. Hy xc nh trnh t axit amin do cc trnh t nucleotid di y quy
nh:
a. 5 AUGCAGUGCAAUCCUGCA 3
b. 5 AUGCAGUCCAAUCCUGCA 3
c. 5 AUGUAGUGCAAUCCUGCA 3
98

d. 5 AUGCAGCUGCAAUCCUGCA 3
e. 5 AUGCAGUGUAAUCCUGCA 3
7. Mt protein bnh thng c arginine v tr xc nh. Ngi ta tm
thy cc t bin thay th axit amin bng cc axit amin sau: ile,
thr, ser, gly. B ba quy nh arg phi l b ba no?
8. Mt tARN c trnh t i m 5 CCC 3; mt t bin ca n c trnh
t 5 CCG 3.
a. tARN bnh thng mang axit amin no, tARN t bin mang axit
amin no?
b. tARN t bin s nhn bit b ba no v hu qu s ra sao.
9. Mt th t bin do tc nhn gy t bin dch khung sinh ra th
c th c hi bin nh ho cht gy t bin ng hon khng?
10. Hydroxylamine ch gy ra t bin ng hon CT trn ADN.
Nhng t bin sinh ra b ba stop c th c hi bin nh
hydroxylamine hay khng?
Enzym thinkase c trnh t axit amin t v tr 25 - 33 nh sau:
25

26

met ala

27

28

29

30

31

32

33

phe

ser

his

arg

glu

val

leu

Mt t bin (do tc nhn t bin ch gy ra t bin thm vo

hay mt i mt baz) sinh ra mt protein ch cn 30 axit amin nh sau:
25 26

27

met

28

ser

29

leu

30
tyr

ser

pro

24 axit amin ng trc vn bnh thng. Chnh tc nhn gy t bin

trn c dng phc hi t bin v thu c protein c trnh t sau
(axit amin 1-24 v t 34 tr i vn bnh thng):
25 26

27

met ser

28
leu

29
tyr

30

31
ser

32
pro

33
trp

val

leu

Hy xc nh trnh t mARN bnh thng quy nh on protein ny.

99

Chng 11
Kim sot hot ng ca gen
A. t bin vng kim sot s ch tc ng n on ADN ni
n nh v (hin tng tri u - cis domonant).
Cc t bin vng kim sot (nh gen khi ng (promoter) v gen
ch huy (operator)), thng s ch tc ng n on ADN m n l mt
phn trong ; cc t bin ny s khng nh hng n cc gen trn
cc phn t ADN khc.
V d: Trong mt operon, c cc gen p a b c (p = promotor; a,b,c = cc
gen cu trc). Mt chng vi khun c kiu gen p- a+ b+ c+ s c kiu hnh
nh th no? (p+ = promoter hot ng chc nng bnh thng, p- =
promoter b hng, a+, b+, c+ = gen hot ng chc nng bnh thng).
Promoter b hng nn ARN polymerase khng gn kt c, do
khng tng hp c ARN. Operon s khng hot ng.
V d: Mt operon gm cc gen p o r s t (p = promoter, o = operator, r, s,
t = cc gen cu trc). Chng sau y: p+ o- r+ s+ t+ c operator b hng,
sai hng s gy hu qu nh th no? Nu operator l oc hoc o- (oc
v o- = operator b hng khng cho cht c ch (repressor) gn vo gen
ch huy) th hu qu s ra sao?
Gen ch huy b hng nn cht c ch khng th gn vo n. V th
s khng cn c ch iu ho, operon s hot ng lin tc.
B. t bin cht c ch c th nh hng n ton b ADN trong
t bo (hin tng tri lch - trans dominant)
Cc phn t cht c ch c kh nng tng tc vi mi phn t ADN
trong t bo k c ADN m ho vic tng hp ra n.

100

V d: Trong cc operon di y, r = gen c ch, p = promoter v a, b, c

l cc gen cu trc. Hy xc nh xem operon no trong cc operon di
y hot ng lin tc, khng hot ng hoc hot ng iu ho?
1. r-p+o+a+b+c+
2. r+p-o+a+b+c+/r-p+o+a+b+c+
Cc t bo chng 1 s sinh ra cht c ch khng hot ng chc nng,
chng s khng gn vo gen ch huy. i vi chng 2, chng ta phi xem
xt tng ADN ring bit, sau kt hp li. ADN ng trc s khng tng
hp c ARN v promoter b hng. ADN ng sau s lin tc tng hp ARN
v gen c ch b hng. ADN ng trc s sinh ra cht c ch gn vo c
hai gen ch huy. V th, chng ny s hot ng iu ho.

C. Sai hng gen cu trc ch gy ra t bin trn ADN.

V d: Hy xt operon mc B. Gi s cc gen a, b v c u cn
tng hp mt protein. Nhng chng no di y c kh nng tng hp
protein v trong iu kin no?
1.p+ o+ a+ b- c+
2.p+ o+ a- b+ c+/p+ o+ a+ b- c+
3.p- o+ a+ b+ c+/p+ o+ a+ b+ cChng 1 tng hp c ARN, song lun sinh ra sn phm sai hng b.
Nh vy n s khng tng hp c protein cn. Chng 2 tng hp c
protein , v ADN ng trc tng hp c b v c, ADN ng sau tng
hp c a v c; chng b sung cho nhau. Chng 3 khng tng hp c
protein . ADN ng trc khng th tng hp c ARN v n mang p-.
ADN ng sau tng hp c ARN song sn phm protein ca n lun sai
hng c. Kt hp li, n khng tng hp c protein cn thit.
D. xc nh kiu hnh t kiu gen, hy xt ring tng ADN v
xc nh xem trong nhng trng hp no n s khng sinh ra ARN
hoc enzym c hot ng chc nng.
V d: Cc locut gen di y cng tham gia vo operon lac:
z = gen cu trc m ho -galactosidase
i = gen c ch
o = operator
Hy xt xem cc t bo c cc kiu gen di y c th tng hp c
enzym khng, v nu c th trong iu kin no?
101

 -galactosidase ( + = c, - = khng)
Kiu gen

khng c lactose

c lactose

i- o+ z+
i+ o+ z-/i- o+ z+
i+ o+ z+/i+ o- zChng 1 sinh ra cht c ch khng hot ng, v th operon ny lin
tc hot ng (+, +). i vi chng 2, ADN ng trc s khng sinh ra sn
phm z tt, song li sinh ra cht c ch tt. Cht c ch ny gn vi c
hai gen ch huy, v vy n iu ho hot ng ca operon., (-, +). Vi
chng 3, ADN ng sau khng sinh ra sn phm z tt, chng ta c th b
qua. ADN ng trc bnh thng, v vy operon s c iu ho (-, +).
E. Vi cc operon chu c ch th cht c ch phi gn vi cht
ng c ch c hot ho.
V d: Operon trp - mt h thng chu c ch, c trnh t sau:
p o E D C B A // p R
(p = promoter, o = operator , A - E = cc gen cu trc, R = gen ngoi h
thng (khng lin kt vi cc gen trong operon trp) quy nh cht c ch)
Nu cc gen u hot ng bnh thng th trong iu kin no
operon s khng hot ng?
Cht c ch khng t gn c vo operator. Trc ht n phi gn vi
tryptophan hot ho. u tin, operon hot ng v sinh ra
tryptophan. Tryptophan kt hp vi cht c ch, hot ho n n c th
gn vi operator v kho operon li.
F. S phin m nhiu operon vi khun c th b dng li do hin
tng phin m d - hin tng rt nhy cm vi mc tARN mang
axit amin trong t bo.
V d: i vi operon his, mARN y ca n s c to ra khi trong t
bo khng c histidine. Khi c mt lng nh histidine, ARN y v cc
ARN c kch thc ngn cng c to ra. Khi c mt lng ln histidine, rt t
ARN c hnh thnh. Hy gii thch.
Khi khng c histidine, operon c m v qu trnh phin m tip tc
cho n ht operon. Mt lng nh histidine s lm cho cc tARN his gn vi
vng suy gim ca operon, sinh ra cc ARN ngn. Mt lng ln histidine s
hot ho cht c ch, cho php chng gn vo gen ch huy v ng
operon li.

102

G. cc sinh vt nhn chun, cc gen tng cng (enhancer) c th

iu chnh s bn sao ca gen.
V d: Nhng t bin mt on nm pha trc promoter mt gen sinh
vt nhn chun c biu din nh hnh di y v s lng phn t ARN
do chng to ra. Nu k hiu /// = vng b mt, hy gii thch kt qu thu
c.
-150
ARN to ra

-100

-75

-50

-25

gen

100

2 /////////////////
3
4

s phn t

100
//////////////////

10

/////////////////

50
//////////

100

Chng ta thy t bin 3 v 4 gy suy gim phin m. iu cho

php suy on rng vng nm gia -125 v -75 c cha gen tng cng.
t bin 3 gy suy gim phin m nhiu nht, vy c l gen tng cng
nm gia vng -125 v -90.
H. Cc gen sinh vt nhn chun hot ho thng mn cm vi
ADNase I.
Cc gen ang phin m c l c cu trc khng gian lng lo hn cc
gen khng phin m, nn n d b ADNase I tn cng.
V d: ADN tch chit t gan, c v nguyn hng cu c x l vi ADNase
ri sau tch gen quy nh -hemoglobin. Khi kim tra hot ng ca
gen, ngi ta thu c kt qu di y ( + = c gen -globin, - = khng c):

C/khng c -globin

Gan

Nguyn hng cu

Hy gi thch kt qu thu c.
Nu tch c gen sau khi x l ANDase chng t gen khng
hot ng phin m. Nh vy, gen -globin khng phin m gan v
thn, nhng phin m nguyn hng cu.
103

I. i vi cc gen sinh vt nhn chun, cc intron phi c ct b

trc khi to ra ARN hon chnh.
Nu cho mARN hon chnh lai vi ADN lm khun cho n th s xut
hin cc vng trn ADN ni nh v cc intron nh trn s di y (
= ADN , ---- = ARN).

V d: Gen m ho enzym thinkase c tch ra v cho lai vi mARN ca

thinkase trong t bo cht. Kt qu thu c c minh ho bng hnh di
y:

C bao nhiu intron trong gen thinkase?

Chng ta thy c hai vng ADN khng c cc baz b tr trn ARN. Cc
vng ny phi tng ng vi cc intron b loi b. Vy gen ny c hai intron.

Tm tt
1. Cc protein iu ho c th gn vi mi ADN trong t bo.
2. t bin promoter v operator ca gen no th ch nh hng n
gen .
3. xc nh kiu hnh chung, phi bt u bng vic xc nh
kiu hnh ca tng phn t ADN ring bit ri xem xt hiu ng tng
hp ca c hai.
4. Cc gen tng cng thng nm xa gen m n tc ng n.

104

5. Cc gen sinh vt nhn chun thng cha cc intron v cc gen hot ho

rt nhy vi ADNase I.

Bi tp
1. Nhng locut gen di y cng tham gia vo operon lac:
z = gen cu trc quy nh -galactosidase
o = gen ch huy
p = promoter
i = gen c ch
i*= gen sinh ra cht c ch khng gn kt c vi lactose nhng c th
gn kt vi ADN.
Hy xem xt cc kiu gen hon chnh bng di y.
Sinh ra -galactosidase (c:+ hoc khng: -)
Kiu gen

Khng c lactose

C lactose

1 i+p+o+z+
2 i+p-o+z+
3 i-p+o+z+
4 i+p-o+z+/i+p+o+z5 i+p-o-z+/i+p+o+z6.i*p+o+z-/i-p+o+z+

2. Cc k hiu a, b v c l ch gen quy nh -galactosidase, gen

ch huy v gen c ch operon lac, nhng khng nht thit theo trt t
trn. T nhng s liu di y hy xc nh ch ci no dng ch
gen no.
Kiu gen

khng c lactose

1 a+b+c+

c lactose
+
105

2 a+b+c-

3 a+b-c+

4 a+b-c+/a-b+c-

5 a+b+c+/a-b-c-

6 a+b+c-/a-b-c+

7 a-b+c+/a+b-c-

3. Hy vit kiu gen lng bi mt phn s sinh ra -galactosidase nh cm

ng v enzym permease trong mi iu kin vi operon lac.
4. a. Hy cho bit hai loi t bin khc nhau ca operon lac sinh ra galactosidase mt cch c nh.
b. Mi t bin s hot ng nh th no trong trng thi lng bi mt
phn vi dng kiu di?
5. Bn phi xc nh mt operon mi, chu cm ng, trong mt vng k
nhau theo trt t PQRS. Operon ny sinh ra hai enzym, 1 v 2. t bin
mi vng s sinh ra cc enzym sau y ( + = ch c sinh ra khi c
cht cm ng, 0 = khng c enzym, C = enzym c to ra mt cch c
nh)
t bin

enzym 1

enzym 2

P-

Q-

R-

S-

a. Vng no l cc gen cu trc v vng no l vng iu ho?

b. Da vo li gii ca phn a, hy tin on s c mt ca enzym 1 v 2
(+, 0 hoc C) cc th lng bi mt phn di y:
Trng hp 1: P+Q-R+S+/P-Q+R+S+
Trng hp 2: P-Q+R+S+/P+Q-R+S6. on dn u nm gn vng suy gim ca operon trp c cha trnh t
5 UGGUGG 3. Nu trnh t b i thnh 5 AGGAGG 3 th hu
qu s nh th no?
7. Mt gen sinh vt nhn chun di 1,2 kb. t bin mt an di 0,5 0,7 kb vng di gen hoc mt t bin im 0,3 - 0,4 kb vng trn gen
u lm lng mARN do gen ny tng hp gim xung 10 n 50 ln.
Hy gii thch.
8. Operon trp l operon chu c ch v gen sinh ra cht c ch khng do
gen ny m ho. Cht ng c ch l tARN gn kt tryptophan
106

(tARNtrp) v gen m ho n khng thuc operon ny. Hy xem cc t

bin di y c sinh ra enzym khng v t bin no c th l tri
trng thi ng (cis dominant) khi trng thi lng bi mt phn.
a. ARN polymerase khng gn c vo promoter.
b. Cht c ch khng gn c vo gen ch huy.
c. Cht c ch khng gn vi tARNtrp.
9. Ngi ta dng mARN ca mt gen biu hin gan v tu xng to
phn t lai ARN - ADN. ADN dng cho th nghim lai ny l ADN cu
gen c di y . Da trn kt qu th nghim di y, hy gii
thch cc dng phn t lai thu c.
(

= ADN, --- = ARN).

Gan
tu xng

10. Mt operon gi thuyt c trnh t A B C D E, nhng trnh t ca gen

ch huy v promoter cn cha xc nh c. Gen quy nh cht c ch
nm xa operon ny. t bin mt on nhng phn khc nhau ca
operon c tch v lp bn . Cc mt on ny c xc nh nh
sau:
A

Mt on
1
2
3
4
5

/////////////////////
////////////////
//////////////////
////////////////////
/////////////

//////////
Mt on 2 v 3 sinh ra ARN c mc c nh. Mt on 4 v 5
khng tng hp ARN. Xc nh xem promoter v gen ch huy nh
v u?
107

11. ADN ly t cc giai on pht trin khc nhau ca phi v t cc m

trng thnh c x l bng ADNase. Sau chng c kim tra s c
mt ca gen A v B. ( + = c, - = khng). Da trn cc kt qu thu c,
hy xc nh hot tnh ca gen A v B.
Trng th tinh

Phi v

Giai on

Gan

Tu xng
ng thn kinh
Gen A

Gen B

+
12. Mt gen c 3 intron. ARN t bo cht ly t mt c th khng c
enzym ca gen ny c cho lai vi ADN ca gen. Khng c vng no c
to ra th lai ARN - ADN. Hy gii thch kt qu thu c.
13. Gi s vic khng tng hp c insulin nhng ngi b bnh tiu ng c th do sai hng promoter ca gen insulin hoc do mt on
gen ny. Hy xy dng th nghim phn bit hai nguyn nhn
ny.

108

Chng 12
Sinh hc phn t v thao tc vi ADN
A. Nu s nucleotid A khng bng T, hoc G khng bng C th
ADN l mch n.
V d: T t l phn trm baz cc phn t ADN di y, hy xc nh
xem n l mch n hay mch kp.
A

ADN 1

32

18

31

19

ADN 2

17

28

31

24

ADN 3

29

25

29

17

Phn t 1 l mch kp v A = T v G = C. Phn t 2 v 3 l mch

n v A khng bng T v G khng bng C.
B. Hm lng G-C cao th nhit gy bin tnh cao.
Mi cp G-C c 3 lin kt hydro nn cn nhiu nng lng hn ph
hu.
V d: Phn t ADN 1 bin tnh 74 oC v phn t ADN 2 bin tnh
81oC. Hy so snh hm lng G-C ca chng.
Phn t 2 c hm lng G-C cao hn v n bin tnh nhit cao
hn.
C. Phn t ADN mch vng c n im gii hn s to ra n on khi
x l bng enzym gii hn.
V d: Khi ct plasmid pBR322 bng HaeI, c 11 on ADN c to ra. Khi
ct n bng BamHI, ch c mt on c to ra. C bao nhiu Im gii
hn cho mi loi enzym?
ADN ca plasmid c mch vng. Vy c 11 im gii hn cho HaeI v 1
cho BamHI.
D. ADN mch thng c n im gii hn s to ra n+1 on.
109

V d: Mt phn t ADN mch thng c ct bng EcoRI v to ra cc

on 3 kb, 4,2 kb v 5 kb. Hy xc nh bn gii hn ca n.
C ba on, vy phi c 2 im gii hn. Cc im c th phn
b nh sau:
3
3
5

4,2

4,2
5

4,2

E. t bin im gii hn c th lm mt mt hoc mt s on

v lm xut hin mt on mi, di hn.
V d: Khi x l mt gen (on ADN) bng BgIII, n to ra cc on 1,7 kb,
2,1 kb v 3,2 kb. Khi x l on ADN tch ra t mt t bin khng tng
hp c enzym ca gen ny bng BgIII, ngi ta ch thy hai on 3,2 v
3,8 kb. iu g xy ra?
t bin lm thay i mt hoc mt s baz ti im nhn bit
ca BgIII, v, nh vy, enzym ch ct phn t mt ln. on mi l tng
ca hai on 1,7 v 2,1 kb, cho thy hai on ny nm k nhau trn
phn t ADN bnh thng.
F. t bin to ra im gii hn mi s lm mt i mt on c v
xut hin hai on mi ngn hn.
V d: Ta li xt on ADN mc E. Khi x l AND c tch ra t mt t
bin khc bng BgIII, ngi ta thy xut hin cc on 1,3 kb, 1,7 kb, 1,9
kb v 2,1 kb. Hy gii thch kt qu thu c.
Chng ta thy cc on 1,7 v 2,1 kb cng c c ADN bnh thng v
ADN t bin. im gii hn sinh ra hai on ny phi cn nguyn vn
trn AND t bin. Hai on cn li, 1,3 v 1,9 kb c tng l 3,2 kb,
ng bng on b mt. Vy, mt im gii hn mi c to ra ADN
gc.
G. Nu mt on do enzym gii hn to ra cng xut hin khi x l
bng mt enzym v bng hn hp hai enzym th on khng
c im gii hn cho enzym th hai.
V d: Mt phn t ADN mch thng c x l bng cc enzym di y
v kt qu nh sau:
EcoRI: 1,7; 2,1; 3,2 kb.
HaeI: 1,0; 1,4; 4,6 kb.
Hn hp: 0,7; 1,0; 1,4; 1,8; 2,1 kb.
Nhng on gii hn no ca EcoRI khng cha cc im gii hn ca HaeI
v ngc li?
110

on 2,1 ca EcoRI cng xut hin trong hn hp, vy , n khng c

im gii hn ca HaeI. Cc on khc ca EcoRI mt i khi x l bng c
hai enzym, nn chng phi cha cc im gii hn ca HaeI. Tng t, cc
on 1,0 v 1,4 ca HaeI u khng cha cc im gii hn ca HaeI.
H. lp bn gii hn, phi xem xt cc on gii hn c to ra
khi x l bng hai enzym vi cc on to ra khi x l tng enzym
ring bit.
V d: Hy xt li cc on mc G v xy dng bn gii hn.
V 1,0 + 0,7 = 1,7, v, 1,4 + 1,8 = 3,2, nn on 1,7 ca EcoRI phi
c im gii hn cho HaeI t mt u:
E 1,0

H 0,7

Tng t, on 3,2 cng phi c im gii hn cho don HaeI di 1,8 t

mt u:
E

1,8

1,4

By gi, hy xp xp trt t c th c cho cc on EcoRI:

1,7

2,1

1,7

1,7

2,1

3,2

3,2

3,2

2,1

Hy thm cc im gii hn bit ca HaeI vo ng v tr. Mt

trong nhng trt t c th l:
E

2,1

1,0

0,7

1,4

1,8

Chng ta nhn c on 3,1 kb khi x l ADN ch bng HaeI. Nu trt

t l 2,1; 0,7; 1,0; 1,4 v 1,8, th khi x l bng HaeI phi to ra on 2,8
kb. Vy, on 1,7 kb khng nm gia. Nu on 3,2 kb nm gia,
chng ta phi gp, hoc on 3,5 kb, hoc on 3,9 kb khi x l bng
HaeI. iu khng xy ra, vy trt t phi l:
E

1,7
1,0

0,7
H

E
2,1

3,2
1,8

1,4

Nu im gii hn ca HaeI l 0,7 kb t mt u, th ta phi gp

on 0,7 kb khi x l ADN bng HaeI. Bin lun tng t cho on 1,8 kb.
111

I. T bo thu nhn plasmid phi biu hin tnh khng lin quan
n plasmid .
V d: Trong cc th nghim bin np in hnh, cha n 1% s t bo
thc s nhn c plasmid. Bng cch no bn c th tch c s t bo c
plasmid?
Hy thit k cc plasmid c cha mt hoc mt vi gen khng vi cht
khng sinh. Trn cc t bo vi plasmid ny v cy trn mi trng c cha
cht khng sinh. Ch nhng t bo nhn c plasmid mi sinh trng c.
J. Vic xen mt on ADN ngoi lai vo gen thng lm mt chc
nng ca gen.
V d: Mt plasmid cha cc gen khng ampicillin v tetracycline. Plasmid
cha mt im gii hn nm trong gen tetracycline. Plasmid c
x l vi enzym gii hn trn vi ADN ngoi lai, ri em vi E. coli. Bng
cch no bn c th tch c cc t bo c plasmid lai?
Cy cc t bo trn mi trng c cha ampicillin v chn cc t bo
sinh trng c. Cc t bo c plasmid s c gen ampicillin trn vn. Cy
sao chp cc t bo ny vo mi trng cha tetracycline. Nhng t bo
khng sinh trng c l nhng t bo mang cc gen c on xen. Tr li
vi cc t bo gc trn mi trng cha ampicillin ly ra nhng t bo
khng sinh trng c trn mi trng cha tetracycline.
K. ARN nh du phng x hoc cADN c th c dng nh cc vt
d v chng s gn vi on ADN t chng c sinh ra.
Cc phn t s b tr vi ADN v to ra on lai c th pht hin
c.
V d: X l ADN h gen ngi vi EcoRI ri phn tch cc on to ra bng
in di. Cc bng ADN c chuyn sang mng lc ri vi cADN m ho
insulin nh du phng x. Kt qu nh sau: (* = cc bng c nh du
phng x).

*
*
C th kt lun g t th nghim ny?
Hai bng c nh du * c cha cc trnh t b tr vi cADN insulin,
v, chc l nhng phn ca gen insulin. S c mt ca hn 1 bng
cho thy gen ny c cha im gii hn ca EcoRI.
L. xc nh trnh t ca gen, phi c t gc bn gel.
112

V d: Mt phn t ADN c gii trnh t bng phng php MaxamGilbert. Hy xc nh trnh t ADN t bn gen thu c sau:
A

A+G

C+T

Cc baz u 5 s gc bn gen. V n ch xut hin ct C+T,

nn n phi l T. Cc bng tip theo xut hin c ct C v C+T, nn n
phi l C. Trnh t chung l 5TCATCGCC 3.

