A

2.1 . R

Cc. F2 -

1 CC : 2 Cc,

. -

RR rr Rr. F1
.

. F2

F1. Rr Rr 3/4
R : 1/4 rr. 3/4 F2 1/4
.

. Rr

RR R .
. Rr rr 1/2 Rr : 1/2 rr.
, 1/2
1/2 .
2.3 , F2
3/4 : 1/4 . , F2
3 : 1 . C c
. F2
1 CC : 2 Cc : 1 cc.
, CC Cc. CC,
. Cc,

. -

Cc 2/3.
2.4 . Rr ( ) rr ( ). F1 1 Rr (
) : 1 rr ( ).
. Rr Rr, 1/4 RR : 1/2 Rr : 1/4 rr 3/4
(R ) : 1/4 (rr ).
2.6
( ) p
( ). 3 : 1 ,

(Pp).
: PP
Pp. 2 Pp : 1 PP.
, 1/3
.
2.7 .
b . Bb,
bb.
2.10

81
118
74
90

82
39
0
0

Gg
Gg
GG
GG Gg (G)

2.11 -

[P = 1/4 (bb) 1/2 ()

, -

= 1/8] (-

Bb. , -

). -

1 : 2 Bb.

Bb bb,

. (bb)

1 -

(bb) : 1 (Bb). -

, -

, -

Bb.

Bb F1 2/3.
bb

Bb), -

( bb

Bb 1/4.

2.14 -

(
) = (
F1
Bb) (
bb )
= (2/3 2/3) (1/4)
= 1/9
.
F1 (Bb )
(Bb), .
: (1) Bb ( F1) b (
), = 2/3 1 = 2/3 (2)
( F1) b ( ), = 1/3 1 = 1/3. 1/4. , :
= 2/3 ( Bb Bb)
1/4 ( bb ) = 1/6.
.
,
. -

.

, ,
, ,
. S s . , S ss

Ss.

Ss ,
3 (S ) : 1
(ss).
Ss ss 1 Ss ( ) : 1 ss
( ).
.
(s) (S ) .
(ss)
-

, Ss ss
. ,
(ss). ,
.
2.15 . Ww Dd ww dd

9/64 agouti,
,
9/64
,
3/64 agouti
,

. Ww dd ww dd

3/64

. Ww Dd ww dd

3/64

. ww DD WW dd
. Ww Dd Ww dd
2.16
a Bb Cc a

Bb Cc, .
. a a,
3/4. ,
Bb Bb 3/4
C Cc Cc 3/4.
,
(A B C ) 3/4 3/4 3/4 = 27/64.
.
a a 1/4.

CC b Bb
Cc Cc .
,
A B CC 1/4 1/4 1/4 = 1/64.
2.20 . aa BB CC bb
c hc h. F1 a Bb Cc h
agouti .
,
F2 :
27/64
agouti, ,
9/64 agouti
,

, agouti,
,
1/64
, agouti.
. F2
agouti, A B C . , 2/3 a. , 1/3 .
C , 2/3 Cc h,
a BB Cch 2/3 1/3 2/3 = 4/27.

. a Bb Cc h a Bb

Cc h, 1/4 bb
.

. , 1/4 .
. a Bb Cc h a Bb
Cc h, 3/4
. agouti. , 3/4
agouti .
2.24 C
aa bb. ,
C aa bb.

C
. ,
a Bb, aa Bb D a bb.
2.25 . Ww Rr W r.

(1/4 W R, 1/4 W r, 1/4 w R, 1/4 w r ),

, a (II.1, II.3, II.4)

( W r ). -

1/2 W Rr ( -

. a aa

, ) 1/2 W rr ( ,

1/2 a ()

).

1/2 aa ( -

. -

). ,

. , -

1/4 W R ( , ),

1/4 W r ( , ), 1/4 w R

2.29 . -

( , ), 1/4 w r ( -

, ).

Gaucher.

Cc ( ).

. , -

, -

. ,

C. , -

Cc C. Cc 1/2. Cc
C,
Cc 1/2. , 1/2 -

1/2 1/2 = 1/4.