V d: Mt phn t ADN c gii trnh t bng phng php dideoxy. Da

trn bn gel thu c di y, hy xc nh trnh t si lm khun.
ddA

ddG

ddC

ddT

113

Phi c t gc bn gel, v phi nh rng u 5 di cng. Vy trnh

t s l 5CCAGTTCA 3. l trnh t ca si mi c tng hp. Vy trnh
t si khun l 3GGTCAAGT 5.
Tm tt
1. Phn t ADN si n c hm lng A/T v/hoc G/C khng bng nhau.
2. Nhit bin tnh t l vi hm lng G-C.
3. C th xy dng bn gii hn bng cch b xung cc on thu c
do x l hai enzym gii hn.
4. t bin s lm thay i kch thc cc on gii hn.
5. on ADN xen vo mt gen s ph hu chc nng ca gen .
6. ARN nh du phng x hoc cADN c th dng lm vt d xc
nh trnh t ADN.
7. Trnh t ADN c xc nh bng cch c bn gel t gc tr ln.
Bi tp
1. Hai phn t ADN, A v B, c nhit bin tnh tng ng l 73oC v
81oC. C th kt lun g v thnh phn baz ca chng?
2. Nu mt phn t ADN c A/T = 1,5 v G/C = 1,0 th bn cht ca
phn t ADN ny l g?
3. Xc nh xem cc phn t di y l ARN hay ADN, mch n hay
kp?
T l phn trm
A

27

26

23

24

33

17

21

29

33

21

29

17

21

29

29

21

4. Phn u ca mt gen c tch v gii trnh t bng k thut

Maxam-Gilbert. Kt qu nh sau:

114

A+G

C+T

a. Trnh t ca on ny? (Nh ch r u 5, 3).

b. Trnh t ca ARN c tng hp t si ny?
c. Bn axit amin u tin ca protein?
5. Nu mt si ADN c trnh t 3 GCTGAACGTCAG 5 c vi
triphosphate nh du phng x v 10% dideoxy CTP th s xut hin
nhng on c nh du c kch thc bao nhiu?
6. Mt phn t AND mch thng c ct bng PstI, sinh ra cc on 4,3,
5,1, v 6,6 kb. Xc nh cc bn gii hn ph hp vi kt qu ny.
7. Mt gen c bn gii hn sau y cho enzym BgIII:

115

2,1

2,8

1,2

Xc nh cc on s c to ra t bin, trong :
a. im gii hn nm gia hai on u tin b t bin.
b. Mt im gii hn mi c to ra cch u tri ca gen 2,7 kb.
8. Mt ngi ph n 30 tui c thai bit rng cha b b bnh ma
Huntington
- mt bnh do gen tri trn nhim sc th thng
nh. Khng c lch s bnh gia nh chng b. Th nghim
ADN bng enzym gii hn cho thy c mt alen ca bnh ny.
nghim ny c tin hnh trn b b, b, chng b v bo thai.
qu nh sau:
Cha

Chng

Ngi ph n

Bo thai

git
quy
ct
Th
Kt

a. Ngi ph n c b bnh khng?

b. Con b c b bnh khng?
9. Ct ADN mch thng bng cc enzym A v B to ra cc on sau
( tnh bng kb):
A: 1,1; 2,0; 2,7; 3,2
B: 1,5; 3,5; 4,0
Hn hp hai enzym: 0,3; 0,8; 1,1; 1,2; 2,7; 2,9

Bn no di y khng ph hp vi kt qu trn:
B

2,7

4,0

1,5

3,5

4,0

1,5

3,5
1,1 3,2
2,0

2,0

2,7

1,1

3,2

116

A
(a)

(b)
B

1,5
1,1

B
4,0

3,2

2,0

3,5

2,7

(c)
10. Mt phn t ADN b ct bng EcoRI sinh ra mt on 16 kb. Khi
ct phn t ban u bng BgIII, thu c cc on 4,3; 5,5 v 6,2 kb.
Khi x l bng c hai enzym, thu c cc on 2,7; 3,5; 4,3 v 5,5 kb.
a. Bn cht vt l ca ADN?
b. Xy dng bn gii hn.
11. Mt gen di 4,2 kb c xen vo im gii hn EcoRI ca mt plasmid
7,4 kb. Sau , plasmid c x l bng HindIII, to ra cc on 3,8; 4,9
v 2,9 kb. Khi x l bng c EcoRI v HindIII, thu c cc on 1,1; 1,4;
1,5; 2,7 v 4,9 kb. Nhng on no cha thng tin ca gen?
12. Mt on ADN ngi 3,0 kb c trn vi plasmid 7,0 kb cng b ct
bi cng loi enzym sinh ra on ADN ngi. Sau tch cc phn t
bng in di, thu c cc bng tng ng 3, 6, 7, 10 v 14 kb.
a. Hy gii thch?
b. Phn t no c dng bin np t bo?
13. Mt plasmid 6,0 kb c cha cc gen khng ampicillin, tetracycline v
chloramphenicol. N cn cha mt im gii hn cho mi loi ezym sau:
EcoRI, BgIII, SmaI v PstI. Mt gen gy ung th ca chut c tch v
xen vo mi im gii hn. Phn t lai c dng bin np E. coli v
cc th bin np c tnh da vo tnh khng vi cht khng sinh. Kt
qu nh sau (R = khng; S = mn cm).
Enym
Chloramphenicol
EcoRI
BgIII
SmaI
PtsI

Ampicillin

Tetracycline

R
S

S
R

R
R

a. V bn cc im gii hn cho mi enzym.

117

b. Mt plasmid khng c ADN ngoi lai c ct bng hn hp cc enzym di

y v thu c cc on sau (bng kb):
EcoRI + SmaI

0,8; 5,2

EcoRI + BgIII

1,3; 4,7

EcoRI + PstI

2,5; 3,5

EcoRI + SmaI + BgIII

0,8; 1,3; 3,9

EcoRI + SmaI + PstI

0,8; 1,7; 3,5

V li bn da cu a v ch r khong cch.
14. Mt phn t ADN mch thng c nh du bng 32P u 5. Sau
n c ct v to ra cc on nh sau (* = on c nh du
phng x)
HindIII
BgIII
Hn hp

3,0*; 4,0; 5,0*

1,8*; 2,2*; 2,7; 5,3
1,2; 1,5; 1,8*; 2,2*; 2,5; 2,8

V bn gii hn.
15. AND ly t phi chut v t bo limpho ca chut trng thnh c
ct bng enzym gii hn v c d bng cADN quy nh chui nh ca
phn t khng th c nh du phng x. Cc on nh du phng
x xut hin nh sau:
Phi

T bo limpho

Hy gii thch kt qu thu c.

118

Chng 13
Tn s alen v trng thI cn bng
Hardy -weinberg
A. Nu hai alen sinh ra ba kiu hnh khc bit th tn s alen c
th tnh bng cch ly tn s ng hp t + 1/2 tn s d hp t.
V d: Trong mt nhm 200 ngi, 160 ngi c nhm mu MM, 36 ngi MN
v 4 ngi NN. Tnh tn s alen N?, M?
Trc ht hy tnh tn s cc kiu gen:
Tn s kiu gen MM = 160/200 = 0,8
Tn s kiu gen MN = 36/200 = 0,18
Tn s kiu gen NN = 4/200 = 0,02
Tn s alen N, k hiu l q:
q = 0,02 + 1/2(0,18) = 0,02 + 0,09 = 0,11
Tn s alen M, (p):
p = 0,8 + 0,09 = 0,89
Vi mt gen c hai alen, nu bit tn s mt alen th tn s alen kia
bng 1 - tn s alen th nht, v p + q = 1.
Tn s alen cng cn c th tnh bng cch tnh trc tip s alen.
Mt qun th gm 200 c th s c 400 alen mi gen.
Tn s alen M, (hoc p) = [(160 X 2) + 36]/400 = 356/400 = 0,89
Tng t, tn s alen N, (hoc q) = [(4 X 2) + 36]/400 = 44/400 = 0,11
B. Nu mt qun th trng thi cn bng th cc kiu gen phi
c phn b: p2(A1A1) + 2pq(A1A2) + q2(A2A2).
Nu xc sut giao t l A1 bng p, th xc sut c hai alen
cng l A1 bng p x p = p2. L lun tng t cho q. C hai cch xut
hin d hp t, vy ta c 2pq.
V d: Gi s rng trong mt qun th ngi tn s alen li cong (p) l 0,4.
Tnh tn s cc kiu gen trng thi cn bng.
V p = 0,4, nn, q = 1 - 0,4 = 0,6. Vy khi qun th trng thi cn
bng, ta c:
(0,4)2 CC + 2 (0,4) (0,6) Cc + (0,6)2 = 0,16 CC + 0,48 Cc + 0,36 cc.

119

Ch rng, nu alen ln l ph bin trong qun th th cc c th

ng hp t ln chim a s, v s lng cc th d hp t c th vt s lng theo l thuyt tnh t php lai n tnh.
V d: Qun th no di y trng thi cn bng Hardy-Weinberg?
Qun th

A1A1

A1A2

A2A2

0,25

0,5

0,25

0,60

0,2

0,2

0,04

0,32

0,64

Trc ht chng ta phi tnh tn s alen cho mi qun th.

Qun th 1: p = 0,25 + 1/2(0,5) = 0,5; q = 0,5, v q = 1 - p.
Qun th 2: p = 0,6 + 1/2(0,2) = 0,7; q = 0,3.
Qun th 3: p = 0,04 + 1/2(0,32) = 0,2; q = 0,8.
By gi, s dng cc tn s alen tnh tn s cc kiu gen cho
trng thi cn bng. i vi qun th 1, cc tn s s l: (0,5) 2A1A1 =
0,25; 2(0,5)(0,5)A1A2 = 0,5 v (0,5)2A2A2 = 0,25. Tn s thc t ph hp vi
tn s l thuyt, vy qun th ny trng thi cn bng.
i vi qun th 2, tn s cc kiu gen trng thi cn bng l:
(0,7)2A1A1 = 0,49; 2(0,7)(0,3) A1A2 = 0,42 v (0,3)2 A2A2 = 0,09. Bn c th
tnh gi tr 2. Tuy nhin, trong trng hp ny iu khng cn thit
v sai khc gia tn s l thuyt v thc nghim qu ln (v d, 0,42 v
0,2 - tn s cc d hp t). Vy, qun th ny khng trng thi cn
bng.
Tng t, qun th 3 trng thi cn bng.
C. Nu mt qun th trng thi cn bng Hardy-Weinberg th
tn s alen ln (q) c th tnh bng cn bc hai ca tn s kiu
hnh ln.
V q2 = tn s kiu hnh ln, vy q = cn bc hai ca tn s .
V d: Trong s nhng ngi M da en, t l ngi c hng cu hnh lim
l khong 1/10000. Hy tnh tn s cc alen.
q=

1 / 10000

= 1/100 = 0,01

V p+q = 1, vy p = 1 - q,
p = 1 - 0,01 = 0,99
Bn cng c th tnh tn s l thuyt ca cc th d hp t
2pq = 2 (0,99)(0,01) = 0,0198
120

hoc khong 2%.

D. Nu qun th trng thi cn bng th tn s alen ln lin
kt X (q) c th tnh bng: s c th c mc bnh/ tng s c
th c.
Hy nh rng cc c th c ch cn mt alen ln trn X l biu hin
tnh trng.
V d: Mt qun th ngi trn o c 50 ph n v 50 ngi n ng, hai
ngi n ng b bnh m mu. Tnh tn s alen bnh m mu v tn s
ph n mang gen bnh.
Tn s n ng mc bnh l 2/50 = 0,04. Vy q, tn s alen bnh l
0,04. p = 1 - 0,04 = 0,96. Vy tn s ph n mang gen bnh l
2pq = 2 (0,04)(0,96) = 0,0768
hoc khong 8%.
E. Mt qun th c locut gen vi ba alen trng thi cn bng th
tn s cc kiu gen l: p2 (A1A1) + 2pq (A1A2) + 2pr (A1A3) + 2qr
(A2A3) + q2 (A2A2) + r2 (A3A3).
V p + q + r = 1, cc kiu gen s c phn b (p + q + r)2.
V d: Tn s cc alen IA, IB, IO ca h thng nhm mu ABO nhng ngi
M da trng l 0,28; 0,06; 0,66. Hy tnh tn s mi nhm mu.
K hiu p = tn s alen IA, q = tn s alen IB, r = tn s alen IO.
Nhm mu A c kiu gen IAIA v IAIO; vy tn s ca n bng p2 + 2pr =
(0,28)2 + 2(0,28)(0,66) = 0,4480 hoc khong 45%.
Tn s nhm mu B = q2 + 2qr = (0,06)2 + 2 (0,06)(0,66) = 0,828 hoc
3%.
Tn s nhm mu AB = 2pq = 2(0,28)(0,06) = 0,0336 hoc 3%.
Tn s nhm mu O = r2 = (0,66)2 = 0,4356 hoc 44%.
F. Tn s cc th ng hp t cao hn l thuyt l kt qu ca ni
phi.
Ni phi lm tng ng hp t bng vi mc gim d hp t.
V d: Tn s cc kiu gen A1A1, A1A2 v A2A2 mt qun th cch ly l
0,375, 0,25 v 0,375. Tnh tn s alen; xc nh xem qun th c trng
thi cn bng khng, nu khng, gii thch ti sao?
Trc ht phi tnh tn s alen:
p(A1) = 0,375 + 1/2(0,25) = 0,5
q(A2) = 1 - 0,5 = 0,5.
121

Nu qun th trng thi cn bng th tn s d hp t phi l 2pq =

2(0,5)(0,5) = 0,5 hoc 50%; tn s cc ng hp t u l (0,5)2 = 0,25.
Qun th khng trng thi cn bng, s d hp t gim ng bng s
tng cc ng hp t. Hin tng ni phi c th gii thch cho kt qu
ny, tuy nhin cn xem thm chng 14.
Ch : Ni phi c th lm thay i tn s kiu gen, nhng tn s alen
khng thay i.
G. Nu trong mt qun th c (f) c th ni phi th tn s cc
kiu gen c th tnh bng (p2 + fpq) (A1A1) + (2pq - 2fpq) (A1A2) +
(q2 + fpq) (A2A2).
Tn s ng hp t do ni phi bng fpq v do giao phi ngu nhin
bng q2 v p2.
V d: Trong mt qun th rui c 20% s c th ni phi. Cho q = 0,4,
hy tnh tn s cc kiu gen.
Ta c f = 0,2; q = 0,4 v p = 0,6.
Tn s kiu gen A1A2 = p2 + fpq = 0,36 + 0,048 = 0,408
Tn s kiu gen A2A2 = q2 + fpq = 0,16 + 0,048 = 0,208
Tn s kiu gen A1A2 = 2pq - 2fpq = 0,48 - 0,096 = 0,384
G. H s ni phi c tnh bng: 1 - (tn s d hp t quan st c/tn s d hp t theo l thuyt).
V d: Trong mt qun th yn mch hoang di, tn s ng hp t tri,
d hp t v ng hp t ln l 0,67, 0,06 v 0,27. Tnh h s ni phi.
Trc ht hy tnh tn s cc alen:
p = 0,67 + 1/2(0,06) = 0,7
q = 1 - 0,7 = 0,3
By gi, hy tnh tn s d hp t theo l thuyt:
2pq = 2(0,3)(0,7) = 0,42
H s ni phi = 1 - (0,6/0,42) = 0,86
y l h s ni phi cao cho thy hu ht yn mch trong qun th
ny sinh sn bng cch t th phn.
Ch : C cch khc na tnh h s ni phi l: (tn s d hp t theo
l thuyt - tn s d hp t quan st c)/tn s d hp t theo l thuyt.

122

Tm tt
1. Tn s alen c tnh bng: tn s ng hp t + 1/2(tn s d hp t).
2. Trong trng hp tri hon ton, ta tnh tn s alen bng cch gi s
qun th trng thi cn bng v q2 l tn s kiu hnh ln.
3. Hy nh rng tng tn s cc alen phi bng 1.
4. Ni phi lm gim s lng d hp t.

Bi tp
1. Mt qun th ngi c 300 ngi nhm mu MM, 180 MN v 420 NN. Hy
tnh tn s alen v xc nh xem qun th c trng thi cn bng
khng.
2. Vi cc s liu bi 1, gi s c s ngu phi gia cc c th, hy xc
nh tn s cc kiu nhm mu th h sau.
3. Trong mt qun th 200 ngi, 17 ngi c nhm mu A, 52 B, 3 AB v
128 ngi nhm mu O. Nu trng thi cn bng th tn s mi alen
l bao nhiu?
4. Trong mt qun th 300 ngi, c 69 ngi nhm mu MM, 108 MN v
123 NN. Hy tnh h s ni phi.
5. Tnh tn s alen, nu
a. S ng hp t ln gp i s d hp t.
b. S ng hp t ln gp su ln s d hp t.
6. C khong 1/20000 ngi sinh ra mc bnh x nang. Gi s rng bnh
ny do gen ln trn nhim sc th thng quy nh, d rng nhng
nghin cu gn y cho thy c nhiu kiu di truyn khc nhau ca
bnh ny.
a. Tnh tn s alen bnh x nang.
b. T l phn trm ngi mang bnh trong qun th.
7. Bnh m mu xanh - l bnh do gen ln lin kt X quy nh. T l
mc bnh l 64/10000 ph n.
a. Tnh tn s len ln.
b. T l ph n l th mang bnh?
123

8. Mt bnh do gen ln lin kt X c tm thy vi t l 1% nam gii.

a. Tnh tn s alen ln.
b. T l ph n mang gen bnh?
9. Cho p = 0,8, q = 0,2 v f = 0,4, hy tnh tn s cc kiu gen th h
sau.
10. Trong mt qun th ngi ln v ngu phi, tn s alen IA, IB v IO l
0,6, 0,3 v 0,1. Hy tnh tn s mi loi nhm mu.
11. Nu mt gen trn nhim sc th thng c hai alen vi tn s p =
0,8 v q = 0,2; tn s d hp t l 0,2. Hy tnh h s ni phi.
12. Bng ton hc bn hy bo v hoc phn bc li khng nh di
y: u, ht mu vng l do alen tri quy nh; tnh trng ln l
mu xanh; vi thi gian, tn s cy u c ht vng tng cho n khi
3/4 s c th c ht vng.
13. Mu sc v c sn chu u do mt gen c ba alen kim sot: CB
(nu), CP (hng) v CV (vng). Alen nu tri so vi hai alen kia; alen
hng tri so vi alen vng. iu tra mt qun th c sn, ngi ta thu c cc s liu sau:
Nu:

472

Hng:

462

Vng:

66

Tng cng: 1000

Hy tnh tn s alen khi qun th trng thi cn bng.
14. Trong mt qun th ngi c kch thc rt ln, tn s cc nhm mu
MN nh sau: 0,486 MM, 0,332 MN, 0,182 NN.
a. Tnh tn s alen.
b. Xc nh qun th c trng thi cn bng khng?
c. Nu qun th khng trng thi cn bng, hy tnh h s ni phi.

124

Chng 14
Tn s alen do t bin, di nhp v chn lc
A. t bin t lm thay i tn s alen trn mt th h.
Tc t bin ca mt gen thng vo khong 10-4 n 10-6/th
h; mt s t bin cn c tc nh hn nhiu.
V d: Xt mt qun th m cc c th khi u u c kiu gen AA.
Nu - tc t bin thun A a l 10-5. Tnh tn s alen A sau mt
th h.
Khi u p = 1. Tn s ca A s gim 0,00001 sau mt th h. V
khi u q = 0 v s khng c hi bin. Vy p = 1 - 0,00001 = 0,99999.
B. Vi mt gen c hai alen, s thay i tn s alen ph thuc c
vo tc t bin thun , v tc t bin nghch : p =
q - p v q = p - q.
Tc t bin thun Aa () lm gim s lng alen A, cn t bin
nghch aA () lm tng s alen A.
V d: Xt mt qun th c p = 0,8; q = 0,2. Nu = 5 x 10-5 v = 2 x
10-3, hy tnh tn s alen sau mt th h.
125

p = q -p = (2 x 10-5) - (2 x

) - (5 x 10-5) (8 x 10-1)

10-1

= (4 x 10-6) - (40 x 10-6) = -3,6 x 10-5

q = p -q = (5 x 10-5) ( 8 x 10-1) - (2 x 10-5) (2 x 10-1)

= (40 x 10-6) - (4 x 10-6) = 3,6 x 10-5
Vy p1 = 0,8 - 0,000036 v q1 = 0,2 + 0,000036
C. Nu ch tnh n tc ng ca t bin, th tn s alen
trng thi cn bng q = /( + ) v p = /( + ).
V d: Tnh tn s alen trng thi cn bng vi cc s liu mc B.
q = /( + ) = 5 x 10-5/[(5 x 10-5) + (2 x 10-5)] = 5 x 10-5/7 x 10-5 = 0,71
p = /( + ) = 2 x 10-5/[(5 x 10-5) + ((2 x 10-5)] = 2 x 10-5/7 x 10-5 = 0,29
Ch rng tn s trng thi cn bng khng ph thuc vo tn s
alen ban u. Cng cn ch rng v d ny l tnh hung l tng v
trong thc t khng ch c t bin tc ng n tn s alen.

D. Tn s alen ln sau cuc di nhp c c tnh bng: tn s ban

u ca qun th gc - (kch thc nhm di nhp c X hiu s tn
s alen gia qun th ban u v nhm di nhp c).
Nu qun th di nhp c tn s cao hn qun th gc, q s tng. Nu
qun th di nhp c tn s thp hn qun th gc, q s gim. Bng ton
hc, c th biu din: q = q - m(q - qm), vi m = kch thc nhm di nhp.
V d: Trong mt qun th gm 900 con bm, tn s alen quy nh cu
t chuyn ng nhanh ca mt enzym (p) bng 0,7, v alen quy nh
cu t chuyn ng chm (q) l 0,3. 90 con bm t qun th ny di nhp
n mt qun th c q = 0,8. Tnh tn s alen ca qun th mi .
Trc ht chng ta tnh m = 90/900 = 0,1. V q = 0,8 v qm = 0,3, nn
q = 0,8 - 0,1(0,8 - 0,3) = 0,8 - 0,05 = 0,75
V p + q = 1, nn p = 1 - 0,75 = 0,25.
E. Trong mt qun th gm cc c th gc v cc c th di nhp,
t l phn trm alen do cc c th di nhp ng gp bng:
[tn s alen gc (q) - tn s alen hn hp (q)]
[tn s alen gc (q) - tn s alen nhm di nhp (qm)]

126

v d: Trong mt qun th ngi, tn s alen M, (p), l 0,75 v tn s alen

N, (q), l 0,25. Trong nhm di nhp, tn s N l 0,4. Qun th hn hp c
q = 0,32. Tnh tn s alen N trong qun th hn hp.
Theo cng thc
m = (q - q)/(q - qm)
= (0,25 - 0,32)/(0,25 - 0,4) = -0,07/ -0,15 = 0,47 hoc 47%.
F. Nu cc kiu gen khc nhau c ph hp khc nhau th tn s
kiu gen sau chn lc s l: ph hp nhn vi tn s ban u.
V d: Trong mt qun th, tn s kiu gen AA = 0,25, Aa = 0,5 v aa =
0,25. Nu ph hp tng ng ca cc kiu gen ny l 1, 0,8 v 0,5, hy
tnh tn s cc kiu gen v tn s alen sau mt th h chn lc.
Trc ht ta tnh tn s cc kiu gen sau chn lc:
AA = (1) (0,25) = 0,25
Aa = (0,8) (0,5) = 0,40
Aa = (0,5) (0,25) = 0,125
Lu rng tng cc tn s ny khng bng 1, v mt s c th b
chn lc o thi. By gi, tn s cc kiu gen c tnh bng tn s mi
kiu gen/tng tn s:
AA = 0,25/0,775 = 0,322
Aa = 0,4/0,775 = 0,516
aa = 0,125/0,775 = 0,162
Ta tnh tn s alen = tn s ng hp t + 1/2 tn s d hp t.
p(A) = 0,322 + 1/2(0,516) = 0,58
q(a) = 0,162 + 1/2(0,156) = 0,42
G. Tn s alen ln sau mt th h chn lc bng qs = q(1 - sq)/(1
- sq2), vi s = h s chn lc.
V d: Tn s alen quy nh cu t chuyn ng chm ca mt enzym
(q) khi u l 0,7. Nu h s chn lc l 0,6, tnh tn s alen sau mt
th h chn lc.
qs = q( -sp)/(1 - sq2) = (0,7)[(1 - 0,6)]/[1 - (0,6)(0,7)2]
= (0,7 - 0,42)/(1 - 0,294) = 0,28/0,706 = 0,396
H. Nu kiu gen ng hp t ln gy cht th tn s alen ln
sau mt th h chn lc bng q/(1 + q).
V d: Trong mt qun th, mt alen ln khi u l trung tnh v c
tn s l 0,3. Mi trng sng bin i lm cho cc kiu gen ng hp t
ln cht hon ton. Tnh tn s alen sau 1 v 2 th h chn lc.
q1 = q/(1 + q) =0,3/1,3 = 0,230
127

 tnh cho th h th hai, chng ta phi bt u vi tn s alen mi:

q2 = 0,23/1,23 = 0,187
Phi ch rng s cn rt nhiu th h loi b mt alen. N s cn
tn ti trng thi d hp t.
I. Nu d hp t c u th chn lc so vi cc ng hp t, th tn
s1
s alen ln trng thi cn bng (q^) s l:
, vi s1 = h
s1 + s 2
s chn lc cho ng hp t tri, v s2 = h s chn lc cho
ng hp t ln.
V d: Nhng ngi c kiu gen d hp t v hemoglobin hnh lim c u
th chn lc nhng vng st rt lu hnh. Nhng ngi mc bnh hng
cu hnh lim thng cht. Nu h s chn lc ca cc ng hp t c
kiu hnh bnh thng l 0,2, hy tnh tn s alen hng cu hnh lim
trng thi cn bng.
Cho s2 = h s chn lc ca Hbs/Hbs v s1 = h s chn lc ca
HbA/HbA. V cc c th hng cu hnh lim cht, nn s 2 = 1. Vy q^ =
0,2/(1 + 0,2) = 1,7.
Ch rng p^ = s2/(s1 + s2) = 1/1,2 = 0,83.
J. Nu c t bin v chn lc cng tc ng th tn s alen ln
trng thi cn bng sau mt th h chn lc l khong
/s
v ca alen tri l /s.
V d: ng, tc t bin ca alen quy nh ng bnh thng
thnh alen ln l 2,5 x 10-6 v h s chn lc l 0,1. Tnh tn s alen ln
khi qun th trng thi cn bng.
q^ = 2,5 x 10-6/10-1 = 0,005
Mt ln na, cn lu rng q^ khng ph thuc vo tn s alen ban
u.
Tm tt
1. Tn s alen trng thi cn bng ph thuc vo tn s t bin thun
v nghch, v h s chn lc.
2. Qun th di nhp c tn s alen cao hn qun th n nh s lm
tng tn s alen qun th n nh; c tn s alen thp hn s lm
gim tn s alen qun th n nh.
3. Tn s alen ln sau mt th h chn lc bng qs = q(1 - sq)/(1 - sq2).
4. Nu ng hp t ln gy cht, th tn s alen ln sau mt th h
chn lc bng q/(1 + q).
128

5. Cc d hp t s c duy tr trong qun th nu chng c u th chn

lc so vi c hai ng hp t.