. F4 (1/2)4 = 1/16.

, -

2.26 .

IV.5,

a, .
. aa,
.
. a ()
aa (),
a .
a
, aa (II.2,
II.5) .

IV.4,
IV.6
.7. .7 ,
,

.
. IV.5 IV.6
(V.1), a

IV.5 IV.6 , -

a,

, -

a .

IV.5 , -

. Neurospora

a. -

crassa Saccharomyces cerevisiae

.
. ,

.
3.8 . ,
,
1.
3.10 . : , .
. : ,
.
3.14 .
1/2. ,
(1/2)3 = 1/8.
.

,
. P (
) = 1 P (
).
()
1/8. , P (
) = 1 (1/8 + 1/8) = 3/4.
3.15 , -

IV.5 a 2/3. ,
IV.5 a V.1 2/3 1/2 = 1/3.

( IV.6 a) = ( .3 a

a
V.6)
= [( .6 a

a
.3) ( .3

a
IV.6)]
= [(2/3 1/2) 1/2)] = 1/6.
1/6 1/2 = 1/12
IV.6
a V.1. ,
a
1/12 1/3 = 1/36.
, IV.6

V.1 .

3
3.1 .
3.3 .
3.6 . ,

. -

(c +c). O

, c +Y. -

c +c x c +Y. -

c +

() .

3.17 . -

(cY)

(c +Y).

.
.
3.19 . 17 + 26 = 43 .
.
.
, . ,
. . ,
. ,
,
.

, .
3.20
1/2.
,
(1/2)5 = 1/32.
3.23
,

3.25 . ww vg +vg +

w +Y vgvg. F1 w Y vg +vg
( , ) w +w
vg +vg ( , ).
.
, F2 3/8
, 3/8
, 1/8 1/8 .
. F1
, ww vg +vg +
w Y vg +vg.
. F1
, w +w
vg +vg w +Y vgvg. : 1/4
, 1/4 , 1/4
1/4
.
,
.
3.28 ( ) .
, .
b .

bb,

. ,

. , -

50%

(.., BB, bb) 1/4

(.., Bb)

. -

1/2. /a, -

, -

a AY, -

3.30 . -

1/4 -

1/2.

, -

, -

(). c -

(1) bb CC dd () P =

c -

, c c c c
+

1/4 cc. , 1/4.

. 1

c c : 2 c +c, 2/3
.
3.32 . ,
: aY. a
,
F1 1.
. () (P = 0)
a F1.
. F1 b Cc Dd.
, P = 1/2.
. , a b Cc Dd
b Cc Dd.
. , F1 Aa.
(1/4
) a. , 1/2
(..,
Bb Bb 1/2 b).
, F2
1/4 1/2 1/2 1/2 = 1/32.
+ +

(1/2 1/4 1/4 1/4) = 1/128, (2) aY Cc

Dd () P = (1/4 1/4 1/2 1/2) =

1/64, (3) bb CC dd P = (1/4 1/4
1/4 1/4) = 1/256 (4) aa bb Cc Dd () P = (0 1/4 1/2 1/2) = 0.
3.33 ww
: ww
. w + .
Punnett,

ww Y:

Ww

w+
www+

Y
wwY

w+O

YO

ww Y . (w)
(w ) , , (ww )
.

ww Y w +Y -

Punnett:

w+

wY

w+wY

wYY

w+w

wY

w+ww
wwY
,

ww

w+Y
YY

3.37

(
).
.

3.39 . -

. -

. -

( a = ):

(-2)

(a),

(-5).

(aa) -

( ).

, -1 -2, .

. -

(aa = ): -

(aa),

(a)

(aa). , -

. -

: ( ),

, -

( )

-6 -7 -

, -

). -

. -

, -

. ,

3.45

3.42 () ,

. () -

, -

. -

. () -

h -

1/4 -

, -

, .

. ,

() . , -

h + -

, -

h ,

3.43 () -

90% . , 10%

( aa

aa aa). ()
. ()
,

. ()
.
,
.

, ,
. ()

h +, ,
, .
4
4.2 : w 1/w 1, w 1/w 2,

w 1/w 3, w 2/w 2, w 2/w 3, w 3 /w 3 .

4.6 I AI B I Ai.

(I AI A, I AI B, I Ai, I Bi ) : A (P =
1/2), AB (P = 1/4) B (P = 1/4).
. P = 1/2 1/2 = 1/4.
. P = 0,
O.
. P [( AB) ( B)] = (1/2 1/4) (1/2
1/4) = 1/64.
4.8 O, i /i.
4.10 C R/C W C R/C W -

10

1 C W/C W : 2 C R/C W : 1 C R/C R. , .

4.12 F/F G N/G N f/f

G O/G O.
. F1
F/f G N/G O
.
. G
, 9:3:3:1
F2. 3/16 (F/ G O/G O), 6/16
(F/ G N/G O),
3/16
(F/ G N/G N), 1/16
(f/f G O/G O), 2/16
(f/f G O/G N) 1/16 (f/f G N/G N).
. F/f G N/G O
f/f G O/G O 1/4
(f/f G O/G O), 1/4 (f/f
G O/G N), 1/4 (F/f G O/G O) 1/4
(F/f G N/G O).
4.18 9:7 F2
9:3:3:1 A/a B/b A/a B/b F1. 9/16
A/ B/ 1 A/A B/B : 2
A/a B/B : 2 A/A B/b : 4 A/a B/b.
A B
, A/A B/B

.
A/A B/B 1/9 1/9.
4.19 9:7 F2 9:3:3:1,

A/ B/ A/ b/b,
a/a B/ a/a b/b .
. , ,
.
4.20 . A/a B/b A/a B/b
9/16 A/ B/, 3/16 A/ b/b, 3/16 a/a B/
1/16 a/a b/b. A/ b/b, a/a B/ a/a b/b
,
. A/ B/ . ,
9 : 7 .
. .
.
. a/a B/b A/a B/b
5/8 (1/8 A/a b/b + 1/2
a/a /) 3/8 (A/a B/).
4.23 C/c
Y/y
/agouti.
3 : 1 C/ , c/c
. 2
: 1 agouti :
Y/Y , Y/y y/y agouti.
agouti, c/c Y y,
c/c Y/y c/c y/y .

. ,

27/64 . 1 27/64 =

: C/ Y/y

37/64 -

c/c /y. ,

. F2 27 : 37 .

. -

1 Y/y Y/y -

,
A B

2 : 1 agouti. ,
C/c Y/y c/c Y/y.

C:

. C/c Y/y C/c Y/y ,

A/ B/ c/c.

Y/Y ,

F2 3/4
3/4 1/4 = 9/64. 1 9/64 = 55/64

1 : 2

: 1 agouti.

F2 9 : 55 .

C/c C/c 1 :

, Y/y.

4.25 . Y/Y R/R () y/y r/r

() F1 Y/y R/r (-). F1

.
F2

F2 1/16 (Y/Y R/R ), 1/8

oo (Y/Y R/r ), 1/16 (Y/Y

r/r ), 1/8 (Y/y R/R ), 1/4 (Y/y R/r ), 1/8 (Y/y r/r )
1/4 (y/y /) . F1
1/4 (Y/Y R/R ), 1/4 - (Y/y R/r ), 1/4 (Y/y R/R )
1/4 oo (Y/Y R/r ).
. Y/Y R/r (oo) Y/y r/r ( ) 1/4 oo (Y/Y R/r ), 1/4 - (Y/y R/r ), 1/4 (Y/Y r/r ) 1/4 (Y/y r/r ) .
. Y/Y r/r () y/y R/r
() 1/2 - (Y/y R/r )
1/2 (Y/y r/r ).
4.27 .
,
:
A/ B/ C/. F1 A/a
B/b C/c , ,
A/ B/ C/ (3/4 3/4 3/4) =

F2
2.

() () .

4.32 . , H/H H/h h/h . , H/H .