Bi tp
1. Mt ng nghim nui cy E. coli cha 1011 t bo, tt c u nhy vi
phag T1. Nu tc t bin thnh khng l 3 x 10 -8 th s c bao
nhiu t bo khng sau mt th h.
2. Trong mt qun th Drosophila ln, tn s alen ebony (e - mt nu g
mun) l 0,32. Nu tc t bin e+ e l 2 x 10-5 v e e+ l 106, hy tnh:
a. Tn s alen sau mt th h.
b b. Tn s alen khi qun th trng thi cn bng.
c. Tn s alen cn bng nu alen ebony c ph hp l 0,9.
3. Nu tc t bin A a ln gp nm ln a A, hy tnh tn s alen
a trng thi cn bng. V tnh a, nu (A a)/(a A) = 10; (A a)/
(a A) = 2.
4. Tn s alen quy nh nhm mu Duffy ngi Caucasia l 0,492. Tn
s alen ny qun th California da en l 0,094. Alen ny khng
tm thy nhng ngi Phi da en. Tnh t l alen ny nhng ngi
M da en c ngun gc t nhng ngi Caucasia.
5. Trong mt qun th ngi, 16% s ngi c mt xanh. 20% s ngi ca
qun th di nhp n mt qun th m ch 9% s ngi c mt xanh.
Nu gi s rng mt xanh c di truyn nh mt tnh trng ln trn
nhim sc th thng, hy tnh tn s alen mt xanh qun th mi.
6. Cc kiu gen AA, Aa v aa c tn s bng nhau trong qun th. Nu
ph hp tng ng ca cc kiu gen l 1,0, 0,8 v 0,3 v khng c
t bin. Hy tnh tn s cc kiu gen v tn s alen cho th h sau.
7. Hai alen transferrin TfA v TfB c tm thy chim b cu. Trng ca
nhng con chim ci d hp t khng khun hn trng ca c hai loi
chim ci ng hp t. Nu h s chn lc ca trng c kiu gen TfA/TfA
l 0,31 v c kiu gen TfB/TfB l 0,22, hy tnh tn s alen trng thi
cn bng.
8. Tn s alen ln b l 0,2 v h s chn lc l 0,8. Hy tnh tn s alen
sau 1 v 2 th h.
9. Nu tn s alen d l 0,25 trong qun th di nhp v 0,5 qun th
hn hp, v nu tc di nhp l 0,1. Tnh tn s alen d qun th
ban u.
129

10. Nu tc t bin t chut lang thnh khng lang l 3 x 10 -5 v

h s chn lc ca alen khng lang l 0,2, hy tnh tn s alen trng
thi cn bng.
11. Chng ci cc ngi do alen tri trn nhim sc th thng quy
nh. c tnh tn s t bin t bnh thng thnh ci cc l 4,5 x 105
v ph hp ca alen ci cc l 0,25. Tnh tn s alen ci cc
trng thi cn bng.
12. Nu s = 0,8 v q = 0,4, hy tnh tn s cc kiu gen B1B1, B1B2 v
B2B2.
13. Warfarin git cht th do n lm cho mu th khng ng c; rt
may, th xut hin tnh khng li n nh t bin nhy thnh
khng. Khi c mt warfarin, ph hp cho cc kiu gen ss, rs v rr l
0,68, 1 v 0,37. Hy tnh cc tn s trng thi cn bng sau khi s
dng lu di warfarin.
14. Tn s cc c th b tc ng bi mt alen tri, him, trn nhim
sc th thng l 4 x 10-6. Nu 70% s c th chu tc ng ca alen
ny cht trc khi sinh sn, hy tnh tc t bin t alen bnh thng thnh t bin.
15.

Trong mt qun th tn s cc kiu gen nh sau:

AA
0,984064

Aa
0,015872

aa
0,000064

Nu tc t bin A a = 5 x 10-6 v tc t bin ngc bng

0, hy tnh h s chn lc cho a.
16. Nu q = 0,01 v cc ng hp t ln cht trong d con, hy tnh
tn s alen sau 1 v 2 th h.

130

Chng 15
Nhng bI tp kh
1. Hai gen ln quy nh mu son (v) v nu (br) rui gim
Drosophila cng nh hng n mu mt. Nhng rui t bin c
hai gen ny u c mt mu trng. Cho rui ci mt son thun
chng lai vi rui c mt nu thun chng. F1 cho t phi c F2.
Hy xc nh t l kiu hnh F2 trong nhng trng hp sau:
a. Gi s gen mt son lin kt X v gen mt nu trn nhim sc
th thng.
b. Gi s c hai gen u trn nhim sc th thng v khng lin kt.
c. Gi s hai gen u trn nhim sc th thng v lin kt bi 20
n v bn .
d. Gi s hai gen u trn nhim sc th thng v lin kt bi 60
n v bn .
2. Drosophila Mu mt trng (Xw) l gen ln lin kt X; mu c
(cn) v nu (se) l nhng tnh trng ln do gen trn nhim sc
th thng quy nh. Nhng con ci ng hp t v nhng con c l
bn hp t v gen mt trng s c mu mt trng, bt k nhng alen
khc cng c mt trong kiu gen. Thm vo , cc ng hp t v
gen mt c v mt nu cng u biu hin mt trng. Ngi ta thc hin cc php lai sau y, ri cho F1 t phi c F2.
Php lai

B m

ci mt trng X c mt nu

ci mt nu X c mt c

ci mt c X c mt trng

ci mt trng X c mt trng

Cc t l phn ly F2 thu c nh bng di y. Hy ghp cho ng cc

t l vi cc php lai trn.
A

C
Ci

4 trng

3 c

3 nu

1 nu

D
c

3 nu

17 trng
9

3 c
1 trng

3 c
131

1 trng
3.

1 trng

3 nu

Ch khng lng Mexico c rt t lng. Khi cho lai ch khng lng vi

nhau, thu c 2/3 s con khng lng v 1/3 c lng thng. Khi cho i
con khng lng ni phi, ngi ta cng thu c kt qu tng t. Khi cho
i con c lng thng ni phi, thu c cc ch con c lng thng.
Gi s rng khi cho lai mt ch con khng lng vi mt ch lng qun
thun chng ta thu c s ch con khng lng v lng qun bng
nhau. Hy tin on tn s kiu hnh t cc php lai sau:
a. ch lng qun X ch lng qun
b. ch lng qun X ch lng thng

4. Loi Anser alchemister c c Hong gia s dng nhiu nm v n c

kh nng trng vng khi nui chng bng cc kim loi khc nhau.
Mt chng trnh nhn ging c thc hin v tch c nhiu
t bin. Gen ln hg c kh nng hnh thnh trng c v bc thu
ngn (trng vng) v lm cho g mc bnh i ngoi ra thu ngn.
Gen ln th hai sn lm cho trng c vt thic v g b bnh nt ng
trng. Cui cng, gen tri Fe lm cho trng c v ph r st v b nam
chm ht. Fe gy cht trng thi ng hp t; nhng phi nh vy
b non trc khi hnh thnh v. Cc t bin khc nhau c ch con
ng trao i nh di y:
sn

hg

Fe

ch thic thu ngn st vng

Cho hai c th d hp t v c ba gen trn lai vi nhau.
a. T l i con sng st c kh nng trng vng?
b. T l i con sng st c kh nng trng thic?
5.

Bn th t bin Drosophila c tn l jaunty wing (cnh vy) (j),

faty body (thn bo) (ft), balloon wing (cnh trn) (ba) v clot eye
(mt dnh) (cl). S dng s liu t cc php lai di y xy
dng bn di truyn thch hp:
Php lai A:
Rui ci cnh vy, thn bo thun X rui c cnh trn, mt dnh
thun

F1

ton b kiu di
132

Php lai B:
Rui c F1 t php lai A X rui ci cnh trn, vy, thn bo, mt
dnh

253 rui c cnh vy, thn bo

245 rui ci cnh vy, thn bo
243 rui c cnh trn, mt dnh
259 rui ci cnh trn, mt dnh

Php lai C:
Rui ci F1 t php lai A X rui c cnh trn, vy, thn bo, mt
dnh

cnh vy, thn bo

160

cnh trn, mt dnh

162

cnh trn, vy, thn bo

158

mt dnh

165

thn bo, mt dnh

cnh vy

thn bo, cnh trn, mt dnh

cnh trn, vy

thn bo

77

thn bo, cnh trn

78

cnh vy, mt dnh

75

cnh trn, vy, mt dnh

80

cnh vy, thn bo, mt dnh

kiu di

cnh trn

cnh trn, vy, thn bo, mt dnh

tng cng
6.

1000

Di y l mt phn bn nhim sc th X ca Drosophila:

mk

scr
133

1,0

6,0

22

mk = thn xm
l

= gy cht

scr = ngoi hnh khng gn

Alen gy cht l c duy tr rui ci d hp t v nhm sc th X
nh du bng mt on c bit c cha t bin tri Bar (B) v
cc o on bao ph ton b nhim sc th. nhng con rui ci
ny, mt nhim sc th X c cha alen ca t bin B v cc o
on; cc gen khc trn nhim sc th ny u l kiu di. Nhim
sc th X kia c s xp xp cc gen bnh thng, ging nh rui kiu
di, tr gen l. Rui ci nh vy c cho lai vi rui c thn xm v c
ngoi hnh khng gn. Nhng con rui ci con khng biu hin t
bin Bar c gi li. Nhng rui ci c lai vi rui c kiu di v
phn tch i con ca chng. Hy tnh ton cc tn s; gi s rng h
s trng lp vng gen ny l 0,5.
7. Tt cm ic c theo di hai gia nh ngi aixlen c ph h
sau:
Gia nh A

Gia nh B

a. Xc nh kiu di truyn ca tnh trng ny gia nh A.

b. Cng nh vy i vi gia nh B.
c. Tt cm ic hai gia nh ny l do mt hay nhiu gen quy
nh? Hy gii thch.
8.

Chng vi khun lys+his+val+ c dng lm th cho, cn chng

lys- his- val- l th nhn. Cc th bin np u tin c tch ra trn mi
trng ti thiu c cha his v val.
a. Nhng kiu gen no s sinh trng trn mi trng ny?
134

b. Cho cc khun lc ny sinh sn trn mi trng ti thiu + his, 75%

cc khun lc ban u sinh trng c. Nhng kiu gen no s sinh
trng c trn mi trng ny?
c. Cho cc khun lc ny sinh sn trn mi trng ti thiu + val, 6% s
khun lc sinh trng c. Nhng kiu gen no sinh trng c trn
mi trng ny?
d. Cui cng, cho cc khun lc ny sinh sn trn mi trng ti thiu.
Khng khun lc no sinh trng c. Da vo thng tin ny, xc
nh xem kiu gen no sinh trng c trn mi trng ti thiu +
his? trn mi trng ti thiu + val?
e. Da trn nhng thng tin trn, tm xem gen no nm gn gen lys.
f. Th nghim bin np c lp li, song a nui cy c cha mi trng ti thiu + lys + his. 50% khun lc sng c. Cho cc khun lc
ny sinh sn trn cc mi trng khc nhau xc nh kiu gen
ca chng. Kt qu nh sau:
val-his+lys+

val+his-lys+

37

val+his+lys-

Da vo kt qu trn, hy xc nh trt t cc gen.

9.

Nhng chng E. coli no di y c th thu phn lactose v trong

nhng iu kin no?
a. i+o+p+z+/i+p+o-z+
b. i-p-o+z+/i+p+o+zc. i+p+o+z+/i+p-o-z+
d. i+p+o+z+/i+p+o-ze. i-p+o+z+/i+p-o-zf. i+p+o+z-/i+p-o-z-

10. Bn ang th nghim vi mt sinh vt ngoi nhp v pht hin ra

operon sn xut t l operon chu cm ng. Operon gm bn vng
theo trnh t sau:
QRST
Tuy nhin, v tr ca gen ch huy (operator) v gen khi ng
(promoter) cn cha xc nh c. Bn thy rng gen c ch nh v
xa operon ny. c bit, cn c hai gen khc nhau cho vic sn xut t.
Nhng mt on khc nhau ca operon ny c tch v lp bn
. V tr cc mt on nh sau ( ///// = mt on):
Q

T
135

Mt on
1

////////

////////////

//////////

///////////////////

Mt on 1 lm cho t c sn xut ra lin tc; mt on 2, 3, 4 lm cho

t khng c sn xut ra.
a. Vng no c th l operator?
b. Vng no c th l promoter?
c. Kiu gen lng bi mt phn di y c to ra v kh nng sn
xut t ca n c xc nh ( - = khng sn xut t; I = chu
cm ng).
Kiu gen

Kh nng sn xut t

Q+R-S+T+/Q-R+S+T+

Q+R+S+T-/Q+R+S-T+

Q+R+S-T+/Q+R-S+T+

Da vo nhng thng tin b sung ny, xc nh xem vng no

cha promoter?
11. Bng di y lit k hot ng enzym ca bn th t bin e, r, u
v y cng tc ng n -galactosidase E. coli. C mt t bin
gen z, mt t bin gen ch huy, mt t bin promoter v mt
gen i. Xc nh xem t bin no gen no. IPTG c dng lm
cht cm ng cho operon Lac.
Chng
Kiu di

Hot enzym
-IPTG

+IPTG

1000

t bin e

t bin r

t bin u

1000

1000

t bin y

1000

1000

Flacz+/kiu di

2000

/e

1000

/r

1000

/u

1000

2000
136

/y

Flacz-/kiu di

2000

1000

/e

/r

/u

1000

1000

/y

1000

12. Kt qu lai cc chng vi khun t bin mt on c trnh by di

y ( + = cc th ti t hp kiu di c khi phc li). Hy v bn
cc t bin. Khng ch gii hn bng cc loi bn thng thng.

2
3
4
5

13. t bin RII phag T4 s sinh trng v to ra nhng vt tan ln trn

chng vi khun B; cc t bin rII s khng sinh trng trn K12 - mt
th tim tan lambda. Cc php lai sau y c thc hin chng B.
i con ca cc php lai c lm long n nng 10 -7 v c s
dng ti nhim chng B. Di y l s lng cc vt tan thu c:
1x2

250

1x3

250

2x3

250

i con ca cc php lai trn cng cn c lm long n nng 10-4

v cy ln K12. S lng cc vt tan thu c nh sau:
1x2

50

1x3

25

2x3

75

Hy v bn cc t bin ny v ch r khong cch gia chng.

14. Trnh t baz di y l trnh t gn u 3 ca ARN -globin ca
ngi. Tip l trnh t axit amin gn u C ca protein bnh thng v ca ba th t bin: Olmstead, House v Max.
137

5AAUGCCCUGGCCCACACUUAUCACUAAGCUAGCUUGACUAUGUG 3
bnh thng : ala-leu-ala-his-thr-tyr-his-C
Olmstead:

ala-arg-ala-his-thr-tyr-his-C

House:

ala-leu-ala-his-thr-C

Max:

ala-leu-gly-pro-his-leu-ser-leu-ser-C

a. B ba stop no c s dng hemoglobin bnh thng?

b. Cho bit t bin xy ra mi th t bin.
15. Mt phn t ADN c tch chit v phn tch thnh phn baz gm:
18% T, 32% A, !0% C v 40% G.
a. Bn cht vt l ca ADN ny l g?
b. Bn thc hin th nghim bin tnh v thu c kt qu di y:

Hp th

Nhit
Ti sao y l kt qu khng mong i?
c. Hy gii thch kt qu ny?
Ph lc
Gii p bi tp
Chng 1: Li gii
1. a. Bnh ny do gen ln qui nh. Nu hai v chng c cng kiu hnh
(bnh thng) sinh con, m mt trong s con ca h b bnh th c hai
b m phi d hp t v cp gen ny. Khi h d hp t th kiu hnh
ca h phi th hin tnh trng tri [ Khi nim (KN) 1-B].
b. Xc sut a con th hai b bnh l 1/4. Qui c B = bnh thng, b = b bnh. Php lai s l Bb x Bb . 1/4 trong s con ci ca h
s l bb (KN 1-H).
138

2. Yy x yy.
Chng ta thy xut hin hai kiu hnh vi t l xp x 1:1 v
khng bit alen no l tri t php lai ny. Hai kiu hnh gi n
mt gen (KN 1-B) v t l phn li 1 : 1 gi n php lai ng hp t
x d hp t (KN 1-D).
3.
a. Tnh trng cnh ngn l tnh trng tri nhng gy cht trng thi
ng hp t.
b. 1/2 cnh ngn : 1/2 bnh thng.
c. Tt c bnh thng. Ss x ss (KN 1-A).
Chng ta thy hai kiu hnh vi t l phn li 2 : 1. V c hai con
rui c cng kiu hnh nn tnh trng cnh ngn phi l tnh trng tri
(KN 1-E). Rui b (m) cnh ngn phi d hp t v rui m (b) bnh
thng phi ng hp t v cp gen ln. Do vy:
Ss x ss

1/2 Ss : 1/2 ss (KN1-D)

4. Mu en l tnh trng tri, mu nu l tnh trng ln v c hai ch b
m u d hp t. i con c t l phn li xp x 3 : 1 v c hai
ch b m c cng kiu hnh (KN1-C). Do vy php lai n gin nht
s l: Bb x Bb.
5. Tt c AB; hoc 1/2 AB , 1/2 A; hoc 1/2 AB, 1/2 B;
hoc 1/4 A : 1/4 AB :1/4 B : 1/4 O.
Cc php lai c th l:
AA x BB
AA x BO
AO x BB
Hoc

AO x BO

Cc kiu gen khc nhau cho ra cng kiu hnh, mt kt qu ca trng

hp a alen (KN 1-G).
IAIA x IBIB sinh ra tt c IAIB (AB)
IAIO x IBIB sinh ra

1/2 IAIB (AB) 1/2 IBIO (B)

IAIA x IBIO sinh ra

1/2 IAIB (AB) 1/2 IAIO (A)

IAIO x IBIO sinh ra

1/4 IAIB (AB)

1/4 IAIO (A)
1/4 IBIO (B)
139

1/4 IOIO (O)

6. Ll x Ll . Trong L: ui di, l : ui ct.
Ta thy c ba loi kiu hnh vi t l phn li xp x 1 : 2 : 1. Mt trong
s cc kiu hnh (ui ngn) l tnh trng trung gian gia ui di v
ui ct. Nh vy, chng biu hin tnh tri khng hon ton (KN1-E).
7. Mt l tnh trng tri v y l php lai gia hai rui d hp t.
Chng ta thy c hai kiu hnh v t s phn li xp x 3 : 1. C hai b
m c cng mt kiu hnh (KN1-C).
8. Trng-1, trng-2 : Ww, trng-3 : WW, en-1 : ww
V chng ta thy c hai kiu hnh khc nhau i con trong php lai
trng-1 x trng-2, nn tnh trng mu trng phi trng thi d hp t
(Ww) v en phi ng hp t ln (ww) (KN1-B v 1-C). T s phn li
nhn c khng phi l 3 : 1, nhng nh rng i con xc sut biu
hin mu en l 1/4 (KN1-H).
V trng-1 x trng-3 sinh ra con u c mu trng. Trng-3 phi
ng hp t (WW) (KN1-A). Php lai th ba c th l php lai phn
tch v nhn c t s phn li 1:1 (KN1-D), kt qu ny cng c th
suy ra c t nhng thng tin bit.

9. 1/6.
Php lai th nht s l se+se x se+se. Tnh trng mt i con s l
se+se+ hoc se+se; 2/3 trong s chng c th s l se+se (KN1-C v 1-E).
php lai tr li sinh ra rui mt nu en th c hai rui b m phi
d hp t. Nu c hai b m u d hp t th 1/4 s rui con c mt
nu en (KN 1-C).
Do vy, 2/3 (d hp t) x 1/4 = 2/12 = 1/6
10. a. chestnut : CcCc; palamino : CcCw; cremello : CwCw
b. Cho giao phi chestnut vi cremello.
Chng ta nhn c ba kiu hnh t php lai gia cc c th c cng
kiu hnh v t l gia cc kiu hnh gn vi 1 : 2 : 1 gi n tnh
tri khng hon ton (KN1-E).
Nu chng ta gi C = gen qui nh mu sc, Cc = chestnut v Cw =
cremello, CcCw = phi l palamino.

Php lai s l:
Cc Cw x Cc Cw

1/4 CcCc : 1/2 CcCw: 1/4 CwCw

140

Php lai gia palamino vi mt ng hp t no cng s sinh ra mt

na palamino (KN1-D). Do vy nhn c tt c palamino th lai
chestnut (CcCc) vi cremello (CwCw).
11. Xm : Cch, chinchila cch cch ; xm sng cchch hoc cchc ; bch tng: cc
Trc tin, ta lit k cc kiu gen v kiu hnh c th:
Xm : CC, Ccch, Cch, Cc
Chinchilla: cch cch
Xm sng : cchch v cchc
Himalayan : chch , chc
Bch tng : cc
nhn c th Himalayan trong php lai th ba, mt trong hai b m
phi c alen ch v phi c mu xm. Do vy th b m xm l Cch ,
chinchilla phi l cch cch , v php lai th nht phi l:
Cch x cch cch

1/2 Ccch (xm) : 1/2 cchch (xm sng) (KN1-D v 1-G)

Trong php lai th hai, hy th vi kiu gen cho mu xm sng l cchc.
Php lai s l:
Cch x cchc

1/4 Ccch (xm) : 1/4 Cc (xm) : 1/4 cchch (xm sng) : 1/4 chc
(Himalayan)
Cn kh nng khc cng cho mu xm sng, l kiu gen cchch , cng
sinh ra i con c cng t l phn li kiu hnh, nhng trong trng hp
ny th Himalayan con c kiu gen l chch .
12. a. Tc vng: bb
nu sng: bbmb
nu: bmbbmb hoc Bb
nu xm: Bbmb
en: BB
Hai php lai u cho thy mu vng v en l ng hp t. Gi s cc
kiu gen ny l bb v BB (tng ng vi mu vng v mu en). Php
lai th ba v th t cho ra nhiu kt qu, nhng u l kt qu ca di
truyn n gen. Cc kt qu cng chng t rng chng ta ang
gp trng hp di truyn a alen v chng ta c thm t nht l mt
alen th ba. Ch rng mt s php lai gia nu x nu sinh ra tt c

141

nu. Kt qu ny cho thy mu nu c th cng l ng hp t, hy

gi kiu gen ny l bmbbmb.
Xt php lai th ba. Chng ta gi s rng tc vng l ng hp t v
tc nu c th cng nh vy. Nu php lai l bb x bmbbmb th tt c
i con s l bbmb. c tc mu nu sng, phi c tnh tri khng
hon ton gia tc nu v tc vng. Hy xem kt qu th hai c th c
trong php lai ny. Ta thy mt na s con c tc vng m tc vng c
kiu gen ng hp t, v cho 1/2 s con c tc vng th mi b m
phi cho mt alen b. Do vy ngi c tc nu trong trng hp ny phi
d hp t v alen b. Chng ta cng bit rng kiu gen khng th l
bbmb v kiu gen ny cho mu nu sng. Chng ta xut c ba
alen: B, b v bmb. By gi chng ta s s dng alen th ba biu din
th d hp t tc nu.
Vy, tc nu c th l: Bb hoc bmbbmb.
Ch cn li mt kiu gen ca mu nu xm. N phi l kt qu ca
tnh tri khng hon ton gia mu en v mu nu. Do vy mu
nu xm l Bbmb .
V chng ta thy nhiu t s phn li t cng mt kiu hnh, chng ta
phi cp ti trng hp a alen (KN1-G). Chng ta cng thy mt s
kiu hnh trung gian, di truyn theo mt trt t tri ln nht nh
(KN1-F).
b. 1/4 nu sng
1/4 nu xm
1/2 nu
S dng cc kiu gen phn (a) chng ta c:
Bbmb x bbmb

1/4 bbmb (nu sng)

1/4 Bb

(nu)

1/4 bmbbmb (nu)

1/4 Bbmb (nu xm)
d. 1/2 nu : 1/2 nu xm
bbmb x BB

1/2 Bb (nu) : 1/2 Bbmb (nu xm)

13. Mu xanh nht l kt qu ca tnh tri khng hon ton gia alen
xanh m v mt vi alen khc, alen th hai ny l alen gy cht khi
142

trng thi ng hp t. Kt qu php lai th nht cho thy l php

lai d hp t x ng hp t (KN1-D). V mu xanh nht chim s nhiu
trong php lai th hai, nn c l n l th d hp t. Kt qu ny cp
n tnh tri khng hon ton (KN1-A, 1-B v 1-E). i con trong php
lai th hai c t l phn li gn vi 2:1, gi ti mt gen gy cht (KN1F). Alen qui nh mu xanh nht phi l alen gy cht khi trng thi
ng hp t.
Qui c: GDGD : xanh m
GDGL : xanh nht
Php lai th hai c th vit nh sau:
GDGL x GDGL

1/4 GDGD : 1/2 GDGL : 1/4 GLGL

xanh m

xanh nht

cht

14. a. C
b. Khng
Nhng ngi c nhm mu A c th c kiu gen IAIA hoc IAIO. Nu c
hai ngi c kiu gen IAIO kt hn th 1/4 s con s c kiu gen I OIO hoc
nhm mu O (KN 1-B, 1-D v 1-G).
15.-1 : CACB; -2 : CACC; 3: CACA; trng: CCCC; hng : CBCC.
Kiu gen CBCB : gy cht.
Xt php lai 1 v 3. Cc c th c kiu hnh nh nhau li sinh ra con c
kiu hnh khc nhau. Do vy, y c th l trng hp di truyn a
alen (KN1-G).
Php lai 4 v 5 cho thy mu tri hn mu hng v mu trng v
c th -3 phi l ng hp t (KN1-A). Gi s rng chng ta c ba
alen CA , CB v CC.
Cho CA = th cc c th -1 v -2 phi l d hp t nhng cc
alen khc nhau. Hy gi kiu gen CACB : -1 v CACC : -2.
Cc t s phn li trong cc php lai vi mu hng cho thy mu hng
cng phi l d hp t (KN1-C, 1-D v 1-E). Mu trng phi l ng hp
t (KN1-D). Hy gi CCCC = trng , do vy CBCC = hng. T s phn li
2:1 trong php lai cho thy c mt gen gy cht (KN1-F), nhng kiu
gen no s b cht?
S php lai:
CACB x CBCC

1/4 CACB :1/4 CACC :1/4 CBCC :1/4 CBCB

143

hng

cht

S dng cc kiu gen CACC x CBCC cho php lai th ba cng cho ra cc
t l phn li ng nh t l quan st. Do vy chng ta c ba alen CA ,
CB v CC, kiu gen CBCB : cht; CBCC : hng v CCCC: trng.