H/H W/W h/h w/w . -

F1 H/h W/w
, . F1 F2:

3/16 H/H W/

6/16 H/h W/

3/16 h/h W/

1/16 H/H w/w

2/16 H/h w/w

1/16 h/h w/w

11

12

, -

w. , -

9/16 : 3/16

H/h W/w,

: 3/16 : 1/16

H/h w/w, H/h W/w h/h

W/w H/H w/w.

4.35 .

3/16 : 9/16 :
1/16 : 3/16 .
4.34 ,

4.32

5.1 .

. -

H/ W/

/h w/w

H/H W/

/h W/

/h w/w

H/H w/w

H/H W/

/h W/

#1

H /h w/w

#2

H/H W/

25,21/8 =
3,15 cm
281,7/8 = 35,21 cm.
(s 2).

1:

s = s 2 =

1,70
7

s = s 2 =

(xi x )2
n1

= 0,24 = 0,49 cm

(xi x )2

413,35
7

n1
= 59,05 = 7,68 cm

. (r ) ,

(cov) -

. -

. x

h/h w/w,

y,

xiyi nxy

.
H/H, H.
w/w,

r=

covxy

sxsy

n1
sxsy

(xiyi)

.

.
(n 1)

. :
913,50 8 3,15 35,21

r=

0,49 7,68

3,74

= 0,99

3,76

. -

, -

. , 40

1,0.

= 50 s 2 = 5.

.
5.2
, .
(.. 1, 2, 5 .) .

().

. ,

. -

, 10

= 20 s 2 = 2

( ),

20

= 20 s 2 =2 10

= 30 s 2 = 2. -

13

14

5.4 . AA BB aa bb,
F1 Aa Bb.

50 cm

30 cm,
(50 30)/4 = 5 cm 30 cm. Aa Bb, ,
,

40 cm.

. -

40 cm. , -

Aa Bb, AA bb aa BB 40 cm.

. , -

. F2, 1/16 AA
bb, 4/16 Aa Bb 1/16 aa BB. ,

10 -

6/16 = 3/8 40 cm.

= 25 s = 5, 20 = 42

. -

s = 5 10 = 55 s = 5.


.
5.3 .

( ) , . F1
,

. , F1

,
, .
5.10 .
F1
, F2

.
5.13 ,


.
-

). -

5 cm

-. ,

1 . T,

F1 0, 1, 2

AA BB CC DD EE FF (160 cm)

3 . , -

aa bb cc dd ee ff (100 cm)

F2

AA

BB CC DD EE ff (150 cm) aa bb cc dd ee FF
(110 cm), F2 AA BB CC DD EE FF (160 cm)
aa bb cc dd ee ff (100 cm),
.
,
.
, , .
, F2 .
, ,
AA BB CC dd ee ff (130 cm)
aa bb cc DD EE FF (130 cm).
F1 130 cm
(Aa Bb Cc Dd Ee Ff ), F2 160 cm (AA BB CC DD
EE FF ) 100 cm (aa bb cc dd ee ff )!
5.14 F1 Aa
Bb Cc Dd Ee -.
F2 . F2
(2 3 ) (0 1 -

(1/2)5.


(1/2)5.
0
,
, .
,
(. )
.

F2
11 (1/2)10 = 11/1.024,
F2

165 (1/2)10 =
165/1.024. ,
F2 -

[1 (176 (1/2)10)] = 848/1.024.

:
. n = , s = , t = , a = -

15

16

F1 #1

F1 #2

F2

F2

(1/2)10

(1/2)5

(1/2)5

(1/2)

5(1/2)

5(1/2)

10

5(1/2)

(1/2)

5(1/2)

10

(1/2)

10(1/2)

10(1/2)

10

5(1/2)

5(1/2)

25(1/2)

10

10(1/2)

(1/2)

10(1/2)

10

(1/2)

10(1/2)

10(1/2)

10

5(1/2)

2
3

5
5

5
5
5

5
5

5
5

10(1/2)

50(1/2)

10

10(1/2)

5(1/2)

50(1/2)

10

10(1/2)

(1/2)

10(1/2)

10

}
}

b =
x! = (x)(x 1)(x 2) (1), 0! = 1.
p
:
p (s, t ) =

n!
s!t !

p (0, 10) =

p (1, 9) =

p (2, 8) =

p (3, 7) =

10!

ab
s

( )( )
( )( )
( )( )
( )( )
0

0!10! 2
10!
1!9!
10!
2!8!
10!
3!7!

11 + 165
1.024

2
1

1
2

848
1.024

2
1

10

1.024
10

AD
. ,
,
. ,
.
. -

11

1.024

, . -

1.024

45

1.024
120

11

1.024

1.024

5.18 SHR . ,

. ,
. (

5.15 . , -

, -

TIS

AD -

, -

.)

5.20 .

13,0 9,7 = 3,3 cm. -

HN2 R/S = 3,3/4,6 = 0,72.

, b,

5.26 ()

() -

:
, -

xiyi nxy
b=

covxy

sxsy

n1

(xi x )2

. =

xiyi nxy

VA/VP: 0,165
, 0,061

(xi x )2

0,144 -

n1

. , x

. ,

y -

5.27

xi = 111, x = 11,1
yi = 108,5, y = 10,85
(xi x )2 = 51,4
xiyi = 1.257,5
b=

1257,5 10 11,1 10,85

51,4

.


, =

53,15

= 1,04

51,4

.
-

. 1,04
. h

F1:
, F1
.

F1
, -

2 4 -

5.22 (S ) 14,3

9,7 = 4,6 cm. (R )

17

18

6
vg+ m/vg m+

vg+ m
()

6.2 2 , 2 =
16,1 < 0,01. < 1%.
,

vg+ m/vg
m+

( a/a). -

vg m+
()

vg+ m
()

vg+ m/ vg+ m vg+ m/ vg m+

-

vg m+
()

vg m+/vg+ m

vg m+/vg m+

vg+ m+
()

vg+ m+/vg+ m vg+ m+/vg m+

vg m
()

vg m/vg+ m

,
/ : b/b 1:1 (203:197), / :

a/a (240:160). a/a.

/a.
,

.
6.5 m ,
Xm/Xm vg +/vg +
Xm +/Y vg/vg . F1
Xm/Y vg +/vg, . F1
, m . vg m
, vg + m/vg + m vg m +/vg m + . F1 vg + m/vg m +,
.
F1 F1 ,
Drosophila ,
:

vg m/vg m+

,
vg m/vg m .
F2,
.
,
vg m
. m
2, 3
4.
6.6

, , (

).

. (A B C D E F ) = 0,40 0,45 0,35 =

,
.
50%,
50 . c b 66
(= 7 + 8 + 38 + 13), c b

0,063 6,3%.

. ( C d e f ) = 0,40 0,05 0,35 =

0,007 0,7%.

. ( b c D E f ) = 0,10 0,05 0,15 =

0,00075 0,075%.

. (a B C d e f ) = 0,10 0,05 0,35 =

0,00175 0,175%.

. (a b c D e F ) = 0,40 0,05 0,15 =

50%.

0,003 0,3%.

6.10 , -

6.15 D/d

P/p, 95% 5%

. a b /a b

(a b , a b) -

/h .
. 47,5% D P h d
p h, 2,5% D p h d P h.
. 23,75% d P H, d P
h, D p H D p h, 1,25% D
P H, D P h, d p H d p h.
6.19 ,
o
.
. F m W, f M w.
. M f W, m F w.
. F w M, f W m.
6.21. . (
):
P: + + dp/ + + dp b hk +/ b hk +
F1 ( ): + + dp/ b hk +
b hk dp/ b hk dp


(SCO)
(DCO), :

90% (a b,

a + b +) 10%. ,
45% a b +, 45% a +b, 5% a b 5% a + b +.
6.11 7 mu,
7% (3,5% a b
3,5% a + b +)
93% (46,5% a + b 46,5% a b +).
a + b +
.
. = 0,035 (a + b +) + 0,465 (a b +) = 0,50.
. = 1,
,
a + b +.
6.14

. , a
b 20 mu,
/a b
20% (10% A b 10% a B ) 80% (40% 40% a
b). , ,

19

20

l l

+ + dp

305

b hk +

301

,
,

b hk dp

171

SCO

+ + +

169

SCO

+ hk dp

21

SCO

b + +

19

SCO

+ hk +

DCO

( -

b + dp

DCO

,

. ,
l. , -

. DCO

l. -

: b .