Chng 2: Li gii
1. v di l tnh trng tri.
rrLl x rrLl
RrLl x Rrll
R-ll x RRLl
RrLl x RrLl
Rrll x rrLl
Trong R = , r = nu, L = di, l = ngn.
Trc ht hy xt cc php lai ch cho ra hai kiu hnh, nhng php lai
chng t rng c mt gen trng thi ng hp t. Php lai 1 v 3 l
nhng php lai nh vy. php lai 1 ta ch thy mt nu v php
lai 3 ch thy mt . Gen qui nh v mu mt phi l ng hp t
t nht l mt trong hai b m. Qua hai php lai ny chng ta khng
th xc nh c mu hay mu nu l tnh trng tri. php lai 1,
c hai b m u c cnh di v c t l phn li 3:1 i con. Do
vy, cnh di phi l tnh trng tri v php lai ny phi l Ll x Ll. L
lun tng t cho php lai 3 (KN 2-B).
php lai 2, chng ta thy t l phn li xp x 3:3:1:1, cho thy mt
gen l d hp t c b v m. Chng ta thy t l phn li 3 : 1
nu. Do vy mu l tnh trng tri v d hp t c hai b m: Rr x
Rr. Vy t s phn li 3:3:1:1 cho thy php lai l: AaBb x Aabb (KN2D).
php lai 4, ta thy c 4 kiu hnh vi t l phn li xp x 9:3:3:1.
y l kt qu ca php lai gia hai cp gen d hp t RrLl x RrLl
(KN2-C). php lai 5, chng ta thy mt t s phn li 1:1:1:1, kt qu
ca php lai phn tch hai cp tnh trng (KN2-E). V chng ta xc
nh c mu v cnh di l tnh trng tri, nn cc kiu hnh ny
l cc th d hp t. Mu nu v cnh ngn l cc th ng hp t
ln.
2. dydy unc+unc x dy+dy uncunc.
144

Qui c dy = ngn, dy+ = di, unc+ = mp, unc = thon.

Chng ta thy c 4 kiu hnh, cho nn c th c hai gen (KN2-A). Bn
nhm kiu hnh vi s lng gn bng nhau, cho thy mt gen trng
thi d hp t c hai b mv gen kia l ng hp t. Chng ta khng
th ni rng alen no l tri, nhng t l phn li 1:1:1:1 gi ti kt qu
ca php lai phn tch hai tnh trng (KN2-E).
3. ta l tnh trng tri i vi trng, t nht c ba gen v c tng tc
gen. Kiu hnh F1 chng t mu ta l tri i vi tt c cc mu
khc.Chng ta thy xut hin nm kiu hnh F 2, vy phi c t nht 3
gen. Nh rng, nu chng ta ch c hai gen v tnh tri hon ton,
chng ta ch c bn kiu hnh (KN2-A v 2-F). Thng thng vi ba gen,
chng ta c th c tm kiu hnh. y chng ta ch c nm kiu
hnh, nh vy phi xy ra tng tc gen (KN2-J).
4. 7/16.
Hai ngi c cng kiu hnh kt hn sinh ra mt kiu hnh mi. Kt qu
ny cho thy c hai gen (KN2-J), mi mt gen c th sinh ra mt kiu
hnh t bin. Nghe c bnh thng phi l tnh trng tri i vi
ic v s php lai l:
AAbb x aaBB
ic ic
tt c AaBb
(nghe c)
Php lai tip theo l:
AaBb x AaBb . Php lai ny sinh ra:
9/16 A-B- : nghe c
3/16 A-bb : ic
3/16 aaB- : ic
1/16 aabb : ic
5. 5 c lng: 1 khng c lng.
Trc tin vit s php lai: HhSs x HhSs.
S dng nguyn tc nhn xc sut (KN2-G) d on cc kiu gen.
1/4HH .............................cht
3/ 4 S- .........3/ 8 HhS- c lng
1/2Hh
1/4ss ...........1/8Hhss khng c lng
3/4S- ..........3/16 hhS- c lng
1/4hh
145

1/4ss ...........1/16 hhss c lng

Php lai cho mt t l 10/16 c lng: 2/16 khng c lng. Chng ta
ch c th m c nhng con rui cn sng, do vy ta c t l 5 c
lng : 1 khng lng. Chng ta s dng nguyn tc nhn xc sut
(KN2-G) v tip theo l sa i kt qu v c gen gy cht (KN2-H).
6. a. 9/64
b. 3/64
c. 3/64
Qui nh :A/a = bnh thng/bch tng, B/b = nu/xanh, D/d = bnh
thng/i tho ng, P/p = bnh thung/PKU
S php lai:
AaBbDdPp x AabbddPp
Tnh xc sut ca tng trng hp bng cch nhn cc xc sut ring
(KN2-G).
a. 3/4bnh thng x 1/2nu x 1/2bnh thng x 3/4bnh thng
= 9/64
b. 3/4bnh thng x 1/2xanh x 1/2i ng x 1/4PKU = 3/64
c. 1/4bch tng x 1/2xanh x 1/2 i ng x 3/4bnh thng =
3/64
7. a. 3 tri : 1 ln.
b.1 tri : 1 th t bin b : 2 th t bin a.
C hai t l phn li F1 u bt ngun t t l 9:3:3:1 (KN2-J). c
t l phn li 15:1 th t nht con mang mt alen tri phi biu hin
tnh trng tri; A-B-, A-bb, v aaBu biu hin kiu hnh tri.
Php lai cn tm l AaBb x aabb, l php lai phn tch hai tnh trng,
do vy chng ta c t l phn li 1:1:1:1 (KN2-E). Ta c:
1/4 A-B- bnh thng
1/4 A-bb bnh thng
1/4 aaB- bnh thng
1/4 aabb th t bin
c t l phn li 9:4:3 trong php lai b, mt alen t bin c ch
hot ng ca gen khc nh sau:
9 A-B- bnh thng
3 A-bb th t bin b
3 aaB- th t bin a
1 aabb th t bin a (KN2-J).
Php lai cn tm li l php lai phn tch hai tnh trng AaBb x aabb
146

Php lai to ra:

1/4 A-B- bnh thng
1/4 A-bb th t bin b
1/4 aaB- th t bin a
1/4 aabb th t bin a (KN2-E)
8. a. 9/32
b. 0
c. 9/32
d. 0
e. 1/8
tnh cho tng phn ta s dng nguyn tc nhn xc sut (KN2-G).
a.3/4 A- x 1/2 B- x 1 C- x 3/4 D- = 9/32
b. Tt c cc c th u phi c kiu gen Cc , khng c th no l cc.
c. 3/ 4A- x 1/2 bb x 1C- x 3/4 D- = 9/32
d. Tt c cc c th u phi c kiu gen Cc , khng c th no l cc.
e. 1/2 Aa x 1/2 Bb x 1 Cc x 1/2 Dd = 1/8
9. Mu en l tnh trng tri v c hai con ch u d hp t v gen ny.
hoc lng di hoc lng ngn l tnh trng tri v trng thi d hp
t, con ch kia l ng hp t ln. T s phn li st vi 3:3:1:1, gi
n php lai c dng sau y: AaBb x Aabb (KN2-A v 2-D). Mu en
phi l tnh trng tri khi m 3/4 s ch con c mu en. Chng ta
khng th kt lun c lng di hay lng ngn l tnh trng tri.
Php lai c th l DdLl x Ddll hoc Ddss x DdSs.
10. a. 1/8
b. 3/8
V tnh trng nhiu ngn l t gp, nn c th gi nh rng bt c
c th no b bnh u trng thi d hp t. C hai ngi con phi d
hp t v tnh trng li cong v b h bnh thng l ng hp t ln.
Hy qui c P/p = nhiu ngn/bnh thng v C/c = li cong/ li thng.
S php lai c th vit nh sau:
PpCc x ppCc
v s dng nguyn tc nhn xc sut (KN2-G) tnh:
a. 1/2 bnh thng x 1/4 thng

= 1/8

b. 1/2 nhiu ngn x 3/4 li cong = 3/8

147

11. a.

ta

AABBCC

Trng-1 aaBBCC
Trng-2 aabbCC
Trng-3 aabbcc
b. 1 ta : 1 : 2 trng
c. 4 trng : 1 : 1 nu : 1 xanh : 1 ta
V tt c F1 c mu ta, nn mu ta phi l tnh trng tri.
php lai 3 chng ta thy c 5 loi kiu hnh, do vy phi c t nht ba
gen (KN2-F) v Trng-3 th h P phi ng hp t ln v c ba gen.
Phi phn tch qua nhiu bc mi c th tr li c cc cu hi. Trc
ht phi xc nh c kiu di truyn ca mi mu mt. Xem xt
php lai vi trng-1. F2 c t l phn li xp x 3 : 1, cho thy ch c
mt gen l d hp t F1. Gi s gen A l gen ln trng-1. Vy kiu
gen ca trng-1 l aaBBCC (KN2-K).
trng-2, ta thy c ba kiu hnh vi t l phn li 9 : 4 : 3, cho thy
trng-2 l ng hp t ln v hai gen (KN2-J). Gi s aabbCC = trng2.

S ca php lai 2 c th l:
AABBCC x aabbCC

AaBbCC
( ta)

t phi
9A-B-CC ta
3 A-bbCC
3 aaB-CC trng
1 aabbCC trng
(KN2-C v 2-J)
F1 ca php lai 3 phi d hp t v c 3 gen. S dng phng php v s
v nguyn tc nhn xc sut (KN 2-G) tnh cc tn s.
1/4 aa .................... 1/4 trng
3/4C- .... 27/64 A-B-C- ta
3/4B148

1/4cc......

9/64 A-B-cc

3/4C-.....

9/64 A-bbC-

xanh
3/4 A
1/4bb
1/4cc...... 3/64 A-bbcc
nu
C 128 nhn con v t l thu c gn st vi t l d on.
F1 ca php lai 2 l AaBbCC v php lai phn tch l AaBbCC x aabbcc.
Chng ta nhn bit kiu gen ny bng php lai phn tch v nhn c:
1/4 aaBbCC ta
1/4 AabbCC
1/4 aaBbCC trng
1/4 aabbCC trng
(KN2-E)
F1 ca php lai 3 l AaBbCc, php lai phn tch l AaBbCc x aabbcc,
php lai phn tch 3 cp gen, chng ta c 8 nhm bng nhau:
1/8 AaBbCc ta
1/8 Aabbcc nu

1/8 AabbCC
1/8 AaBbcc xanh

Bn nhm cn li u l aa v do vy u c mu trng.
12.a. AABB x aabb; A qui nh mu trng v aabb cng cho ra mu
trng.
b.Tt c trng; 3 trng: 1 .
F1 gi n mt php lai ch c 1 th ng hp t, nhng nu ng
nh vy th khng c mu g ngoi mu trng F2. Do vy chng ta
cho rng phi c 2 gen. Nu chng ta chia hai nhm cy cho 8, ta s
nhn c t s 13:3, mt bin dng ca t s 9:3:3:1 (KN 2-J). Mu
trng ca F1 phi l d hp t v c 2 gen AaBb. Mt php lai gia 2 d
hp t sinh ra:
9 A-B- trng
3 A-bb trng
3 aaB-
1 aabb trng
149

Hai b m u trng phi l AABB v aabb.

Nu nh F1 c lai tr li vi m: AABB x AaBb, chng ta nhn c tt c
trng A-B- . Nu lai vi b aabb x AaBb, chng ta nhn c 1/4 AaBb
(trng), 1/4 Aabb (trng), 1/4 aaBb (), 1/4 aabb (trng) hoc 3
trng: 1 .
13. Dng 1: aaBBcc
Dng 2: AABBcc
Dng 3: AAbbcc
Dng 4: aabbcc
Chng ta thy c 5 kiu hnh trong php lai 4, do vy c 3 gen phi d
hp t (KN2-F). Do vy dng bch tng 4 phi l aabbcc. php lai 2,
chng ta nhn c kt qu nh php lai hai c th d hp t v mt gen
(v d 3 :1). V tt c chut bch tng phi l cc, nn ch locus ny l
d hp t F1, do vy dng bch tng 2 l AABBcc. Php lai 1 v 3 sinh
ra chut con c t l xp x 9:4:3, chng t c 2 gen d hp v y l
t s bin i ca t s 9:3:3:1 (KN2-J). Cho A v C l d hp trong
php lai 1 v B, C trong php lai 3. Do vy, dng 1= aaBBcc v dng 3
= AAbbcc.
Php lai 1 sinh ra:

Php lai 3 sinh ra:

9 A-B-C- xm

9 A-B-C- xm

3 aaB-C- nu vng

3 A-bbC-

3 A-B-cc bch tng

3 A-B-cc

1 aaB-cc

1 A-bbcc

bch tng

en
bch tng
bch tng

Ch 1 kiu hnh cn li l mu scla. Kiu gen phi l aabbC-.

Hy xem xt cc t s v i t n gin n phc tp (KN2-K).
14. c v nu l tnh trng ln v tng tc cho ra mu trng. T l
gn vi 9:3:3:1, do vy chng ta c cc c th F 1 l cc th d hp t
kp (KN2-A,2-B v 2-C).
Hy gi s+ = kiu di, s = c; br+ = kiu di, br = nu. Php lai l:
s+s+brbr x ssbr+br+

Tt c s+sbr+br

T phi
9 s+-br+-
3 s+-brbr nu
3 ssbr+- c
150

1 ssbrbr trng
15. Ch ci vng: GgBb
Ch c vng: Ggbb
Ch c en: ggBb
Cc t s chng t c 2 gen (KN2-B) phn li theo t l bin i t t
l 3:3:1:1 (KN2-D v 2-J), gi n s tng tc gen. Trc ht hy phn
tch nhm ch theo mu vng v khng phi mu vng. Trong php lai
1, chng ta thy t l 3:1, chng t rng gen qui nh mu vng phi
d hp c 2 con ch. Hy qui c G- = vng v gg = khng phi mu
vng. Mu en v nu xut hin vi t l 1:1, chng t rng 1 d hp
t x 1 ng hp t ln (KN2-B), nhng chng ta khng th ni rng alen
no l tri trong php lai 1. php lai 2 chng ta thy t l phn li 3
en: 1 nu, chng t rng en l tnh trng tri v mi c th u d
hp v gen ny.
Hy qui nh B = en v bb = nu. Ch ci vng phi d hp v c 2
gen : GgBb. Ch c vng phi l: Ggbb. Ch c en phi l gg nu
khng n c th li l vng, do vy ch c en l ggBb.
16.12 : 4 c : 3 mn : 1 trng.
Lp php lai AaBbCc x AaBbCc v s dng nguyn tc nhn xc sut
tnh cc kh nng (KN2-G); Sau xc nh kiu hnh cho cc
kiu gen (KN2-H).

3/4 C- ...27/64 A-B-C-

3/4 B1/4 cc ...9/64 A-B-cc

3/4 A3/4 C- ...9/64 A-bbCmn

1/4 bb
151

1/4 cc ... 3/64 A-bbcc

cht
3/4 C- ...9/64 aaB-C-
c
3/4 B1/4 cc ...3/64 aaB-cc
c
1/4 aa
3/4 C- ...3/64 aabbCtrng
1/4 bb
1/4 cc ... 1/64 aabbcc
cht

Chng 3: Li gii
1. a. 39 cp.
152

b. 39
Mi nhim sc th c 2 bn copy v 2 x 39= 78. Gim phn lm gim
i mt na s nhim sc th v 1/2 x 78 =39 (KN3-A).
2. a. 64. V n = 6, 26 = 64 (KN 3-B).
b. 1/64
Xc sut 1 giao t nhn c A l 1/2 . Tng t, xc sut nhn
c B,C,D,E hoc F cng l 1/2. Xc sut nhn c tt c s bng
tch ca cc xc sut ring:
1/2 x 1/2 x 1/2 x 1/2 x 1/2 x 1/2 = 1/64
3. Khong 8 triu. Hai mi ba cp ngha l n = 23, v 223 = 8 x 106 (KN3B).
4. a. 2c
b. 2c
c. 4c
Cc t bo thi im cui ca gim phn I cha mt na s nhim
sc th, nhng trng thi kp. V ADN ca trng (sau gim phn II) l
do s phn chia ca nhim sc t, nn cc t bo cui ca gim
phn I phi c s lng ADN l 2c. Mt t bo tin hnh nguyn phn c
23 cp nhim sc th n. Nu 1 b nhim sc th n = c th 2
b = 2c. Nhim sc th c nhn i trc khi bt u gim phn;
do vy 2 x 2c = 4c (KN3-D).
5. f.
Cc nhim sc th tng ng s kt cp trong gim phn. Mi giao
t s nhn mt nhim sc th t mi cp A,B,C hoc D (KN3-A). S th
tinh kt hp 2 t bo vi b nhim sc th nh trn. Do t bo r l t
bo sinh dng nn n dng lng bi.
6. a. 1/8
b. 1/8
c. 3/8
S cc t hp nhim sc th l 2n, trong n = s nhim sc th
khc nhau (KN3-B). Bi vy c 2 3 hay l 8 t hp. Xc sut nhn c
mt nhim sc th trong mt cp l 1/2 ; do vy xc sut nhn
c c 3 nhim sc th ca b hoc ca m
= (1/2) 3= 1/ 8 .
Cc hp t c th mang 2 nhim sc th ca b v 1 nhim sc th
ca m l P'Q'R; P'QR' v PQ'R' (KN3-A). V c 8 t hp khc nhau nn
t l l 3/8 .
7. Giao t ca la m c 21 nhim sc th, giao t ca ng c 10 nhim
sc th. Thm ch nu 10 nhim sc th ca ng c th kt cp vi
10 nhim sc th ca la m, d iu rt kh xy ra, th vn cn
153

11 nhim sc th ca la m khng th kt cp, v nh vy, n c

th phn li mt cch ngu nhin trong gim phn. Mi giao t s nhn
c b nhim sc th khng hon chnh, v hu ht giao t khng c
kh nng sng st. Nu s th tinh thc hin c th hp t c th c
thm cc nhim sc th (th ba) hoc c th mt i mt s nhim
sc th (th mt, th khng) (KN3-C).
8. a. 7
b. 14
c. 21
nhn ca ng phn, mt trong cc nhn ca ht phn l do kt qu
ca gim phn, v vy n l n bi (KN3-A). T bo l l lng bi. Ni
nh l kt qu ca s th phn gia 1 nhn ht phn n bi v 1
nhn lng bi to ra mt t bo tam bi.
9. a. 240
b.120
c. 60
Tinh bo s cp l lng bi v s tri qua gim phn to ra 4 t bo,
do vy 4 x 60 = 240. Tinh bo th cp tri qua mt ln phn bo
gim nhim v s phi tri qua mt ln phn bo na nn 2 x 60 =
120. Tinh t l n bi v s trng thnh tr thnh tinh trng (KN3A).
10.a. 60
b. 60
Non bo s cp l lng bi v s tri qua gim phn, nhng ch mt
trng c to ra t mt non bo s cp (KN3-A). Non bo th cp s
phn chia to ra 1 trng v 1 th cc.
11.256 . S t hp l 2n, trong n = s nhim sc th khc nhau (KN3B).

154

Chng 4: Li gii
1. Ngi v chung thu. V cha ca ngi ph n ny b bnh mu kh
ng, ng ta phi cho con gi ca mnh mt nhim sc th X (KN4-B).
Ngi ph n ny bnh thng, chng t c ta phi trng thi d hp t
v gen ny. a con trai ca c ta b bnh, n phi mang mt alen
mang bnh. Alen mang bnh ny phi c ngun gc t m.
2. Ngi ph n tc thng kt hn vi ngi n ng tc qun. Do c s
khc bit v gii tnh, ngi ph n phi ng hp t v gen ny. Nu
c ta c kiu gen d hp t th 1/2 con trai s c tc qun v 1/2 s c
tc thng (KN4-D). Ngi ph n khng th l ng hp t v tnh trng
tri; v nu nh vy th c con trai, c con gi s nhn c nhim sc
th X mang gen tri v tt c s c tc qun. Nu ngi ph n ny
ng hp t v gen ln (thng) kt hn vi ngi n ng tc qun th
tt c con trai s nhn c nhim sc th mang gen tc thng v tt
c con gi s nhn c nhim sc th mang gen tc qun (KN4-E).
3. Gen qui nh tnh trng cnh di l tri i vi cnh ngn v nm trn
nhim sc th thng. Gen qui nh mt mu tri i vi mu
nu v lin kt vi gii tnh.
Ta qui c: + = cnh di, s = cnh ngn, XR = mt v Xr = mt nu
ss; XRXR x ++; XrY
Trc tin chng ta xem xt s phn li F 1 v thy rng mt , cnh
di phi l tnh trng tri. F2 c bn kiu hnh, c th cho rng c hai
gen. Nhng ch hai trong bn kiu hnh xut hin con , chng t
mt trong hai gen ny lin kt vi gii tnh (KN4-C).
Hy xem xt kiu gen ca m. Nu di cnh l tnh trng lin kt vi
gii tnh th tt c rui F1 s c cnh ngn. Gi nh ny khng ph
hp vi kt qu; Do vy gen qui nh di cnh phi nm trn nhim
sc th thng.
V tnh trng mu mt, rui F1 phi d hp t v tt c cc gen.
Mt na s rui F2 s nhn nhim sc th X bnh thng v mt na
s nhn nhim sc th t bin (KN4-D).Chng ta nhn c 47 rui
mt nu v 53 rui mt chng t rng tnh trng mu mt lin
kt vi gii tnh. Tnh trng cnh di c rui v c rui c t l
phn li xp x 3:1, t l phn li ca mt gen trn nhim sc th th155

ng. rui t l phn li chung l 3:3:1:1 , cho thy mt gen nm trn

nhim sc th thng v mt gen lin kt vi gii tnh (KN4-F).
4. a. Lng ngn l tnh trng lin kt vi gii tnh nhng l mt gen ln
gy cht. Chng ta thy c s khc bit c v kiu hnh c v s lng
gia rui v rui , iu cho thy gen lin kt vi gii tnh l gen
gy cht bn hp t (khng c rui lng ngn) (KN4-C v 4-G). V
gii ci c hai kiu hnh cho nn lng ngn phi l tnh trng tri v
php lai s l:
XSXs x XsY
b. i con tt c u c lng di v phn u c hai gii. c
rui lng di, rui m phi ng hp t v php lai s l:
XsXs x XsY
5. Mo d hp t v alen qui nh mu lng lin kt vi gii tnh. Chng
ta thy c s khc nhau v kiu hnh con v con chng t tnh
trng lin kt vi gii tnh (KN4-C). Mo phi d hp t v phi c nhim
sc th X bt hot (KN4-H).
Hy qui c Xb = mu en v Xo = mu vng. By gi php lai s l:
XbXo x XoY

XoXo

XbXo

XbY :

(vng) (en v vng) (en)

XoY
(vng)

6. Mu sc lng l tnh trng lin kt vi gii tnh v gii l gii ng

giao t. Chng ta thy s khc bit v kiu hnh gia gii v gii
cho thy c s lin kt vi gii tnh KN4-C). V tt c cc c th ca mi
gii ging nhau v kiu hnh nn b m khng th l d hp t (KN4-D
v phn lu ). Ta lp php lai theo cch thng thng:
XGXG x XgY
Trong trng hp ny th c chim trng v chim mi u c mu xanh,
v chim con l XGXg v chim con l XGY. Kt qu ny khng ph hp
vi kt qu thc tin. Do vy c th c sai lm khi chng ta cho
rng gii l gii ng giao t.
Nu cho rng gii l gii ng giao t th php lai by gi s l:
ZGW x ZgZg
(xanh)

(vng)

ZgW

ZGZg

(vng) (xanh)

156

7. Mu sc lng l tnh trng lin kt vi gii tnh, xanh l tnh trng tri
v gii c hai alen qui nh mu sc. Ch c php lai th ba cho bit
tnh trng ny lin kt vi gii tnh v c s khc nhau v kiu hnh
gia ging v ging (KN4-A, 4-B v 4-C). Php lai th hai cho bit
xanh l tnh trng tri i vi vi vng. Nhc li rng, loi chim, gii
l gii ng giao t (ZZ). Nu chng ta qui c G/g = xanh/vng , cc
php lai c th c vit theo s sau:
ZgZg x ZgW

ZGZG x ZgW

ZgZg x ZGW

ZgZg

ZgW

ZGZg

(tt c vng)

ZGW
(tt c xanh)

ZGZg

ZgW
(xanh)

(vng)
8. F1: tt c xm; tt c vng. F2: 1/2 xm, 1/2 vng c hai gii.
Chng ta thy c s khc nhau v kiu hnh trong cc gii F 2, chng
t tnh trng c lin kt vi gii tnh (KN4-C). F 1 cho thy mu xm l
tnh trng tri i vi vng. Rui F 1 ca php lai phi d hp t v hai
kiu hnh c rui F2 l kt qu ca s phn li nhim sc th X
rui F1. Hy t X+ = xm v Xy = vng.
Php lai s l:
X+X+ x XyY

X+Xy x X+Y
(xm)

(xm)

X+X+

X+Xy

X+Y

XyY

(xm) (xm) (xm) (vng)

Php lai cn tm l:
XyXy x X+Y

X+Xy XyXy

X+Y

XyY

(xm) (vng) (xm) (vng)

9. a. Da c vy l mt tnh trng lin kt vi gii tnh.
b. 1/2
Chng ta thy s biu hin bnh khc nhau gia con trai v con gi,
cho thy bnh do mt gen lin kt vi gii tnh qui nh (KN4-C). Da c
vy khng th lin kt vi nhim sc th Y, v nu nh vy th tt c
con trai u biu hin bnh (KN4-B). V tt c cc c con gi, v
157

khng c cu con trai no b bnh, cho nn da c vy phi l tnh trng

tri lin kt vi nhim sc th X (KN4-E).
Nu qui nh S/s = c vy/ bnh thng, th b m s l XsXs x XSY.
Tt c cc c con gi phi d hp t. Mt na s cc chu s nhn c
nhim sc th XS (KN4-D).
10.a. kiu di (thn , cnh di) , thn vng, cnh ct.
b. 3 kiu di: 3 thn vng, cnh di : 1 thn , cnh ct : 1 thn
vng, cnh ct c hai gii.
c. 3 kiu di : 1 thn , cnh ct; 3 kiu di : 3 thn vng,
cnh di : 1 thn , cnh ct :1 thn vng, cnh ct.
Hy qui c X+ = , Xy = vng, vg+ = di, vg = ct

XyXy, vg+vg+ x X+Y, vgvg

X+Xy, vg+vg
(kiu di)

XyY, vg+vg
(vng)

(KN4-B)

S dng phng php nhn xc sut tnh t l phn li ca F2 .

3/4 vg+ ............................. 3/8 X+-,vg+1/2 X+
1/4 vgvg ........................... 1/8 X+-,vgvg
3/4 vg+ ............................. 3/8 Xy- ,vg+
1/2 Xy1/4 vgvg............................. 1/8 Xy-, vgvg
(KN4-F)
Vi php lai nghch:
X+X+,vgvg x XyY,vg+vg+

X+Xy, vg+vg x X+Y, vg+vg

Tt c rui ci F2 s nhn X+ t F1 v gen trn nhim sc th thng
s cho t l phn li 3:1. Gii s nh v d trn (KN4-E v 4-F).
11. Rui m mang mt gen qui nh mt mu cam trn nhim sc th
thng v rui b cng mang mt gen qui nh mt mu cam lin kt
vi gii tnh. V rui F 1 biu hin kiu di (mt ), chng phi c hai
158

gen khc nhau (xem chng 2), v gen qui nh mu cam phi l gen
ln. F2 c s phn li khc nhau cc gii, do vy chng ta phi cp
ti tnh trng lin kt vi gii tnh (KN4-C). Nu c hai gen u lin kt
vi gii tnh, chng ta s c rui mt mu cam F1. Nhng kt qu thu
c khng nh vy. Do vy s c mt gen trn nhim sc th thng v
mt gen lin kt vi gii tnh. Rui phi mang gen trn nhim sc th
thng. Nu rui mang gen lin kt vi gii tnh th rui F1 s c mt
mu cam. Vy rui phi mang gen lin kt vi gii tnh. Mt gen trn
nhim sc th thng v mt gen lin kt vi gii tnh s cho t l phn
li 3:3:1:1 t nht l rui F2 (KN4-F). Trong s rui F2 chng ta
nhn c t l 5 mt mu cam : 3 mt mu , mt t l bin i
ca t l 3:3:1:1 (xem chng 2).
Ta qui c a = mu cam lin kt vi gii tnh, v b = mu cam trn nhim
sc th thng. Cc php lai by gi s l:

bb, X+X+

x ++, XaY

+b, X+Xa
F1

()

F2

x +b, X+Y
()

3 X+-, +

3 X+-,+

1 X+-,bb
cam

3 Xa-,+cam
1 X+-,bb
cam
1 Xa-,bb
159

cam
12.a. XN+X+V, ++; X++Y, ee .
b. Rui F1 cnh bnh thng : 3 bnh thng, mt , thn xm : 3
bnh thng, mt son, thn xm : 1 bnh thng, mt , thn
en : 1 bnh thng, son, thn en v u hai gii.
Rui F1 cnh khuyt:
3 khuyt,
thn xm

mt

3 bnh thng, mt
, thn xm

3 bnh thng, mt ,
1 bnh thng, mt
thn xm
, thn en
1 khuyt,
thn en

mt

1 bnh thng, mt ,
thn en
Chng ta thy khng c rui cnh khuyt, do vy cnh khuyt phi l
gen gy cht khi trng thi bn hp t (KN4-G). V rui c mt mu
son cho nn rui m phi d hp t v gen qui nh mu son
(KN4-C v 4-D).
Nu k hiu XN= cnh khuyt, XV = son, X+ = bnh thng v +/e =
xm/en. Chng ta c th vit s php lai nh sau:

XN+X+V,++ x X++Y,ee

XN+X++,+e
(xm,khuyt)

X++X+V,+e
(xm)

XN+Y,+e
(cht)

X+VY,+e
( son,xm)

T l phn li F2:
X++X+V,+e x X+VY,+e

3 X++-,+-

bnh thng, mt , thn

xm

160

1 X++-,ee

bnh thng, mt , thn

en
3 X+V-,+-

bnh thng, mt son,

1 X+V-,ee

bnh thng, mt son,

thn xm
thn en
Php lai vi rui ci cnh khuyt l:
XN+X++,+e x X+VY,+e
Mt na s rui c s nhn XN+ v cht (KN4-D v 4-G); Nhng rui
sng st s c cnh bnh thng v mt . Trong s rui c, 3/4 s c
thn xm (xem chng 1).
T l phn li rui :
3 XN+X+V, +-

Cnh khuyt,
thn xm

mt

1 XN+X+V, ee

Cnh khuyt,
thn en

mt

3 X++ -, +-

Cnh bnh thng,

, thn xm

mt

1 X++ -, ee

Cnh bnh thng,

, thn en

mt

13.C hai gen, mt gen ln trn nhim sc th thng qui nh mt

mu nu v mt gen ln lin kt vi gii tnh qui nh mt mu
trng. Bt c rui ng hp t/ bn hp t v gen qui nh mu trng
no cng s cho mt mu trng, d c mt cc gen khc. F 1 biu hin
kiu di chng t c hai gen (xem chng 2) v F2 c s khc nhau v t
l phn li gii v gii , chng t rng t nht c mt gen lin kt vi
gii tnh (KN4-C). Mt na s con F2 c mt mu trng, y l t
l phn li ca mt gen ln lin kt vi gii tnh (KN4-D). Chng ta nhn
c t l phn li 3 : 1 nu, l t l phn li ca mt gen trn
nhim sc th thng. T l phn li rui F2 l 3 : 1 nu, cho
thy tt c rui c t nht mt nhim sc th X bnh thng (X+).
Ta qui c: X+-A- = ; X+-aa = nu; Xw = trng
Php lai s l:
X+X+aa x X+ YAA

X+XwAa

X+YAa
161

(tt c )

t phi
3 X+-, A-
1 X+, aa nu

3 X+Y a-

1 X+Y aa nu
3 XwY A- trng
1 XwY aa trng

Chng ta nhn c mt t l phn li bin i ca t l 3:3:1:1 trong

s rui F2, cho thy c mt gen trn nhim sc th thng v mt
gen lin kt vi gii tnh (KN4-F, xem chng 2). S rui nhn c gn vi
t l 4:3:1.
14.a. 3 : AA, BB v AB.
b. Ch c mt, hoc A hoc B.
Rui gim s to c c hai dng tiu n v trong cc t bo, chng
c th t hp thnh dng AA, BB v AB. ngi, ch mt trong hai
nhim sc th X l hot ng (KN4-H), do vy mt t bo ch to c
hoc AA hoc BB. ngi tn ti c hai dng cu trc, nhng ch mt
dng c mt trong mt t bo c th.

Chng 5: Li gii

162

1. a. Lin kt gen.
b.WS/ws.
c. 10,7 n v bn .
Chng ta thy rng php lai ny l php lai phn tch v cc lp kiu
hnh chia thnh 2 nhm c tn s khc nhau. Do vy cc gen phi lin
kt khng hon ton (KN 5-A v 5-D). Cc lp c tn s cao nht l:
ng mu, ngn (W-S-) v m, di (wwss). V vy kiu gen ca con
ci l WS/ws (KN 5-B). tnh tn s trao i cho (RF), ta c:
RF=17/158 x 100% =10,7% =10,7 n v bn (KN 5-A).
2. a. Lin kt gen.
b. Sp xp kiu i ( trans).
c. 28, 6%.
y l php lai phn tch. Nu cc gen khng lin kt, chng ta thu
c t l 1:1:1:1 (KN 5-A v 5-B). Cc alen trng thi ng (cis) s xut
hin lp c t l ln v l kiu hnh cao, trn v thp, nhn. V
vy gen quy nh tnh trng cao, nhn nm trn cc nhim sc th
khc nhau. Nu chng ta gi T/t l cp gen quy nh tnh trng
cao/thp v R/r: nhn, trn th php lai l Tr/tR X tr/tr v cc th ti t
hp l TR v tr. Tn s ti t hp l: (105+110) x 100%/751 = 28, 6%
3. c

a 3

10

Ta bt u vi cc gen c khong cch ln nht: a-e = 25. V a-f = 7,

chng ta th t f nm gia a v e:
a 7

18

Gen d c th nm hai pha ca f theo th t afde hoc adfe. V f-d =

8 v d-e = 10 nn trt t phi l afde. V vy ta c trt t gen sau:
a 7 f 8 d

10

V b-d = 12 v a-b =3, chng ta c:

a

10

Nu c-f = 13 th c phi cch a 6 n v bn v bn tri, ng

theo gi thit c-b = 9. Chng ta c c bn di truyn ca cc gen
k tip nhau v c khong cch bn cng k tip (KN 5-L).
4. Gen d nm gia. Hy nhm cc lp tng h vi nhau (KN 5-E):
DeF

76

dEf

82

def

30

DEF

24
163

Def

15

dEF

18

deF

DEf

B m phi c kiu gen DeF/dEf (KN 5-B) v cc lp DEf v deF cha

trao i cho kp(KN 5-F). Hy so snh cc lp khng c ti t hp vi
cc lp c trao i cho kp. Cc alen eF v Ef u di cng vi nhau
trong c hai trng hp. V vy D phi nm gia (KN 5-G). Chng ta
vit li kiu gen ca b m: FDe/fdE
5. a. dy 35, 5

b 5, 4 bt

b. 0,74.
F1 cho ta bit tt c 3 alen t bin u l ln. Gi +/dy l cp gen
quy nh tnh trng thn mnh/thn b, +/b: trng/en, +/bt: lng
thng/lng qun. S lai l:
dy + + /+ b bt X dy b bt/dy b bt
Ta sp xp cc nhm kiu hnh tng h nh sau(KN 5-E):
+bbt: 301,

dy++: 305

++bt: 8, dyb+: 6
+++: 169,

khng ti t hp
trao i cho kp

dybbt: 172:

trao i cho n gia dy v

b
+b+: 19, dy+bt: 21: trao i cho n gia b v bt
Ta c trao i cho kp to ra ++bt v dyb+ (KN 5-F). Cc alen quy
nh tnh trng lng cong v thn b lun lin kt vi nhau c dng
b m v dng con c trao i cho kp. V vy alen b phi nm
gia (KN 5-G). Chng ta v s trao i cho kp khng nh
iu . Vi trt t gen nh vy, chng ta suy ra lp +++ v dybt cha
trao i cho n gia dy v b; cc lp +b+ v dy+bt cha trao i
cho n gia b v bt. Chng ta tnh khong cch gia dy v b:
RFdy-b= (169+172+8+6) x 100/1000 = 35, 5
Khong cch gia b v bt:
RFb-bt = (21+19+6+8) x 100/1000 = 5, 4% (KN 5-H)
Tn s trao i cho theo l thuyt:
= 0,355 x 0,054 = 0,019
Tn s trao i cho thc t:
= 14/1000 = 0,014
Vy CC = 0,014/0, 019 = 0,74 (KN 5-H)
6. a. k+cd/+e+
164

b. k

e 5 cd

Cc gen ny lin kt vi nhau (KN 5-A). Php lai ban u l:

kcd+/kcd+ X ++e/++e
v F1 c kiu gen kcd+/++e vi mt trt t gen cha xc nh. Hai lp
kiu hnh cui cng c tn s thp nht v l cc lp c to ra do
trao i cho. Hai lp kiu hnh ny s cho php xc nh trt t gen.
Nu trt t gen l kcde, trao i cho kp s to ra k++ v +cde. Vy
trt t gen phi l: kecd (KN 5-F v 5-G). V vy:
RF(k-e)= (32+34+2+1) x 100/1000 = 7n v bn
RF(e-cd)=(25+23+1) x 100/1000 = 5 n v bn
(KN 5-H)
7. Hai gen lin kt nhau nhng cch nhau trn 50 n v bn .
Gi dy v b l gen quy nh tnh trng cnh b v mt nu tng ng.
F1 cho thy cnh b v mt nu l tnh trng ln v F 1 phi c kiu
gen l dyb/++. T l F1 khc 9:3:3:1 nn cc gen ny khng phn ly
c lp m chng lin kt nhau (KN5-A). V khng c trao i cho
rui gim c nn chng ta c th kim tra c khong cch gia hai
gen . T l y l theo kiu lin kt ca hai gen cch nhau trn
50n v bn v xp x 5:1:1:1(KN 5-K). Theo l thuyt, t l
phn ly s l 100: 20: 20: 20: 20. Do vy ta cn kim tra chnh xc
bng cch tnh khi bnh phng
8. a. Gen quy nh tnh trng trn v nht lin kt, tnh trng c mu
phn ly c lp.
b. RS/rs; Sp/sp.
c. 20,5 n v bn .
4 lp kiu hnh c t l ln v bng nhau cho thy c mt gen khng
lin kt (KN 5-J). Nu hai gen khng lin kt, chng c th to ra t
l1:1:1:1 trong s nhng lp c tn s ln. Cc lp kiu hnh trn/dt vi
c mu/nhn v c mu/nhn vi nht/khng nht c t l nh vy Vy
cc gen trn v nht lin kt. Cc alen tn ti trng thi ng RS/rs (KN
5-A v 5-B)
RF = (38+33+40+35) x 100/710 = 20,5%
9. Gen A v B lin kt, cch nhau 20 n v bn ; C phn li c lp.
Chng ta khng thy 8 lp kiu hnh c t l khng bng nhau, do vy
c t nht 2 gen lin kt nhau (KN 5-A). Nu tt c 3 gen lin kt, chng
ta c 2 lp c t l ln ch khng phi 4 lp. V vych c 2 gen lin kt
(KN 5-B v 5-J ). xc nh hai gen no lin kt, chng ta xt hai gen
cng c mt hai lp c t l ln:
n ho, nhanh nhn:

780 + 810
165

Khng n ho, chm chp:

820 + 790

V ch hai lp kiu hnh c t l ln nn cp A/a v B/b lin kt. C

khng lin kt v to ra t l 1:1:1:1 vi c hai cp A/a v B/b.V vys
lai l:
Ab/Ab cc

X aB/aB CC

F1

Ab/aB Cc

ab/ab cc

Php lai phn tch:

Ab/aB Cc

V cc alen sp xp trng thi i F1 nn cc th ti t hp l AB

v ab.
RF = (192 + 208 + 195 + 205) x 100/4000 = 20 n v bn
10. Qr = qR = 42, 5%; QR = qr =7, 5%.
Trc ht ta lp php lai:
Qr/qR X

qr/qr

Ta nhn thy cc lp Qr v qR c tn s ln (KN 5-B). V RF(Q-R) = 15%,

nn 15% th h con s l dng ti t hp (QR v qr), (KN 5-I). Cc lp tng h s bng nhau, do vy QR = qr =7,5%. Qr = qR = 42.5% (KN 5E).
11. + b stw

0,393

fz + +

0,393

+++

0,072

fz b stw

0,072

+b+

0,032

fz b stw

0,032

+ + stw

0,003

fz b +

0,003

Trc khi tnh ton, hy xc nh nhng trao i cho n gia fz v

b, l + + + v fz b stw; v cc trao i cho n gia b v stw,
l + b + v fz + stw. Cc trao i cho kp l + + stw v fz b +.
V fz-b = 15, b-stw = 7, nn tn s trao i cho kp l thuyt =
0,15 x 0,07 = 0,0105. Vy
CC = 0,6 x 0,0105 = 0,006
V cc lp tng h c tn s bng nhau, nn tn s ca mi lp + + stw
v fz b + s l 0,003.
Khong cch gia fz-b = 0,15 = (tn s trao i cho n + tn s
trao i cho kp)/tng s = (tn s trao i cho n +
166

0,006)/tng s; vy tn s trao i cho n = 0,144 v tn s ca

mi lp = 0,072.
Khong cch gia b-stw = 0,07 = (tn s tro i cho n +
0,006)/tng s; Vy tn s trao i cho n = 0,064, tn s ca
mi lp = 0,032.
Tn s cc lp khng c ti t hp cn li th h con l: 1, 000 - 0,214
= 0,786 v tn s ca mi lp = 0,393 (KN 5-I).
12. Ab, Cd; Ab, cD; aB, Cd; aB, cD.
S lai:
AB/AB;CD/CD

AB/ab; CD/cd

ab/ab;cd/cd
X

ab/ab; cd/cd

Cc lp c tn s t nht s c to thnh t s ti t hp ca hai b gen

(KN 5-B).Cc lp ny s l: Ab Cd, Ab cD, aB Cd v aB cD.

167

Chng 6: Li gii
1. a. Gen ln nm trn nhim sc th thng.
b. I-1, I-2, II-1, II-2, III-4, III-5
c. 2/3
d. 1/6
V b m bnh thng sinh ra mt a tr b bnh, nn gen quy nh
tnh trng bnh phi l gen ln (KN 6-B). Gen ln ny khng th lin
kt vi nhim sc th X. I-1 c th mang gen ln ny II-3 b
bnh . Bt k 2 c th b m bnh thng no c con mang gen ln
v biu hin tnh trng ln u phi l dng d hp t (xem chng 1).
V II-3 phi c kiu gen pp nn tt c nhng a tr bnh thng :III4v III-5 phi l d hp t.
V s giao phi gia II-1 v II-2 theo s lai: Pp X Pp nn c hai c
th III-1 v III-2 u c 2/3 kh nng c kiu gen Pp. l nhng c
th bnh thng do vy n khng th c kiu gen pp. V III-3 c 2/3
kh nng c kiu gen Pp v v III-4 l PP nn xc sut c hai u
c kiu gen Pp l:2/3 x 1 = 2/3 (KN 6-H). Nu hai c th d hp t giao
phi vi nhau, th xc sut con b bnh l 1/ 4. Do vy xc sut
cui cng l: 2/3 x 1/4 = 1/6.
2. Gen lin kt Y, gen tri lin kt X, gen ln lin kt X, gen tri nm trn
nhim sc th thng.
Tnh trng lin kt Y ch c truyn t b cho con trai. Nu l tnh
trng tri th t nht mt trong hai b m phi mang gen . Nu
l gen ln lin kt X th tt c con gi phi c X mang gen bnh thng
(tri) t b (KN 6-B, 6-C).
3. A: Gen ln lin kt X, gen ln trn nhim sc th thng.
B: Gen tri lin kt X, gen tri trn nhim sc th thng.
168

C: Gen lin kt Y, gen tri trn nhim sc th thng, gen ln trn

nhim sc th thng, gen ln lin kt X.
D: Gen tri trn nhim sc th thng.
ph h A, tnh trng c di truyn bi gen ln v n b ngt qung
mt th h (KN 6-B). Nu ngi ph n l d hp t th 1/2 s con trai s
mang bnh (KN 6-C). Nu gen nm trn nhim sc th thng, chng ta
s c 1/4 s con mang bnh.
ph h B, gen quy nh tnh trng phi tri. Nu n l gen ln,
c hai b m phi l ng hp t v tt c th h con phi b bnh.
Nu gen ny lin kt X, tt c con gi s b bnh (KN 6-E).
ph h C, gen quy nh tnh trng lin kt Y, v ch con trai biu
hin bnh. Nu l gen tri trn nhim sc th thng, ngi ph n
phi c kiu gen ng hp t ln v ngi chng ca c y phi c kiu
gen d hp t (KN 6-A). Mt na s con ca php lai ny (Aa X aa) phi
b bnh. Nu l gen ln lin kt vi X v ngi ph n mang gen
bnh (d hp t), th c 2 ngi con gi phi nhn nhim sc th X
bnh thng (mang gen tri) v c 2 con trai mang gen ln X a. Nu
l gen ln trn nhim sc th thng v ngi ph n l d hp t th
1/4 s con phi b bnh.
ph h D, tnh trng do gen tri quy nh . Nu l gen ln, cc
con u phi b bnh. N khng th l gen tri lin kt X, v nu vy
th tt c con gi phi mang bnh (KN 6-E).
4. A nm trn nhim sc th 6, B nm trn nhim sc th 2, C trn
nhim sc th 13, D trn nhim sc th 12, E trn nhim sc th 10.
Gen A c mt dng X, Y, Z v nhim sc th 6 c mt c 3 dng
ny. B ch c dng Y v nhim sc th s 2 l nhim sc th duy
nht i vi Y.
L lun tng t cho php suy ra cc gen khc(KN 6-J).
5. Nu tnh trng l ln th b v m bnh thng phi c con biu hin
bnh (KN 6-B). Nu mt tnh trng c xc nh bi gen ln lin kt
vi X, nam gii mang alen ny s biu hin bnh. V c th b I-a khng
biu hin bnh, nn anh ta khng th mang gen t bin nm trn X.
Mt tnh trng c xc nh bi gen tri lin kt X phi xut hin tt
c con gi ca ngi b b bnh (KN 6-E).