, -

F1
(dp + +/+ b hk) -

(SCO)
(DCO) ( ), -

SCO -

(l /+) :

: dp b hk + + + -

dp b,

a ++

405

dp + hk + b + b hk.
(RF):
RF(dp b) = [(171 + 169 + 6 + 8)/1.000]
100% = 35,4%
RF(b hk) = [(21 + 19 + 6 + 8)/1.000] 100%
= 5,4%
: dp b
35,4 mu, b hk 5,4 mu, dp hk 40,8 mu.
. = 1
= 1 DCO/ DCO
= 1 (14/1.000)/0,354 0,054)
= 1 0,73 = 0,27
6.25
(l )
: : l /+ +/Y , F1: +/Y , l /Y
(), l /+ , +/+ .

+b+

44

SCO

+++

48

SCO

a b+

DCO

DCO a b l.

a + +, + b l.

a + +/+ b l. 44 + b +
b l, 48
+ + +
a b 2 a b + (
). , ,
.
, -

(RF)

RF(a b ) = [(48 + 2)/499] 100% = 10%

RF(b l ) = [(44 + 2)/499] 100% = 9,2%

.

(

6.26

) -

a c 14% 0,5% = 13,5% (6,75%

a c +, + + b) 12% 0,5% = 11,5%
c b (5,75% a + +, + c b).
[100% (13,5% +
11,5% + 0,5%) = 74,5%]
(37,25% a + b,+ c +). , :

. ( ) .
a c 14% (7%
a c, + +) c b 12% (6%
+ +, c b).
.

a+b

37,25

745

+c+

37,25

745

a c+

6,75

135

(0,12 0,14) 100% = 1,68%.

++b

6,75

135

a ++

5,75

115

0,3, -

+ cb

5,75

115

30%

Acb

0,25

+++

0,25

, 1,68% 0,30 = 0,50%

(0,25% a c b + + +).
6.29 . (. -

. -

):

a, b
a, c
a, d
a, e
b, c
b, d
b, e
c, d
c, e
d, e

902
973
957
497
875
945
497
930
498
496

98
27
43
503
125
55
503
70
502
504

9,8
2,7
4,3
50,0
12,5
5,5
50,0
7,0
50,0
50,0

21

22

a, b, c d ,

, -

2,

50%. e

,
Sharpen Up
. 83 367 + 260
= 627 . , , Sharpen Up

83/627 = 13,24%.
2
.
: Sharpen Up

,
/
. . c
C. Sharpen Up (cc),
(1) cc CC C ( ),
(2) cc Cc 1/2 Cc (), 1/2 cc
() (3) cc cc cc ().
( = 1/3),
, ,
627 [(1/3 0) + (1/3 1/2) + (1/3
1)] = 314. 627 314 = 313

.

.
, :

. :

b + d a + c/b d + a c +, e/e + b d a c/b d a c, e/e.

b + d + a + c + . b d, d a, a c.
b + d +
a + c + b d a c. , e + e. :
(b + d + a + c + e +) = (

e +)
= 1/2 (0,055 0,043
0,027) 1/2
= 1,6 105
= 0,0016%.
6.31 , 2

.



.
2

Sharpen Up, -

(13,24%) -

. ,

314 0,1324 = 42

313 0,1324 = 41. -

45 38 -

. , = [(45 42) /42

+ (38 41) /41] = 0,43, df = 1, 0,50 < P <

/-

0,70. , ,

Sharpen Up.

( ). -

: Sharpen Up

/ .
(1) cc
CC 1/2 Cc (), 1/2 cc () (2) cc cc cc ().

( =
1/2),
627 [(1/2 1/2) + (1/2
1)] = 470. 627 470 = 157.