169

(a)

(b)

6. 1/8. Cc c th I phi c kiu gen IBIOv IAIO tng ng sinh ra 1

a tr c nhm mu O. By gi chng ta xt cc kiu gen c th c
ca mi c th. II-1 phi l IOIO, II-2 l IAIO, II-5 l IAIB, II-6 phi l IBIO,
hoc nhng a tr khc phi c nhm mu AB hoc B. III-2 phi l IA
IO, III-3 c xc sut 1/ 2 l ng hp t v gen IB v 1/ 2 l IB IO. Nu III-3
c kiu gen IB IO s c 25% kh nng a tr c kiu gen IOIO. V vy
p s l:
1/ 4 x 1/ 2= 1/8 (KN 6-H).
7. a. Gen tri lin kt X.
b. Gen tri trn nhim sc th thng, gen tri lin kt X, gen lin kt Y.
c. Gen ln lin kt X, gen lin kt Y.
d. Gen tri lin kt X, gen lin kt Y, gen tri trn nhim sc th thng.
Chng ta ch c th loi b nhng kiu di truyn hon ton tri vi ph
h.
(a), chng ta loi b trng hp gen tri lin kt X, v cc con gi II
u bnh thng (KN 6-E). Kh nng gen tri trn nhim sc th thng
c th xy ra ngay c khi chng ta khng thy con gi biu hin bnh .
Hai ngi ph n I-1 v II-4 c th l d hp t cho d kh nng ny t xy
ra ( KN 6-A v 6-G).
(b) chng ta loi b tt c nhnh trng hp gen tri v tnh trng ny
ngt qung mt th h (KN 6-B). Gen lin kt Y c truyn t b sang
con trai (KN 6-F).
(c), chng ta loi b trng hp gen ln lin kt X v tt c con trai
ca II-5 u bi hin bnh (KN 6-C). Nhng ngi ph n u mang
bnh do vy gen ny khng th lin kt Y. Nu I-1 v II-1 u d hp t
v gen ln nm trn nhim sc th thng (KN 6-G), th 1/2 s con
phi biu hin tnh trng .
(d) chng ta loi tr trng hp gen lin kt Y v n gii cng mang
tnh trng ang xt. Ta cng loi tr c trng hp gen tri lin kt X v
II-4 c cc con gi bnh thng (KN 6-E). Chng ta loi tr trng hp gen
tri trn nhim sc th thng v c hai c th bnh thng to ra con
b bnh (KN 6-B).
8. a. 1/ 2
b. 1/ 4
Nhng ngi biu hin tnh trng tri lun l d hp t (KN 6-A), do vy
ngi n ng c th c kiu gen Hh v con trai c 1/ 2 kh nng c
kiu gen Hh. V anh ta phi c kiu gen hh. Nu ngi n ng c kiu
gen Hh, ta c php lai: Hh X hh v nhn c s con c kh nng b
170

bnh Huntington l 50% (chng 1). V vy xc sut h c mt a

tr b bnh Huntington l: 1/ 2 x 1/ 2= 1/ 4 (KN 6-H).
9. a. 243/1024
b. 135/512
c. 9/1024
(a), v tt c nhng a con c cng kiu hnh, nn mi a tr s c
cng kh nng c kiu hnh khng c rng hm. V vy (3/ 4) 5 =
243/1024 (KN 6-G).
(b), n = 5, s = 3, t = 2, p = 3/4 & q = 1/4.
V vy P = 5!/3! 2! (3/4)3 (1/4)2 = 135/512
Khi xc nh c phng trnh, chng ta nhn cc xc sut vi nhau,
v xc sut xut hin mi kiu hnh l mt hng s. Do vy :
3/ 4 x 3/ 4 x 1/ 4 x 1/ 4 x 1/ 4 = 9/1024 (KN 6-G).
10. a. 27/64
b. 9/4096
Vi 3 a tr, cng thc l: 3!/2! 1! (3/4)2 (1/4) = 27/64. (KN 6-J)
Xc sut 1 b trai hoc 1 b gi bnh thng = 3/ 4 x 1/ 2 = 3/ 8.
Xc sut 1 b trai hoc 1 b gi b bnh l 1/ 4 x 1/ 2 = 1/ 8.
Chng ta nhn cc xc sut vi nhau v c :
3/ 8 x 1/ 8 x 1/ 8 x 3/ 8= 9/ 4096 (KN 6-H).
11. 1/ 8.
Ngi v (2) phi l d hp t Aa v nhn A t b. V khng cp n
bnh ny b m ngi chng, nn ta gi s m anh ta l AA v ngi
chng nhn alen A t m. Vy xc sut ngi chng (1) nhn a t b
anh ta l 1/2. Nu 2 c th d hp t giao phi, xc sut con mang
tnh trng ln l 1/ 4. V vy P = 1/ 4 x 1/2 = 1/ 8 (KN 6-G v 6-H).
V khng bit r lch s bnh, nn mi a tr u c kh nng b
bnh nh nhau.
12. a. 3/8
b. 5/16
Gi T l gen quy nh kh nng chuyn ho (nm c v ca PTC). t l
khng c kh nng chuyn ho (khng nm c v ca PTC). V vy ta
c php lai: Tt x tt, mt php lai phn tch, s c 1/ 2 c hi ngi
con b bnh hoc bnh thng (xem chng 1). Do vy p = q = 1/ 2, n =
4, s = 2 = t. Suy ra P = 4!/2! 2! (1/2)2 (1/2)2 = 3/8 (KN 6-J).
(b), n = 6, s = t = 3. Vy P = 6!/3! 3! (1/2) 3 (1/2)3 = 5/16. Ch rng
khi n tng th P gim do c nhiu kh nng khc na.
171

13.a. 1/32
b. 3/ 4
Gi M/m l t l khng c/c rng hm, A/a = bnh thng /bch tng v
T/t = bnh thng/Tay-Sachs. Ta c php lai mm Aa Tt x Mm Aa Tt, vi
kt qu to l 50% s con c rng hm, 75% c kiu da bnh thng v
75% c kiu hnh bnh thng i vi bnh Tay-Sachs. trng hp (a),
chng ta nhn cc xc sut vi nhau s c: (1/2) x (1/4) x (1/4) = 1/32
(KN 6-H). trng hp (b) c 50% kh nng s con khng c rng hm v
25% kh nng b bnh Tay - Sachs. V chng ta cn 1 trong 2 trng hp
m khng phi l c hai nn chng ta cng cc kh nng li: 1//2 + 1/4 =
3/4 (KN 6 - I).
14.a. Dng B.
b. Dng A v C.
Chng ta cn xc nh cc dng c nhim sc th 11; cc dng ny
hy vng s tng hp c enzim. Chng ta thy rng dng b c nhim
sc th 11 v dng A v C c nhim sc th s 5 (KN 6-K).
15. Tnh trng l tnh trng tri lin kt nhim sc th X v gy
cht nam gii. Chng ta thy s khc nhau v kiu hnh i con gi
n s lin kt gii tnh (KN 6-A, xem chng ). Chng ta khng thy
nhng ngi con trai b bnh nhng li c nhng ngi con trai c kiu
hnh bnh thng; iu chng t nhng ngi con trai nhn
c alen ln. Vy ngi m phi c kiu gen d hp t v tnh trng ny.
Gi XI : gen gy bnh v Xi: gen quy nh kiu hnh bnh thng.
Php lai khi s l:
XI Xi

Xi Y

XI Xi
b bnh

Xi X i
bnh thng

Xi Y

XI Y

bnh thng

cht

Chng 7: Li gii
1. 34.
Gim phn Q v R to ra cc giao t vi 7 v 10 nhim sc th tng
ng. Hp t c 17 nhim sc th v khng nhim sc th no kt cp
c. con lai hu th, mi nhim sc th phi c gp i ln. Con
172

lai c 2 b nhim sc th t Q (2 x 7) v 2 b nhim sc th t R (2 x

10). Vy con lai l th d t bi (KN 7-H).
2. Kiu gen ca ngi con trai c th l Xb XbY. Hy bt u bng vic xc
nh kiu gen ca b m. Lun coi rng h bnh thng tr khi c bng
chng tri li. Kiu gen ca ngi ph n phi l XBXb v ngi n ng l
XbY. Nu ngi con trai nhn c Xb t m v Y t b, anh ta c kiu gen
XbY v c kiu hnh rng nu nhng l bnh thng. Nhng kiu hnh
khc, c bit l tinh hon pht trin km gi n hi chng
Klinefelter, XXY. S phn li khng bnh thng c th xy ra gim
phn I ngi cha to thnh giao t XbY (KN 7-G). Hoc s phn li
khng bnh thng c th xy ra gim phn II ngi m to ra 1
trng c XbXb.
3. a. Ch thy 8 kiu hnh m theo l thuyt phi l 25 = 32.
b. Con ci F1 c th l d hp t do mt o on cha c v d.
c.

b 3
+

d 5

c 5
+

e
+

o on
Chng ta d on c mt o on v ta thy s kiu hnh t hn
mong i (KN 7-A). Chng ta c th xc nh v tr ca cc u ca
on o bng cch tnh tn s trao i cho gi cc gen.
RF gia a v b = 10/200 x 100 = 5 n v bn , ng bng khong
cch trn bn ; khng c o on.
RF gia b v c = 6/200 x 100 = 3 n v bn , nh hn mong
i.
RF gia d v e =10/200 x100 = 5 n v bn , nh hn mong
i.
Chng ta thy khng c ti t hp gia c v d, v vy c v d phi nm
trong on o. Khong cch t b ti im t gy c th tnh t cc
th ti t hp gia b v c-d, hoc bng 3 n v bn . L lun tng
t vi im t gn e v khong cch t im t n e = 5
n v bn ( KN 7-F). V vy nhim sc th b o on rui
ci ban ul:
a
10

15 18

e
35

40

on o
4. Bng 1 l q, 2: p, 3: r, 4: u, 5: t, 6: s.
Nu mt mt on cho php biu hin mt alen ln, th mt trong
cc bng c di ng vi on mt phi tng ng vi gen cn tm (KN
7-B). Cc bng 1, 2 v 3 phi tng ng vi cc gen p, q v r theo mt trt
173

t no . Cc bng 2 v 3 cng b mt trong nhng mt on B v D;

cc kiu hnh p v r c mt trong cc dng ny.
V vy 1 = q. So snh cc mt on 2 v 3, c hai u ko di n
bng 3. Gen p ch xut hin B; vy 2 = p v 3 = r. Mt on B v C
cng ko di ti bng 4, mt on C v E ko di n bng 4 v 5. V u
l tnh trng duy nht c tt c 3 mt on, nn u = 4.
L lun tng t th t = 5, s = 6 (KN 7-D).
5. a. Chuyn on
b. Sh cch dim t 19 n v bn .
Bn bt th thng lin quan vi chuyn on ( KN 7-E). V kiu hnh
Sh cng l bn bt th, nn b m Sh phi mang chuyn on. sh bn
bt th v i con Sh hu th l kt qu ti t hp gia gen v im
t gy. Tn s trao i cho s cho ta thy khong cch gia Sh v
im t (KN 7-F)
RF = (195 + 185)/2000 x 100 = 19 n v bn .
6. a. 9 nhim sc th, c 3 b A, B, C.
b. Bt th.
Gim phn mt th t bi s to ra cc giao t c 2 b nhim sc th
(xem chng 3). S th tinh s to ra : 2n+n = 3n hoc 3A, 3B v 3C. V
con lai l th tam bi, n s bt th (KN 7-H, xem chng 3).
7. a. Tt c con ci c thn xm; mt on phn nhim sc th X ca
con c.
b. 2 ci thn xm : 1 c vng.
Trc ht, vit s lai vi k hiu X+ : thn xm v Xy : thn vng.
Xy Xy x X+Y

X+Xy : XyY
Xm vng
Vic s dng tia X lm chng ta ngh ti kh nng t gy nhim sc
th. Chng ta gp kiu hnh ln khng mong mun, gi n mt
mt on nhim sc th X ca rui c (KN 7-B). Cc nhim sc
th X ca rui ci F1 vng c th l:
y
//////
Nu chng ta lai rui ci ny vi mt rui c thn xm khng b chiu
x, th php lai l (X*: nhim sc th X b mt on):
XyX* x X+Y
174


X+Xy : X+X* : X*Y : XyY
xm xm cht vng
Cc con c nhn c nhim sc th X mt on b cht (KN 7-C).
8. 34. Gim phn A v B s to ra cc giao t c 11 v 6 nhim sc th.
S th tinh s to ra mt con lai vi 17 nhim sc th v khng nhim
sc th no c th kt cp c. Nu mi nhim sc th con lai c
gp i th s to ra mt th a bi hu th (KN 7-H).
9. iu khng th xc nh c. Gi Xa: m mu. Php lai s l: XAXa x
XaY.
Hi chng Tcn c kiu gen : XO. Nu s khng phn li nhim sc
th xy ra b, th s to ra giao t XY v mt giao t khng c nhim
sc th gii tnh. Con gi c th nhn Xa t m.
Nu s khng phn li nhim sc th xy ra m th trng c th
khng c nhim sc th X v con gi c th ly Xa t b (KN 7-G).
10. S khng phn li nhim sc th xy ra trong gim phn II ngi
n ng bnh thng. Cn theo di cc nhim sc th gii tnh ngi
n ng qua c 2 ln phn chia ca gim phn. Hai nhim sc th Y
phi c ngun gc t ngi n ng (KN 7-G, xem chng 3).

Gim phn I

Chuyn ng v cng mt
cc

Giao t YY

175

Gim phn II

11.a. Chng ta thy tn s ti t hp gia a v b l 5% m ng l tn s

phi l 19%; v vy c th xy ra o on gia a v b.
b.

b
o on

Chng ta thy tt c nhng kiu hnh mong i, nhng tn s ti t hp

t hn mong i. Mt mt on c th lm gim khong cch bn ,
nhng kiu gen bit v tn s th h con gi cho chng ta thy
khng c mt on. Kh nng khc s lm gim tn s ti t hp l mt
o on (KN 7-A). Tn s trao i cho gia a v b l: 5/100 x 100
= 5%. V vy a phi cch im bt u ca o on cha b l 5
n v bn .
12. c, d, a, b. Chng ta thy rng phi xy ra o on to nn mi
chng mi; chng ta tm o on n gin nht to ra chng
mi. Chng d c th nhn c t c bng o on 9 4 3 thnh 3 4 9.
Nu trt t 8 7 3 4 d b o th chng ta nhn c trt t a. Cui
cng, o on 6 5 4 a to ra trt t b (KN 7-A).
13. Chuyn on t u ca nhim sc th X sang nhim sc th Y.
Php lai rui c F1 thn xm vi rui ci thn vng theo l thuyt s
to ra cc rui c thn vng. Vic chiu x gi n 1 t gy nhim
sc th. Gi X+: thn xm v Xy: thn vng, vit s lai:
Xy Xy x X+Y

X+Xy : XyY
Rui ci xm rui c vng
Theo l thuyt tt c rui c u mu vng. Rui c thn xm c
th nhn c mt alen bnh thng v mu thn t con c b chiu
x. Cch gii thch ng tin cy nht l mt mnh nh nhim sc th
X b gy v gn vi nhim sc th Y.
+
X

+
Y

Chuyn on gia X v Y c cha + s lm c ch alen vng t m (KN

7-F v 7-G). Nu rui c ny c lai vi rui ci vng, th tt c cc
rui c i con s nhn c nhim sc th Y c chuyn on v c
thn xm.
14.a. Nhim sc th 15 v 21 bnh thng; nhim sc th 21 bnh thng, nhim sc th 15 chuyn on; nhim sc th 21 b mt on,
nhim sc th 15 chuyn on; nhim sc th 21 mt on; nhim
sc th 15 bnh thng.
176

b. 1/3
Cch tt nht l v chuyn on v theo di nhim sc th 15 v 21
qua gim phn.

Gim phn s sinh ra

Giao t bnh thng

Giao t bnh th- Giao t mt on

Giao t lp on
ng v chuyn
on

S th tinh to ra

C th bnh thng

C th bnh thng c
mang chuyn on

177

C th b bnh
Down

Khng sng c?

(KN &-C v 7-G, xem thm chng 3)

Tnh hung thc thng phc tp hn ta v trn y, v thng thng c
hn 4 loi giao t c to ra, ph thuc vo nhim sc th 21 chuyn
on di hay ngn. Tham kho chuyn on Robertson v hi chng
Down lin quan trong ti liu.
15.a. Gen notch quy nh cnh x c kt hp vi mt mt on; gen
facet quy nh mt que nm trong mt on ca gen cnh x notch.
b.
Mt on notch
2 vb

10 vb

facet

dwarf

Gi XN: notch, gen quy nh cnh x; Xd: dwarf, thn ln (ngn); Xf:
facet, mt que. Php lai 1 cho thy rui ci cnh x ban u l d hp
t v tnh trng cnh x l tnh trng tri. Vic khng c con c cnh
x cho thy gen quy nh cnh x ng thi l gen gy cht ln. (xem
chng 4).
S lai ca php lai 1 :
XN+ X++ x X+dY

XN+ X+d

X++ X+d

X++ Y

Cnh x

kiu di

kiu di

XN+ Y
cht

Php lai 2 cho thy rng cnh x l gen gy cht v chng ta li thy
khng c rui c cnh x. N cng cho php ta tnh tn s trao i
cho gia gen quy nh cnh x v gen thn ngn. V nhim sc th
X ca rui ci F1
N

d
178

Rui c kiu di l cc th ti t hp v tn s trao i cho RF =

34/334. 100 = 10 n v bn (xem chng 4 v 5). V vy gen n
cch d 10 n v bn .
Php lai 3 a ra 1 gen mi: gen quy nh mt que. Tnh trng ny
xut hin mt s rui ci th h con. Nu gen quy nh mt que l
tri, th tt c rui ci th h con phi l mt que, nhng kt qu
khng phi nh vy. Do vy gen quy nh mt que l ln v s xut
hin ca tnh trng ny mt vi rui ci cho thy rng c mt mt
on (KN 7-B). Nu gen quy nh cnh x lin kt vi mt mt on,
th mt on lin kt X s gy cht khi trng thi ng hp t hoc
rui c bn hp t (KN 7-C). Gi s gen mt que nm trong 1 mt
on lin kt vi cnh x th nhng con ci cnh x thn ngn l kt
qu ti t hp gia gen quy nh cnh x v gen quy nh thn
ngn.
Con ci ny c kiu gen:
N

d
+

Chng ta c th vit s lai:

N

+
X

+ +d

+f

+ f +
Y

+ + d
Thn ln

cht

cnh x, mt que

kiu di
Php lai 4 cho php chng ta lp bn gen d v f v tn s trao i
cho c tnh bng vic kim tra rui c i con ca php lai ny.
RF = 60/500 x 100 = 12 n v bn . V gen cnh x c th l mt
mt on cch d 10 n v bn v v gen mt que nm trong
on mt, nn on mt c th di t nht 2 n v bn (KN 7-F).
Bn gen ph hp nht l:
f

12
2

d
10
179

//////////////////////
Mt on notch

180

Chng 8: Li gii
1.
a

31

c 19, 0

Tin hnh ging nh phn tch cc gen lin kt (xem chng 5). Cc lp
c tn s cao nht: +bc v a++ l nhng dng khng phi do ti t hp
v nhng lp c tn s t nht l do trao i cho kp: l a+c v
+b+.
Do vy, chng ta c th xc nh trt t cc gen l acb. V:
RFb-c

= (34 + 14 + 28)/400 x 100 = 19, 0 n v bn

RFa-b

= (52 + 58 + 34 + 28)/400 x 100 = 43, 5 n v

RFa-c

= (52 + 14 + 58)/400 x 100 = 31 n v bn

.
bn .
.
2. 14 n v bn . Php lai l: nic- thi+ x nic+thi-. Kiu gen mc c
trn mi trng ti thiu l nic+thi+ v l kt qu t s trao i
cho gia hai gen. Dng ti t hp khc l nic- thi- s khng mc c trn
mi trng ti thiu. Do vy, tng s th ti t hp l 2 x 35 = 70.
V RF = 70/500 x 100 = 14 n v bn .
3. a______ 10 ___ ____ 8 ___ c . ___ 15.5 ____ b
Phn tch php lai u tin v phn bit bo t no l PD, NPD v TT.
Cc lp bo t I v VI = PD, II v VII = NPD, II, IV v V= TT. V PD = NPD,
nn a v b khng lin kt nhau. (KN 8-B v 8-C). By gi, hy phn bit
mi gen mi bo t l kt qu ca s phn ly ln phn chia I hoc
II (KN 8-E). iu ny cho php tnh khong cch gia gen v tm
ng (KN 8-F):
a- tm ng = 1/ 2(10 + 7 + 2 + 1)/100 x 100 = 10 n v bn
.
b- tm ng = 1/ 2(10 + 18 + 2 + 1)/100 x100 = 15, 5 n v
bn .
Tin hnh tng t php lai 2. V PD >>NPD nn a v c lin kt (KN 8C). Khong cch c tm ng l:
c- tm ng = 1/ 2(5 + 12 + 3)/100 x 100 = 8 n v bn .
By gi tnh khong cch a-c theo cng thc = (1/ 2 TT + NPD)/tng
s bo t x 100 (KN 8-F).
a-c = [1/2(5 + 8 + 12)/100 + 3/100] x 100 = 15, 5 n v bn .
181

Nu a v c nm mt pha ca tm ng, a-c c th cch nhau 2

n v bn . V kt qu ny khng ng nn a v c phi hai
pha ca tm ng.
4. 24.
Khong cch gen-tm ng = 1/2 s bo t phn ly ln phn chiaII/
tng s bo t x 100. V vy t l phn trm phn ly ln phn chia
th 2 = 2 x 12 = 24 (KN 8-E v 8-F).
5. .a v c lin kt v cch nhau 5, 5 n v bn ; b v d lin kt v
cch nhau 25 n v bn .
xc dnh s lin kt, hy so snh so snh PD v NPD. trng hp 1 v
3, PD = NPD, suy ra a khng lin kt vi b hoc d. php lai 4, PD >>
NPD suy ra b lin kt vi d (KN 8-C).
Chng ta tnh khong cch gia b - d, a - c theo cng thc:
(1/2 TT + NPD)/ tng s bo t x 100 (KN 8-D).
a-c = (1/2, 9 + 1)/100 x 100 = 5, 5 n v bn .
b-d = (1/2, 3 + 10)/100 x 100 = 25 n v bn .
6. 4.
V NPD c to thnh t trao i cho kp bn si, nn chng ta c
th coi nh khng c NPD v a v b nm rt gn nhau (KN 8-C). Khong
cch gia cc gen bng = (1/2TT + 0)/100 x 100 = 2; suy ra TT = 4.
7. ad_____ 6, 5 ____ _________12, 5 ______trp
Trc ht hy phn bit cc bo t do phn ly ln phn chia th 2:
l cc lp IV, V, VI i vi ad; v III, V, VI, VII i vi trp (KN 8-E). Khong
cch gia gen v tm ng l:
ad- tm ng = 1/2(18 + 6 + 2)/200 x 100 = 6, 5 n v
bn .
trp - tm ng = 1/2(30 + 6 + 2 + 12)/200 x 100 = 12, 5 n
v bn .
By gi, phn loi cc lp I v V l PD, II v V l NPD v III, IV v VII l
TT (KN 8-B v 8-C). V PD >> NPD nn cc gen ny lin kt. Ta tnh:
RF = [1/2(30 + 18 + 12) + 8]/200 x 100 = 19 n v bn .
8. _____ 23 ____ x ___ 6 ___ y ___ 7 ___ z
Chng ta phi phn tch ng thi hai gen. Hy bt u bng vic
phn loi kiu bo t.

Bo t

x-y

x-z

y-z
182

II

III

IV

VI

VII

VIII

Chng ta khng thy c NPD, do vy c 3 gen lin kt nhau (KN 8- C).

By gi, hy tnh khong cch gia gen v tm ng (KN 8-F).
x-tm ng = 1/ 2(70 + 4 + 4 + 2 + 6)/186 x 100 = 23 n v
bn .
y- tm ng = 1/ 2(70 + 18 + 4 + 4 + 2 +6)/186 x 100 = 28
n v bn .
z- tm ng = 1/ 2(70 + 18 + 14 + 4 + 4 + 6)/186 x 100 = 31
n v bn .
V, tnh khong cch gia cc gen (KN 8-D).
x-y = 1/ 2(18 + 4)/186 x 100 = 6 n v bn .
y-z = 1/ 2(14 + 4 + 2 + 6)/186 x 100 = 7 n v bn .
x-z = 1/ 2(18 + 14 + 4 + 4 + 2 + 6)/186 x 100 = 13 n v bn
.
C 3 gen ny u nm v mt pha ca tm ng.
9. a. PD = 80, NPD = 4, TT = 16.
b. PD = 51, NPD = 7, TT = 42.
Chng ta s dng cng thc: khong cch gia cc gen = (NPD + 1/2
TT)/ tng s bo t x 100 (KN 8-D).
i vi (a): 12 = NPD + 1/ 2(4 NPD), suy ra NPD = 4.
i vi (b): 28 = NPD + 1/ 2(6 NPD), suy ra NPD = 7.
10. 28 n v bn .
Chng ta li s dng ng thc:
Khong cch = (NPD + 1/2 TT)/tng s bo t x 100. Vy = PD =
12 NPD.
Khong cch = 7 NPD/25 NPD x 100 = 28 n v bn .

183

Chng 9: Li gii
1. a. ara nic gal hoc nic ara gal.
b. Chn lc theo gal, sau m s khun lc gal+ara+ v gal+nic+.
Nu trt t ara nic gal l ng th s khun lc gal+ ara+< gal+nic+. V
nic v ara ng bin np 40% thi gian nn chng phi gn v lin k
nhau. Nu gal gia th nic+gal+ phi c tn s ng bin np ln hn
nic+ara+ (KN 9-A).
Nu chng ta chn lc theo gal+ v th t l ara nic gal, th kiu gen
gal+ara+nic- ch cn hai trao i cho ng thi, trong khi gal+ara+
ch i hi mt trao i cho. Nu trt t l nic ara gal th gal+ ara+
ch cn mt trao i cho v tn s thu c phi ln hn gal+nic+ (KN
9-C).
2.. a v b gn nhau, c khng lin kt vi a v b.
Chng ta lu rng c+ t xut hin cng vi cc gen khc cho thy c rt xa
a v b. Cc gen khng lin kt s khng nm cng mt phn t AND. S
ng bin np s i hi 2 trao i cht ng thi (KN 9-B).
3.
Hfr-2

c
85

e
75

Hfr-1

0/90 5
20
b
30

60
50

a
chng 1, trt t truyn gen, c xc nh bi s xut hin gen u
tin, l a d b c e. i vi chng 2, trt t l b d a e c. Nhim sc th vi
khun c dng vng.
Trc ht xc nh thi gian truyn gia cc gen:
a-d = 20 pht, b-d = 10 phtV nhn t F c truyn sau cng v
v b (ti v tr 20 pht) xut hin sau 40 pht tip hp, nn nhn t F
thi im 60 pht, a cch nhn t F 10 pht (ti v tr 50 pht). L
lun tng t cho php tm ra cc gen khc v nhn t F th 2 (9-C).
4. 3 gen. Gen A: cc t bin 1, 4, 8; Gen B: cc t bin 2 v 5; Gen C: cc
t bin 6 v 7.
184

C l th t bin 3 cha mt mt on trm ln cc gen A v C. Hy

bt u bng vic tm ra cc t bin khng b tr cho nhau. Cc
t bin ny phi trong cng mt gen (KN 9-F). Cc t bin 2 v 5
trong cng mt gen. u tin chng ta ng rng cc t bin 1, 3, 4, 8
trong mt gen v 6, 7 trong mt gen khc. Nu th t bin 3 thuc
gen A n phi vi 6 v 7. iu khng xy ra.
Cch gii thch khc l t bin 3 l mt mt on trm ln A v C (KN
9-G). Kh nng khc l t bin 3 c th thuc gen A nhng l mt t
bin cc (KN 9-J). C hai iu u ng ni rng trt t ca 3 gen
l B A C.
M

5.

B
A

R
E

Hy sp xp theo trt t chng trm ln nhau:

1.

ACTER

2.
3.

TERIU
MBACT

4.I U M B A
V nhim sc th vi khun c dng vng nn ta c trt t biu din
hnh trn (KN 9-C).
6. a. rII-1 c th l mt mt on v rII-2 l mt t bin im.
b. C l cc th hi bin bin revb v revc cha t bin ti im th
hai.
S thiu vng cc th hi bin gi ta ngh n mt mt on, do
vy rII-1 c th l mt mt on (KN 9-K). Nu cc th ti t hp kiu
di no c phc hi t cc th hi bin, th cc th hi bin ny c
th l t bin c ch hoc l cc t bin v tr th hai (KN 9-L).
7. .def.
Chng ta thy rng f v d ng ti np gen t nht. V vy d c l nm
xa f v i hi 2 trao i cho ng thi (trao i cht kp) (KN 9185

A, 9-B, 9-C). Chng ta thy rng e v f c tn s ng ti np cao, do

vy e gn f hn d.
8. t bin C, t bin promoter, hoc t bin cc E hoc D.
V t bin khng to ra s b tr vi C- nn n c th thuc cng mt
mt gen (KN 9-F). Nu th t bin l p th th lng bi l: p- E D C B
A/ p+ E D C- B A. ADN bn trn c th khng to ra ARN v s khng c s
b tr (KN 9-J, xem chng 11). Mt kiu t bin khc cng s khng
b tr l mt t bin cc trong gen nm trc C (KN 9-J).
9. Cc chng 2, 5, 7 l nhng mt on.
5

////////////////

////////////

//////////////////////////
___*___*___*____*___*___*____
1

Nui cy trong mi trng ti thiu cho php pht hin cc th hi bin.

V cc mt on khng phc hi c nn chng khng mc c trn
mi trng ti thiu (KN9-K). bit 2, 5, 7 l nhng mt on, ta hy
phn bit cc chng ny t b s liu th hai.
2

Chng 2 khng cho th ti t hp kiu di, v vy n phi trm ln

hai t bin mt on khc (KN 9-H). Cc mt on 5 v 7 khng trm
ln nhau. Th t bin 2 cho kt qu tng t chng A nn A=2. V B
ti t hp vi 7, nn B phi l 5; do C l 7. Th t bin 9 khng cho
cc ti t hp kiu di vi cc mt on 2 v 7. N phi nm vng c
nh v duy nht trong t bin 5 (KN 9-I). L lun tng t i vi
t bin 8: 8 vng duy nht vi 7, 3 vng duy nht vi 2. Cc t
bin 4 v 6 phi nm trong vng mt on chung cho 2 v 7 cng nh
cho 2 v 5 tng ng. t bin 1 cho cc th ti t hp kiu di vi tt c
cc mt on, n phi nm bn ngoi tt c cc vng mt on. N c
th nm mt trong hai u tn cng ca bn .
10. a.

0. 2
|

|
b

0. 3
|

0. 1 0. 2
|

|
e

0. 5
|

0. 3
|

|
g
186

b.

1.
2.
3.
4.
5.

c. a, b, f trong rIIA; e, c, d v g trong rIIB.

d. Khng t hp no.
Cc t bin b v d phi xa nhau, b v g cng vy, e v f phi nm
gia b v d:
____b ___0.2 ___ f __0.4 ___ e ____ d ___ 0.3 __ g
By gi hy xc nh v tr ca a v c. V a-b = 0,5 nn a phi cch f
l 0. 3 v bn phi. t bin c phi bn phi a, hoc n c th cng v
tr vi f. t bin c phi bn phi a v khong cch c cng k tip
(KN 9-E). Hy tm cc du (-) trong cc php lai; cc t bin ny s b
ph bi cc mt on ring. Mt on 1 phi ko di dn cc t bin
a, c v e v khng c th ti t hp kiu di no xut hin (KN 9-I). Tng t, t bin 2 phi ko di n c, d, g. L lun tng t cho php
xc nh cc mt on 3-5. V a b tr e, vy a phi thuc rIIA, cng
ging nh b v f (KN 9-F). Tt c cc mt on u thiu c, do vy
khng th sinh ra cc th ti t hp kiu di (KN 9-H).
11. c a b pro.
Chng ta phi xy dng s ti np, s dng 2 trt t gen v tm ra
trt t s to ra v kiu di bng trao i cht kp. Trt t ny s cho ra
mt s t th ti np kiu di (KN 9-C). Gi s rng trt t l b a pro.
Nhng php u l:
b+

a-

pro+

b-

a+

pro-

b-

a+

pro+

b+

a-

pro-

Nu trt t nh trn, php lai th nht d on c s lng th ti t hp

kiu di nh hn php lai th 2 v cn 2 cp trao i cho ng
thi cho php lai 1. Kt qu ny khng thy c, do vy b phi nm gn
pro hn a. L lun tng t tm ra trt t ca c tng ng vi a v b.
12. 3%.

187

Chng ta phi tnh tng s c th i con v s con ti t hp. V tt

c cc phage sinh trng trong chng D, nn s khun lc sinh trng c
y phi l tng s c th i con, l 200 x 10 7. Kiu di ch
sinh trng chng C, do vy cc vt tan ny phi l dng ti t hp. Tuy
nhin, s lng thu c ch bng1/ 2 tng s th ti t hp, v cc th
t bin kp s khng sinh trng c. S th ti t hp l 2(30)(10 6) =
60 x 106. Suy ra
RF = (60 x 106)/(200 x 107) x 100 = 3% (KN 9-E).

Chng 10: Li gii

1. GAA thnh GUA hoc GAG thnh GUG. Xt cc b ba quy nh glu v

val, chng ta thy A v tr th hai i thnh U s cho ra mt axit
amin mi.
2. Cc axit amin ch c th l kt qu do thay i mt baz stop
b ba.
UAA c th bin i thnh

CAA gln

UCA ser

AAA lysUGA stop

GAA glu
UGA c th bin i thnh

UAG c th bin i thnh

UAG stop

UAC tyr

UUA leu

UAU tyr

UAA stop

UGG tyr

GGA gly

UCA ser

UGC cys

AGA arg

UUA leu

UGU cys

UGG trp

UAA stop

AAG lys

UUG leu

UAC tyr

GAG glu

UCG ser

UAU tyr

CGA arg

CAG gln

Ch rng, nu axit amin mi l cys th stop b ba b bin i l

UGA.
3. a. t bin b ba quy nh cys thnh stop b ba.
188

b.

t bin A thnh G b ba gln.

c.

Thm U vo b ba leu.
GXX UUA AGA CAA UGC CAU AUU

Ch t bin th 3 cho php bn xc nh trnh t baz ca mARN

bnh thng. V c nhiu axit amin b bin i, nn ta nn ngh ngay
n t bin dch khung bt u t axit amin th hai. Th t bin
l mt protein b ct nn chc chn qu trnh tng hp phi dng
li sm. Th t bin b ch c mt axit amin b thay i nn phi
l t bin nhm ngha.
Di y l cc b ba quy nh 3 axit amin u tin protein bnh thng:
GGX

UUA AGA
G
CUX

G
CGX

v protein t bin:

GGX UUC AAA

U

Nu b ba leu l UUA/G th xen U vo s sinh ra b ba UUU quy nh

phe. B ba leu khng th kt thc bng G v khng c b ba leu no
bt u bng G. V th b ba leu phi l UUA v b ba arg l AGA/G.
Phn trnh t cn li c xc nh bng cch vit ra mi b ba quy
nh mi axit amin v xc nh xem b ba no s cho ta trnh t
mong mun.
4. a. Polymer ny lun phin quy nh ser v leu. UCUCUCUCUC to ra
cc b ba UCU v CUC.
b. Polymer quy nh chui peptit phe (UUC), hoc ser (UCU), hoc
leu (CUU).
c. Polymer quy nh chui peptit gln (CAA), hoc asn (AAC), hoc thr
(ACA).
d. Polymerr quy nh chui peptit ile (AUA), hoc asp (GAU); UGA l
tn hiu kt thc.
e. Polymer quy nh chui peptit met (AUG), hoc asp (GAU); UGA l
tn hiu kt thc.
5. Ba.
Xt on trnh t 5 AGACUC 3 on gia si ARN. Hai b ba l
AGA v CUC. Nu A th hai i thnh C, ta c on trnh t 5
189

AGCCUC 3. Protein t bin s c ser-leu thay cho arg-leu. Nu m trm

nhau s c 3 b ba: AGA, GAC v ACU. Cn ARN t bin s c c l
AGC, GCC v CUU.
6. a.

met-gln-cys-asn-pro-ala

b.

met-gln-ser-asn-pro-ala

c.

met-

d. met-gln-leu-gln-ser-cys
e. met-gln-cys-n-pro-ala
Chng ta c mt cch n gin cc nhm gm 3 nucleotid trn ARN
theo chiu 5 n 3. t bin b ch lm thay i mt axit amin v l
t bin nhm ngha. t bin c lm ngng sm qu trnh tng hp.
Trong t bin d, C c xen vo b ba th ba lm thay i nhiu b
ba v axit amin. Ch rng t bin e l t bin im trn ARN
song n vn sinh ra protein bnh thng do s d tha ca m.
7. AGA. Trc ht hy vit ra cc b ba c th c, gm: AGA, AGG, CGX.
By gi hy vit cc b ba cho cc th t bin:
arg
ile

thr

ser

gly

AUU ACX AGU GGX

AUC

AGC

AUA

UCX

Hy tm baz duy nht b bin i gia cc b ba bnh thng v b

ba t bin. AGG khng th l b ba quy nh arg v cn phI thay
i 2 baz c c cc b ba quy nh ile. Tng t, phi thay i
2 baz bin b ba CGX thnh cc b ba quy nh thr v ile. Ch
cn b ba AGA v kt qu nh sau:
arg
AGA
Ile

thr

ser

AUA ACA AGU

gly
GGA

AGA
8. a.

gly, gly

b. 5 CGG 3 (arg); glycine c th c xen vo mt trong nhng b ba

ca arginine.
190

Ta vit ra mARN b tr vi b ba i m:
tARN bnh thng: 3 CCC 5

tARN t bin: 3 GCC 5

mARN bnh thng:5 GGG 3

mARN t bin: 5 CGG 3

axit amin bnh thng: gly

axit amin t bin: arg

Vic tARN gn mt axit amin vo chui peptit khng ph thuc vo b

ba i m, v th tARN b bin i vn s mang gly. N vn s nhn
ra b ba arg nhng i khi li gn gly vo v tr arg.
9. Khng th. Gi s mt tc nhn gy t bin gy ra t bin xen
mt baz. Cc b ba ng sau u b bin i v nhiu axit
amin cng b bin i theo. Mt ho cht gy nn t bin nhm
ngha hoc t bin im s ch lm thay i mt baz trong mt
b ba. S ch c mt axit amin b bin i v i khng th khi
phc li c chc nng ca protein.
10. Khng. Hy nh rng cc t bin xy ra trn ADN v ADN l mch
kp. Ta vit cc trnh t tng ng trn AND cho mi b ba stop:
UAA: 3 ATT 5

UAG: 3 ATC 5

5TAA 3

UGA: 3 ACT 5

5 TAG 3

5 TGA 3

Khng c C ng vi b ba UAA trn AND nn hydioxylamine khng gy

t bin b ba ny. Nu C trn AND ng vi UAG hoc UGA b t
bin thnh T th ARN s c b ba UAA - mt b ba stop khc.
11.5 AUG GCC UUU UCU CAC CGU GAG GUC CUC ... 3.
Hy bt u bng cch vit mi b ba quy nh mi axit amin. Chng
ta thy rng trnh t axit amin b thay i t v tr 26, v th t
bin xen hoc mt baz xy ra b ba ny.

Bnh thng

25

26

27

met

ala

phe

AUG
t bin

met
AUG

GCX

UUC/U

ser
AGU/C
UCX

leu
CUX
UUA/G
191

Mt mt baz b ba ala s khng sinh ra mt b ba ser no. Chng ta

cng khng th to ra b ba UCX bng t bin xen baz. Nu b ba
ala l GCX th vic xen thm A s sinh ra AGCXUUC/U. V th t
bin, axit amin 26 l leu, nn b ba quy nh n phi l CUX. V th,
b ba ala bnh thng s l GCC. So snh axit amin 27, phe v tyr, b ba
phe phi l UUU. Protein b ct ngn, nh th khung c phi dng li
sau b ba 30, cho thy baz th ba ca b ba 30 phi l U. Nh vy b
ba 30 l CGU. t bin c phc hi mt phn t axit amin 32 cho thy
b ba stop bin i thnh b ba bnh thng. khi phc li trnh
t bnh thng, baz c du sao phi b mt i: UGA* GGU CCU C
UGG GUC CUC ...

Chng 11: Li gii

1.
Chng
1

-lac
-

+lac
192

Chng 1 c gen z b hng, nn khng tng hp c z. Chng 2 khng

tng hp c ARN. Chng 3 to ra cht c ch khng hot ng chc
nng, v vy operon lun m. i vi chng 4-6, chng ta phi xem xt
tng ADN ring bit. Chng 4: ADN ng trc khng tng hp c ARN,
ADN ng sau khng sinh ra z. Chng 5: ADN ng trc khng tng hp
ARN, ADN ng sau khng sinh ra z. Chng 6: ADN ng trc lun sinh
ra z khng hot ng chc nng, v th cht c ch khng th gn vi
lactose. Cht c ch ny gn vi c hai ADN v thng xuyn ng ADN
ng sau.
2..b = gen -galactosidase, a = gen ch huy, c = gen c ch.
Trc ht hy xt t bin im khng sinh ra enzym; kiu gen ny s
gip ta xc nh gen cu trc. Vy trong kiu gen a+b-c+ th b l gen
cu trc, cn a, c l cc gen kim sot. Kiu gen 4 khng cho ta bit
iu g. Hy xt kiu gen 7: ADN ng sau khng sinh enzym. Nu a
l gen c ch th cht c ch ca ADN ng sau s phi gn c vo
promoter ca ADN ng trc v enzym ch c sinh ra khi c lactose.
iu khng xy ra, vy a phi l gen ch huy. Nu a l gen ch huy
th ADN th nht lun m v enzym ca n lun c sinh ra.
3. i-ocz-y+/i+o+z+y-.
to ra -galactosidase nh cm ng th mt ADN phi c gen ch
huy v gen z bnh thng. t nht, mt ADN phi to c cht c ch
bnh thng c th gn vi o+. Hy cho rng i+ cng trn mt ADN vi
o+, z+. ADN ny phi mang y-, nu khng, th vic sinh ra permease s
phi theo c ch chu cm ng. Ton b ADN ny s l i+o+z+y-. to
ra permease mt cch c nh, ADN kia phi l z-y+, v oc gen ch
huy khng c kh nng gn kt vi cht c ch. V ta c gen i bnh
thng trn ADN kia, nn gen i c th l i+ hoc i-. Gi s l i- th ADN
ny s l i-ocz-y+.

4. a.

i-o+z+ hoc i+ocz+

b. Trng hp 1: chu cm ng, trng hp 2: c nh.

operon lun m th cht c ch phi khng gn kt c vi gen
ch huy. iu xy ra khi cht c ch b hng (i-), hoc gen ch huy
193

khng gn kt c vi cht c ch (oc). Nh vy chng s c kiu gen l

i+ocz+ hoc i-o+z+. By gi, hy thit lp cc chng trng thi lng bi
mt phn: i+ocz+/i+o+z+ v i-o+z+/i+o+z+. Trong trng hp 1, cht c ch t
c hai ADN u khng th gn vi oc v qu trnh tng hp enzym l
c nh. Trong trng hp 2, cht c ch t ADN kiu di s gn vo c
hai gen ch huy v qu trnh tng hp enzym l chu cm ng.
5. a. P,Q = cc gen cu trc, R,S = cc gen iu ho.
b. Enzym 1 l chu cm ng trong c hai trng hp; enzym 2 l chu
cm ng
trong trng hp 1 v c nh trong trng hp 2.
V P- v Q- u khng to ra enzym, nn chng phi l cc gen cu
trc, v P = gen 2, Q = gen 1. V R- khng sinh ra enzym no, nn
chc l promoter.. V S- lun tng hp c enzym, c l do cht c ch
khng gn c vi gen ch huy, nn S = gen ch huy. Trong trng hp
1, ADN ng trc khng sinh ra enzym 1 (Q-), nhng s sinh ra enzym 2
bng cch cm ng. ADN ng sau khng sinh ra enzym 2 (P-), nhng
sinh ra enzym 1 bng cch cm ng. Trong trng hp 2, ADN ng sau
khng sinh ra enzym 1 nhng sinh ra enzym 2 mt cch c nh. ADN
ng trc khng sinh ra enzym 2, nhng sinh ra enzym 1 bng cch cm
ng.
6. Hu ht cc bn sao ARN c di y ; c rt t bn phin m
d.
to ra ARN ngn, phin m d th tARN trp mang tryptophan phi c
kh nng gn kt vi cc b ba trp vng suy gim. V UGG l b ba
quy nh tryptophan, nn tARNtrp mang tryptophan s khng nhn bit
c AGG. Khi tryptophan bt u c to ra, n s gn vi cht c ch,
hot ho n n c th gn vo gen ch huy.
7. C th c hai gen tng cng, mt nm pha trc v mt nm pha
sau gen.
S gim s lng mARN to ra cho thy trnh t on gen tng cng
b bin i. Cc gen tng cng c tm thy c hai u ca gen v
c th tc ng trn khong cch ln.
8. a. Khng sinh ra enzym; cis dominant.
b. Lun sinh ra enzym; nu sai hng gen c ch s khng phi l cis
dominant, nhng sai hng gen ch huy s sinh ra cis dominant.
c. Lun sinh ra enzym; khng phi l cis dominant.
phin m xy ra, promoter phi hot ng chc nng. Sai hng
promoter s ch nh hng n on ADN m n nh v trn .
(b), hoc gen c ch hoc gen ch huy b sai hng. Gen c ch b sai
hng c th c khc phc nh cht c ch t ADN khc (tc ng
lch - trans acting). gn c vo gen ch huy, u tin cht c ch
phi gn vi mt cht ng c ch.
194

9. Gen ny trn ADN ca gan c bn intron; cc intron b ct b khi

tng hp mARN. V th n s khng lai vi ADN. Qu trnh ct b
ADN tu xng c khc. Intron th hai v mt phn ca intron th 3
gan vn c mt ARN tu xng.
10. Gen ch huy nm gia B v C; promoter nm gia D v E.
t bin promoter s khng sinh ra ARN, v l kt qu chng 4
v 5. Chng cng b mt on vng gia D v E. t bin gen ch huy
thng gy cho gen khng gn c vi cht c ch v vic sn sinh
enzym l c nh. Chng 2 v 3 cho thy hin tng ny v c hai
chng u b mt vng B. Nh vy vng B l gen ch huy.
11. Gen A: c phin m giai on ng thn kinh v tu xng. Gen B: c phin m t giai on trng th tinh n phi v v gan sinh vt trng thnh. Vic khng pht hin c gen sau khi x l ADNase l do
chng hot ng phin m. Nhng giai on c du tr l gen hot
ng phin m.
12. Khng c hin tng loi b intron. Nu chng b loi b s phi xut
hin cc vng. S loi b cc intron l cn thit c th dch m
hon chnh.
13. Tch mARN insulin t ngi bnh thng v cho lai vi ADN t ngi b
bnh i ng. Nu promoter sai hng th gen vn tn ti v s lai vi
ARN. Nu gen b mt, s khng c hin tng lai.

195

Chng 12: Li gii

1. B c nhiu cp G-C. V mi cp G-C c ba lin kt hydrro, nn n cn

nhiu nng lng hn b ph hu
2. Mch n. mch kp A = T v G = C. T l ny khng gp phn t
ny.
3. 1: ADN mch kp; 2: ADN mch n; 3: ARN mch n; 4: ARN mch
kp. Trc ht, s c mt ca T hoc U xc nh n l ADN hay ARN.
Nu A = T(hoc U) v G = C xc nh n l mch n hay kp. Cc
phn t 1 v 4 l mch kp.
4. a. 5 TTGGTCGATGTAATCATCGCC 3
b. 5 GGCGAUGAUUACAUCGACCAA 3
c. met-ile-thr--ser
Hy c trnh t t di ln, v nh rng u 5 l u di. Nu bng
xut hin c hai ct C v C+T, th l C. Nu bng xut hin c
hai ct A v A+G, th l A. Hy nh rng ARN c phin m theo
chiu 5 n 3 v cc axit nucleic ngc chiu. xc nh trnh t
axit amin, hy tm b ba u tin theo chiu 5 n 3 ri ln theo
trnh t cc b ba.
5. Cc on di 1, 4, 8 v 12 baz.
Phng php dideoxy da trn nguyn tc b tr; cc on s c to ra
ni ddC gn vo, bt u t u 5ca si mi c tng hp.
6.

4,3
4,3
5,1

5,1

6,6

4,3

6,6

5,1
6,6

196

C ba on, vy phi c hai im gii hn. Bt k on no cng c

th l on nm gia, v trt t cng c th thay i. V d nh trt
t sau:
6,6

5,1

4,3

cng ng.
7.

a. 4,9; 1,2.
b. 2,1; 0,6; 2,2; 1,2
Nu im gii hn b mt th ADN khng b ct ti im v s
xut hin on di hn. t bin sinh ra im gii hn mi s lm mt
i mt on c v sinh ra hai on mi nh hn on b mt.