(13,24%)
,

470 0,1324 = 62
157 0,1324 = 21 . 45 38 .
, 2 = [(45 62)2/62 +
(38 21)2/21] = 18,4, df = 1, < 0,001.
, ,

/
.

, .
K 7
7.3
, ( ) , .
,
, .

7.5 p a p +
: 20 PD,
18 NPD 8 . PD PD,
.
7.7 .

23

24

. -

w/w ch,
7.,
w/w ch. ,
w
,

: w/w w ch/w ch.
white

( 4). (w/w ) 0,44%

,
(w ch/w ch)
4,1%. w/w ch 2%
.
, w/w w ch/w ch.


. , -

= [(1/2 + NPD)/ ] 100%. : a-b 19,7 mu, b-c 11 mu, a-c 14 mu. , a-c-b.
.
, a b . (). ,
- :

= [(1/2
MII)/( )] 100%. -

: a-cen 20,7 mu b-cen 6 mu. ,

.

7.9 w/Y ( , )

w ch/w ch ( , ),
w/w ch.

7.A

, -

DNA , -

(w/w ) (w /w )

. -

(w/w ) .

GB4

7.12 -

. , -

, -

: ad w pu

ad w pu

y bi y bi, sm phe sm + phe +.

7.15 . GB4 [(3 109 bp/ 0,32 )/(25 106 bp/)]
38 . G3 ~200 .
.
, ,
. .
.
, DNA
.
GB4 DNA,

, 1 b.
,

,
DNA. , G3,
DNA,
250.000 bp.

: GB4
, G3
.

, 1 b. -

ch

ch

ch

G3, , .


C,
C

, C.

C --C.
7.17
lod lod (4,01)

25 mu. lod 3,
waf, .
, 1 mu
1 b, o 25 b waf.

25

26

8.1 . ( D E F
).
. ( C ).
. ( E F ).
. (

8.9 . -

E F ).
. ( C ).
8.2
,

(. 8.8).
8.4 . , .
.

w +,
w M 4

w M 4 . a, b,

c d . ,

.
.

. B C
:

ABCDE
A B C DA
EBCDE
A B C DA

O mottled -

( )

, -

(, ,

w + -

(, ,

w, w +

( , -

. mottled -

8.7 H -

DNA white .

aced

, -

w + -

.
w

(2) . (3)

. -

. -

mottled .

8.10 .

, -

Rec(8) -

8p23.1 8q22.1.

(4) -

Rec(8) -

8q 8p

M4

(. 8.10).

. 8.10,

8,

. -

Rec(8).

) -

Rec(8).

. -

Rec(8),

8q 8p

Rec(8).

Rec(8). -

8q 8p

Rec(8). ,

, -

. , 1/3

, -

Rec(8).

2/3 , 1/2

, -

8.

. Rec(8) -

. -

. (1)

8 -

. Rec(8),

. -

27

28

. -

Rec(8): -

Rec(8).

, .

, -

6,

. -

FISH, -

DNA / -

DNA (. 16 17).

8.11 .

, . -

. -

, -

8.15

.
.
(. 8.9).
8.13 . Lambert
6.
6,
-

8.17 . O Denton -

, -

. Denton

. -

Lambert .

6 12.

. 6

Lambert -

6 -

, -

12. -

. -

, -

6 .

12.

, -

12 .

. -

Denton, -

(-

6 12

),

. -

8.21 . 45.

. 47.

(.

. 23.

8.12, -

. 69.

1).

. 48.

. -

8.22 . T.

6 12

8.23 . ++

Denton.

c ( c -

6 Denton. ,

).

Denton

c c .

. -

, ,

. , -

. + +

12. -

+ -

. -

. -

, .

8.24 -

. -

( -

1) (.

).

8.12) .