8. a. C.
c. Khng.
V bnh ma git Huntington di truyn tri, nn c nhiu kh nng
ngi bnh l d hp t, v v vy s to ra cc on c t gen bnh thng v gen b t bin. Ngi chng c gen bnh thng. Khi so snh vi
b ca ngi ph n chng ta thy ngi b c hai bng mi c l l do
s xut hin mt im gii hn mi ti on u ca gen bnh thng.
Bng th ba v bng th nm cha ADN ca c gen bnh thng v gen
t bin. Ngi ph n ging vi b, v vy b l d hp t. Thai nhi
bnh thng. Vy ta c th v ADN cho b b nh sau:

bnh thng

t bin

9. (b) v (c).
Hy bt u bng enzym B. C th c ba kiu trt t: 1,5; 4,0; 3,5,
hoc 1,5; 3,5; 4,0, hoc 3,5; 1,5; 4,0. on 1,1 v 2,7 xut hin c khi
x l A v hn hp hai enzym, vy cc on ny khng cha im gii
hn cho B. V khng c on no ca B xut hin khi x l hn hp
enzym, nn mi on ca B u c cha im gii hn cho A. Gi s
trt t (b) l ng. Trt t ny phi sinh ra on 2,0 v 1,5 t pha
phi ca phn t. Ta khng thy cc on , vy trt t (b) l sai. Gi
s (c) ng. Trt t ny phi sinh ra cc on 1,1 v 0,3 t u tri v
0,8; 2,7 t u phi. Ta cng khng gp kt qu . Vy, bn (a)
l ng.
10.a. Mch vng.
b.

E
2,7

3,5
197

3,5

4,3

B
B

5,5

Phn t ny, hoc l mch vng, hoc khng c im gii hn cho

EcoRI. Nu khng c th chng ta phi gp nhng on ging nhau
khi x l bng BgIII v bng hn hp hai enzym. Do vy, on 6,2 kb
phi cha im gii hn ca EcoRI. V tr cc on 3,5; 2,7 v 4,3; 5,5
c th c trt t ngc li.
11.1,5 v 2,7 kb.
on gen xen vo c th b loi b khi plasmid nh EcoRI to ra
on 4,2 kb. V kt qu x l bng hai enzym khng sinh ra on ny
nn n phi cha im gii hn cho HindIII. By gi hy xem xt cc
on cng vo ta c on 4,2 kb; l cc on 1,5 v 2,7 kb.
12.a. Cc phn t 6 v 14 kb l cc dimer (lng phn) ca gen v ca
plasmid.
b. 10 kb.
Khi trn cc phn t vi nhau v cho chng gn li, ngoi nhng phn
t ban u cn c th to ra cc dimer. Cc phn t 3 v 7 kb l cc
n phn ca gen v plasmid, cn 14 v 6 kb l cc dimer.
13.a.

198

V phn t lai duy tr tnh khng vi tt c cc cht khng sinh khi

AND xen vo im gii hn ca EcoRI, nn im ny khng nm gia
gen no, n phi nm u bn ngoi cc gen. Nu tnh khng b
mt th im xen nm gia gen. V hn hp EcoRi + BglII, hoc EcoRI
+ SmaI sinh ra t on hn so vi hn hp EcoRI + PstI, nn im gii
hn ca EcoRI phi nm gia tetracycline v chloramphenicol. Cc v tr
cn li c th c xc nh bng cch cng cc on.

14.

H
3,0
1,8

4,0

2,7

5,0

5,3

2,2

Nh rng phn t AND s c nh du cc u, v n l si kp. Kt

qu x l tng loi enzym cho ta kt qu sau:
HindIII

3,0*

BglII

4.0

5,0*

(2,2; 5,3)

1,8*

2,2*

hoc
(2,2; 5,3)

2,2*

1,8*

Nu trnh t th hai theo BglII l ng th vic x l bng hn hp

cc enzym phi sinh ra on 0,8 kb. Ta khng thy on . Vy ta c:
HindIII:
BglII:

3,0*
1,8*

4,0

(2,2; 5,3)

5,0*
2,2*

Nu on 5,3 nm ngay sau on 0,8 th chng ta s phi gp cc

on 0,1 v 3,1. Khng c cc on . Nu on 2,7 nm sau on
0,8, chng ta s phi gp on 1,5 kb.
15.ADN phi b mt im gii hn v hnh thnh nhng im mi trong
qu trnh pht trin. cADN s gn vi nhng on c cha cc trnh t
ca gen quy nh khng th. V c nhng on mi xut hin, nn cc

199

im gii hn c b mt v nhng im gii hn mi c sinh ra, c

l, do s dch chuyn cc vng ADN.

Chng 13 : Li gii

1. M = 0.34; N = 0,57. Qun th khng cn bng.

Tn s alen:
M = [300 + 1/2(180)]/900 = 300/900 = 0,43
N = 1 - 0,43 = 0,57
Nu qun th trng thi cn bng, ta c tn s kiu gen MM =
(0,43)2 = 0,18, MN = 2(0,43)(0,57) = 0,49 v NN = (0,57)2 = 0,33. Cc
tn s quan st c l : MM = 300/900 = 0,33; MN = 180/900 = 0,20;
NN = 420/900 = 0,47.
2. 0,18 MM, 0,49 MN, 0,33 NN.
Khi khng c cc lc bn ngoi tc ng, trng thi cn bng s t c
sau mt th h. Khi , tn s cc kiu gen s ng nh tnh c
bi 1.
3. IA = 0,05, IB = 0,15 v IO = 0,8.

200

V ta c ba alen nn tn s cc kiu gen c tnh theo cng thc (p +

q + r)2 = 1. Trong cng thc ny, ta c r2 = tn s cc c th c nhm
mu O = 128/200. Vy, r = 0,8. Gi p = tn s alen IA, ta c (p + r)2 =
tn s nhm mu A + tn s nhm mu O = 0,085 + 0,64 = 0,725. V
(p + 0,8)2 = 0,725, nn p = 0,05. V p + q + r = 1, v p + r = 0,85, nn
q = tn s alen IB = 0,15.
4. 0,25.
Trc ht hy tnh tn s alen:
M = [69 + 1/2(108)]/300 = 123/300 = 0,41
N = 1 - 0,41 = 0,59
By gi hy tnh tn s d hp t quan st c:
108/300 = 0,36
v tn s d hp t theo l thuyt:
2pq = 2(0,41)(0,59) = 0,48
H s ni phi:
f = 1 - (tn s quan st/tn s l thuyt) = 1 - (0,36/0,48) = 0,25
5. a. p = 0,2; q = 0,8.
b.p = 0,08; q = 0,92.
Theo u bi, ta c phng trnh:
q2= 2(2pq) = 4pq
Thay p = 1 q, ta c:
q2 = 4q(1 q) = 4q 4q2
0 = 4q 5q2
0 = q(4 5q)
Vy q = 0 hoc 5q = 4
q khng th bng 0, v nu vy s khng c cc ng hp t ln. Vy
q = 0,8. Kim tra li bng cch thay th vo phng trnh Hardy
Weinberg:
q2 = 0,64, 2pq = 2(0,2)(0,8) = 0,32
L lun tng t gii phn b:
q2 = 6(2pq) = 12pq = 12q 12q2
0 = q(12-13q). vy q = 0 hoc = 12/13 = 0,92.
Kim tra li:
q2 = (0,92)2 = 0,846; 2pq = 2(00,8)(0,92) = 0,14
6. a. 0,007
201

b.1,4
V bnh x nang l tnh trng ln, vy tn s alen ln (q) l:
tn s kiu hnh ln =

1 / 20000

= 0,007

v p = 1 q = 0,993
Tn s cc th mang theo l thuyt bnh l:
2pq = 2(0,093)(0,007) = 0,14
7. a. p = 0,92; q = 0,08
b. 0,92
V ph n cn hai alen ln mi biu hin kiu hnh, nn t l ph n
mang bnh l q2. Vy q = 64 / 10000 = 0,08. P = 1 q = 0,92. V nam
gii ch c mt nhim sc th X, vy tn s l thuyt ngi n
ng mc bnh m mu l q, hoc 0,08. 92 % nam gii bnh thng.
8. a. 0,01
b. 0,198 hoc khong 2%
Tn s alen ln chnh l t l n ng mc bnh, bng 0,01. Vy p =
1 q = 0,99. Tn s d hp t : 2pq = 2(0,01)(0,99) = 0,0198.
9. 0,704 A1A1; 0,192 A1A2; 0,104 A2A2.
A1A1 = p2 + fpq = (0,8)2 + (0,4)(0,8)(0,2) m= 0,704
A2A2 = q2 + fpq = (0,2)2 + (0,4)(0,8)(0,2) = 0,104
A1A2 = 2pq 2fpq = 2(0,8)(0,2) 2(0,4)(0,8)(0,2) = 0,32 0,128 =
0,192
10. 0,48 A; 0,15 B; 0,36 AB; 0,01 O.
V qun th trng thi cn bng, nn tn s cc kiu gen c tnh:
(p + q + r)2 = p2 + 2pq + q2 + 2qr + r2
p = 0,6; q = 0,3 v r = 0,1, vy tn s cc nhm mu l:
A: p2 + 2pr

B: q2 + 2qr

AB: 2pq

O: r2

= 0,36 + 0,12

= 0,09 + 0,06

= 0,36

= 0,01

11.0,375.
Trc ht, hy tnh tn s d hp t theo l thuyt:
2pq = 2(0,8)(0,2) = 0,32
H s ni phi:
1 (s liu quan st / s liu l thuyt) = 1 (0,2/0,32) = 1 0,625 =
0,375
12.Phn bc. Tn s ht vng s ph thuc vo tn s alen ca qun th
ban u. Nu chng ta gi s rng s giao phi l ngu nhin v khng
c tc ng ca chn lc, chng ta c th tnh c tn s ht vng cho
202

hai qun th trng thi cn bng. Cho p = tn s alen quy nh ht

vng v q = tn s alen ht xanh.
Qun th
2pq)

Tn s cy ht vng (p2 +

0,7

0,3

0,91

0,5

0,5

0,75

Chng ta thy gi thuyt ban u ch ng nu khi u cc alen

c tn s bng nhau.
13.CB = 0,273; CP = 0,467; CY = 0,26.
Trc ht phi xc nh kiu gen ca mi mu:
Nu:

CBCB, CBCP, CBCY

Hng: CPCP, CPCY

Vng: CYCY
Cho p = tn s alen CB, q = tn s alen CP v r = tn s alen CY.
Tnh tn s mi mu:
Nu: 472/1000 = 0,472
Hng: 462/1000 = 0,462
Vng: 66/1000 = 0,066
Tn s mu vng l r2, vy r =

0,066

= 0,26

Tn s vng + hng = (q + r)2 = 0,528

q+r=

0,528

= 0,727

q = 0,727 r = 0,727 0,26 = 0,467

V q + p + r = 1, vy p + 0,727 = 1, v p = 1 0,727 = 0,273
14.a. M = 0,652, N = 0,348
b. Khng.
c. f = 0,267.
Tn s M:
0,486 + 1/2(0,332) = 0,652
Tn s N:
0,182 + 1/2(0,332) = 0,348
Tn s d hp t theo l thuyt:
2pq = 2(0,652)(0,348) = 0,453
203

V tn s d hp t quan st c gim so vi l thuyt, vy qun th

khng trng thi cn bng. Tn s kiu gen NN theo l thuyt l:
(0,348)2 = 0,121. Ta thy s ng hp t NN tng ln, vy c hin tng ni phi. H s ni phi l:
1 (0,332/0,454) = 0,269

Chng 14: Li gii

1. 3000. V ban u khng c t bo khng, nn khng c t bin

nghch. Vy tn s cc t bo khng bng p = (1011)(3 x 10-8) = 3 x
103.
2. a. p = 0,67997, = 0,32003
b. p^ = 0,05, q^ = 0,95
c. 0,02
204

V q = 0,32, p = 0,68 nn mc thay i tn s alen q sau 1 th

h l:

q = p - q = (2 x 10-5)(6,8 x 10-1) - (10-6)(3,2 x 10-1) =

(136 x 10-7) (3,2 x 10-7) = 132,8 x 10-7 = 1,33 x 10-5
V q1 = 0,3203 v p1 = 0,67997, vy tn s cn bng l:
q^ = /( + ) = (2 x 10-5)/(2,1 x 10-5) = 0,95
p^ = (1 x 10-6) /(2,1 x 10-5) = 0,05
V c t bin v chn lc cng tc ng, nn
q^ =

/s =

(2 x 10-5)/10-1 =

2 x 10-4 = 2 x 10-2 = 0,02

Nh rng chn lc = 1 - ph hp.

3. 0,83; 0,91; 0,67.
V tc nhn duy nht lm thay i tn s alen l t bin, nn chng
ta dng cng thc:
q^ = /( + )
Trong trng hp 1,

= 5, vy q^ = 5/(5 + ) = 5/6 = 0,83

Trong trng hp 2,

= 10, vy q^ = 10/(10 + ) = 10/11 = 0,91

Trong trng hp 3,

= 2, vy q^ = 2/(2 + ) = 2/3 = 0,67

4. 0,219.
Chng ta s dng biu thc m = (q - q)/(q - q m), vi q = tn s qun
th gc; q = tn s qun th hn hp; qm = tn s qun th nhp
c. Thay vo, ta c:
m = (0 - 0,094)/( 0 - 0,429) = 0,219
5. 0,32.
Trc ht chng ta tnh tn s alen trong mi qun th. Nh rng
q = tn s ngi mt xanh. qm =
S dng cng thc:

0,16

= 0,04, v q =

0,09

= 0,03.

q = q - m(q - qm) = 0,3 - 0,2(0,3 - 0,4) = 0,3 + 0,02 = 0,032

6. 0,478 AA, 0,378 Aa, 0,144 aa; p = 0,67; q = 0,33.
Sau mt th h ngu phi, tn s cc kiu gen s bng tch tn s ban
u vi ph hp. Vy:
AA = 1(0,33) = 0,33
205

Aa = (0,8)(0,33) = 0,26
aa = (0,3)(0,33) = 0,1
Tn s tng ng ca cc kiu gen trong qun th mi s l:
AA = 0,33/0,69 = 0,478
Aa = 0,26/0,69 = 0,378
aa = 0,1/0,69 = 0,144
V p = tn s AA + 1/2 tn s Aa, vy p = 0,478 + 1/2(0,378) = 0,67;
v q = 1 - p = 1 - 0,67 = 0,33.
7. 0,58. Cho s1 = h s chn lc ca TfA/TfA v s2 = h s chn lc ca
TfB/TfB. Vy,
q^(TfB) = 0,31/(0,31 x 0,22) = 0,31/0,53 = 0,58
hu ht cc qun th chim b cu, tn s TfB bin ng t 0,55
n 0,59.
8. 0,174, 0,154. V chng ta quan tm n tn s alen th h th
nht v th hai sau chn lc, nn chng ta dng biu thc:
qs = q(1-q)/(1 - q2)
Sau mt th h,
q1 = 0,2[1 - (0,2)(0,8)]/[1 - (0,8)(0,04)]
= 0,2 (1 - 0,16)/(1 - 0,032)
= (0,2) (0,84)/0,968
= 0,174
th h th hai, tn s xut pht l 0,174. D dng vit li cng thc
trn thnh
qs = (q - sq2)/(1 - sq2)
= [0,174 - (0,8)(0,174)2]/[1 - (0,8)
(0,174) ]
2

= 0,150/0,97 = 0,154
9. 0,528.
Cho qm = 0,25, q= 0,5 v m = 0,1. Theo phng trnh
m = (q - q)/(q - qm)
Bin i, ta c
m(q - qm) = q - q hoc q = mq - mqm + q
q - mq = q - mqm, hoc q (1 - m) = q - mqm
q = (q - mqm)/(1 - m)
= [0,5 - (0,1)(0,25)]/0,9
206

= (0,5 - 0,0250/0,9
= 0,475/0,9
= 0,528
10. 0,012.
V c t bin v chn lc cng tc ng, nn
q^ =

/s

. Bit = 3 x 10-5 v s = 0,2, vy

q^ = (3 x 10-5)/0,2 = 1,5 x 10-4

= 1,22 x 10-2 = 0,12

11. 6 x 10 5.
V chng ta c t bin tri nn tn s ca n trng thi cn bng
c tnh bng: tc t bin/h s chn lc. V ph hp = 0,25,
s = 0,75 v = 4,5 x 10-4, nn
q^ = 4,5 x 10-5/0,75 = 0,6 x 10-4
12. 0,476 B1B1; 0,428 B1B2; 0,096 B2B2.
Chng ta s dng cng thc:
qs = (q sq2)/(1 sq2)
= [(0,4 - (0,8)(0,16)]/[(1 - (0,8)(0,16)]
= (0,4 - 0,128)/(1 - 0,128)
= 0,272/0,872
= 0,31
v p = 0,69
Cc kiu gen s c phn b nh sau:
p2 B1B1 = (0,69)2 = 0,476
2pq B1B2 = 2 (0,31) (0,69) = 0,428
q2 B2B2 = (0,31)2 = 0,096
13.Tn s ca r = 0,337, tn s ca s = 0,663.
Trc ht hy tnh h s chn lc bng 1 - ph hp. s1 = h s chn
lc ca ng hp t c kiu hnh bnh thng = 1 - 0,68 = 0,32, v s2
= h s chn lc ca t bin ng hp t = 1 - 0,37 = 0,63. Vy:
q^ = 0,32/(0,32 + 0,63) = 0,32/0,95 = 0,337
V p + q = 1, p^ = 1 - 0,337.
14.1,4 x 10-6.
V c rt t c th b tc ng nn c th coi tn s alen ln, quy
nh kiu hnh bnh thng l 1 v ng hp t tri khng tn ti. V
th cc c th b tc ng u l d hp t, hay 2pq = 4 x 10-6. V q

207

= 1 v 2p = 4 x 10-6 nn p = 2 x 10-6. Chng ta gi s l gi tr

trng thi cn bng, th p^ = /s, hay (p^)(s) = .
(2 x 10-6)(0,7) = 1,4 x 10-6
15.0,778.
Trc ht hy xc nh xem qun th c trng thi cn bng khng
bng cch tnh tn s alen v tn s cc kiu gen.
q = 0,000064 + 1/2(0,015872) = 0,008 v p = 0,992.
Theo l thuyt, tn s aa = (8 x 10-3)2 = 0,000064 v AA = 90,992)2 =
0,984064. Vy qun th ny trng thi cn bng, v
q^2 = /s, hoc s = /q^2 = 5 x 10-6/64 x 10-6 = 0,078
16.0,0099.
V ng hp t cht, nn s = 1. Vy
q1 = q/(1 + q) = 0,01/1,01 = 0,0099
i vi th h hai, chng ta bt u vi q = 0,0099, vy q2 =
0,0099/1,0099 = 0,0098. Tnh hung ny c trng cho nhiu ri lon
ngi do gen ln quy nh, nh bnh Tay-Sachs hoc x nang.

208

Chng 15: Li gii

1. a. 3 ; 3 son : 1 nu : 1 trng
b. 9 : 3 son : 3 nu : 1 trng
c. 2 : 1 son : 1 nu
d. 2 : 1 d son ; 1 nu
Hy vit s lai t u n F1 X F1 cho mi php lai. Nh rng
Drosophila c khng c trao i cho, v cc gen cch xa nhau t 50
n v bn tr ln s phn ly ging nh cc gen khng lin kt.
2. A-2; B-1; C-4; D-3.
Mt trng c th l kiu gen Xw v bt k mt gen no khc, hoc
cncn, sese. Php lai 2 phi l
se/se; ++ X ++; cn/cn
Php lai ny s cho ra t l 9:3:3:1. Php lai 2 ph hp vi A. Trong php
lai 1, rui c mu nu phi c kiu gen X+Y; se/se. Nu rui ci
trong php lai ny l XwXw; +/+, th php lai cc c th F1 s l
X+Xw; +/se X XwY; +/se
V cho kt qu:
3 X+-; +-

3 Xw-; +-

trng

1 X+-; se/se

nu

1 Xw-; se/se

trng

Nhng kt qu phn ly khc nhau theo gii tnh C cho thy c s di

truyn lin quan n gii tnh. Rui ci F2 c t l phn ly ging vi hai
gen trn nhim sc th thng, vy n phI l X+. Php lai khi u
phi l:
X+X+; cn/cn; se/se X X+Y; +/+; +/+
D, chng ta gp t l cho thy F1c mt gen trn nhim sc th
thng trng thi d hp t. Rui c phi l cn/cn; se/se, v rui ci l
cn/cn +/+.
3. a. Ton b lng qun.
b. Ton b lng qun.
ng hp t khng lng gy cht v tnh trng khng lng phi l tnh
trng tri. Chng ta c ba alen vi trt t tri ln nh sau: khng lng >
lng qun > lng thng.
4. a. 9/48
209

b. 1/4
sinh ra trng vng, c th phi c kiu gen +/-, +/- v fe/fe cho cc
gen trng thic, thu ngn v st. V vy, ta c
3/4 x 3/4 x 1/3 = 9/48
(T l 1/3 trn y l v ch c 3/4 s c th mang gen v st sng
st v 2/3 trong s l Fe/fe). cu (b), 1/4 s c th sng st s c
kiu gen sn/sn v trng thic.
5.

32,2

ct

4,5

fa

>50

ba
Gen cnh trn c th u ny hoc u kia. Php lai A cho thy c
bn gen u l gen ln. Php lai B cho thy bn gen ny lin kt. Nh
rng khng c trao i cho Drosophila c. Trong php lai C,
chng ta thy c bn lp kiu hnh chnh, chng t mt gen phn ly
c lp vi cc gen khc. Xt hai gen j v fa, chng ta ch thy c hai
lp kiu hnh, vy j v fa lin kt. Xt fa v ba, ta thy bn lp kiu
hnh, vy gen ba cch cc gen khc >50 n v bn . B qua gen
ba, chng ta c th lp bn cho cc gen j, fa v ct.
6. Ton b rui ci kiu di.
Rui c:

397
78
23
2

m + scr
m++
+ + scr
+++

Rui ci d hp t c kiu gen m + scr/+ l +. Hy tnh tn s trao i

cho kp (DCO) quan st c:
(0,5)(0,05)(0,16) = 0,004
By gi, hy tnh tn s l thuyt cho ton b tm kiu hnh. Cc lp
kiu hnh c l s cht.
7. a. Gen ln trn nhim sc th thng.
b. Gen ln trn nhim sc th thng.
c.Hai gen khc nhau.
V hai ngi bnh thng sinh ra con t bin nn tnh trng l tnh
trng ln. Nu l tnh trng lin kt X th ch 1/2 s con trai b
bnh. Hai ngi mc bnh sinh ra ton b con bnh thng cho thy
c hai gen.
8. a. lys+ his+ val+
lys+ his+ vallys+ his- val+
lys+ his- val210

Kiu gen no c lys+ mi mc c.

b. lys+ val+ hislys+ val+ hisKiu gen no c lys+ v val+ mi mc c.
c. lys+ val+ his+
lys+ val- hisKiu gen no c lys+ v his+ mi mc c.
d. lys+ val+ hislys+ val- his+
Chng ta khng thy c kiu gen lys+ val+ his+.
e. lys+ v val+ nm gn nhau; chng ng bin np 75% thi gian. Trt
t gen c th l lys val his hoc val his lys.
f. val lys his. Nu trt t gen l val lys his th kiu gen val+ lys- his+
phi him gp v n c to ra do trao i cho kp.
9.
-lac
A

+lac

( xem chng 11)

10. Q = operator v T = promoter.
t bin vng operator lm cho gen lun hot ng, v vy mt on
1 ph vng Q phi cha operator. t bin promoter, hoc sai hng
mt trong hai gen cu trc s khng to ra t. V vy mt on 2, 3 v
4 (ng vi cc vng R, S v T) phi l promoter. Trong s cc th lng bi,
nu R = promoter th chng 2 phi c trnh t vng gen kim sot
bnh thng, v do chng chu cm ng, v S v T phi b tr cho
nhau. Do vy R khng phi l promoter. chng 3, nu S = promoter
th ADN th nht khng hot ng v ADN th hai chu cm ng nhng
to ra R sai hng. T phi l promoter.
211

11.e = z; y = i; u = o; r = p.
Cc chng mang t bin p hoc z u khng cho hot (e, r). t
bin operator v gen c ch vn lun cho hot mc no
(u, y). z-/z- v z-/p- u khng cho hot , v th chng ta thc s
khng th phn bit c cc t bin z- v p- t nhng s liu cho.
t bin operator lun cho t nht l 1000 n v hot . V vy u
= o- v y = i-.

12.

Chng ta khng th xy dng c bn thng ph hp vi s liu. V

vy hy c xy dng bn vng trn.
13.

0,04

0,02

Cc vt tan trn B cho ta bit tng s cc khun lc i con. Cc vt

tan trn K12 l cc th ti t hp kiu di v ch c kiu di mi mc c trn K12. Tuy nhin s lng ny ch l 1/2 s th ti t hp, 1/2 cn li
l t bin kp (xem chng 9).
14.a. UAA
b.Olmstead: CUG thnh CGG; House: UAU thnh UAA hoc UAG;
Max: CUGGCC thnh CUG G GCC (xem chng 10).
15.a. Si n; v A khng bng T v G khng bng C.
b. Cc phn t si n khng bin tnh.
c. C l phn t ADN ny c mt s vng b sung nhau:

212