. y/y +/+ ()

+/ pal/pal (), y/+ pal/+

() y/Y pal/+ (). -

y/+

pal/+ ,
. (+)

y/+ pal/+ ,
y pal/+. -

. ,
.
. , -

29

30

F1

y/+ pal/+.
()
(y/+), ()
(y).
,
(
)
( ).
. +/+ eye/eye ()
pal/pal +/+ (), pal/+
eye/+. 4
+.
pal/+ eye/+,
pal/+ eye.
4 .
4 eye.
,
4 .
,
.
. +/+ e/e ()
pal/pal +/+ (), pal/+ e/+. To
3 +.
pal/+ e/+, pal/+ e.
,
. ,
,
(pal/+ e/+).
8.28 . F1 aa.
.
F1 1, 2, a1 a 2,
, 12, 1a1, 1a 2, 2a1, 2a 2 a1a 2,
1/6 , 4/6 a 1/6 aa. Punnet,

35 : 1 aa.
1/6

4/6 a

1/6 aa

1/6

1/36

4/36 a

1/36 aa

4/6 a

4/36 a

16/36 aa

4/36 aaa

1/6 aa

1/36 aa

4/36 aaa

1/36 aaaa

8.32 17
. , 34 .
9
9.1 ,
-

F. F + F F,
.

Hfr F , F
F. F,
. 100 , ,
F .
9.2

9.5 A thy leu +, -

c +,

B thy + leu . DNA A

(3) F, c + .

B, DNA A

F +,

leu B

leu . DNA A
B .

B
.
leu +.
9.6 . ,
c + str R

C A, B,
D, E, F, G H.
c +
str R

( C
, ). ,
c + str R
a +, A,
B, D, E, F, G H.
,
a + c + str R. , a c + str R.
. 1:
c +, 1
c +. (1) F , (2) Hfr
F c + ,
-

, c +.

2: c + 6 , g +, h +, a + b + ,
2 Hfr.
c +, g +, h +, a + b +.
:
(0) c +(6) g +(8) h +(11) a +(14) b +(16).
d +, e + f +
.
-

Hfr F , -

F
,
Hfr. 2
F 16 F amp R, c + . F ,
, 2 , F.
, Hfr.
3: 3 c +
1 g + 3 . -

2 F

2 . .
, .
3 F DNA
c + g + F. -

31

32

, F
3 F ,

. ,

. -

cmlB , cmlB aroA pyrD. ,

(), aroA cmlB pyrD.
9.12
,

.
( ) cheB-eda, cheA-eda, cheA-supD, cheBsupD, eda-supD. edacheB-cheA-supD.
9.13 0,07 mu. E. coli K12()
r +, E. coli B
r + r . ,
E. coli B: 1 mL = ( ) ( /mL)
= 1.000 (672/0,1) = 6,72 106/mL. E.
coli B rIIx rIIy,
r +

rII. 1/2 r + 1/2 rIIxy.

r +
E. coli K12(): 1 mL = 2 (
r +/mL) = 2 (470/0,2) = 4.700/mL. rIIx rIIy
[4.700/(6,72 106)] 100% = 0,07 mu.
9.15
.
:

: c + 3 F 16 -

c + F amp R. , 3 F.
.
. 1
3
2. d +, e + f +
.

9.8 . .
. .
. .
. .
. .
. .
. .
. .
. .
9.10 , . pryD cmlB ,
. , () .
aroA pyrD
,

(). aroA

rII. ,

(
rIIA rIIB ).

9.17

: (1) r +,
. (2) r +,

.

9.20 .
.
. A, F D , B G C E .
9.22 . DNA leu leu +,

9.18

DNA

: (1) -

leu. ,
(leu +) ,


leu + (

).

. .

,
.
(2)
,
. A B: E. coli

K12()
,
-

33

34

leu -

( ,

. , -

3 8

leu ,

. 7 -

).

3 8,

9.26 .

. , -

, .

2, 4 7 .

, -

(

),
.
. (B2, B21,

. .
, .
, .
.
.
.
9.25 . , . ,
. ,

.

. -

B27, B28, B32, 901, LB2 D) , .

c75
D,
.
. c75 . .
.
,

. I



. , -

, -

c75 ,

. ,
.
9.27 . . , . ,
, ,
, .
.
.
, ,
.

, ,
.
.
.
, ().
.

-

,
I 1,0.
,
,
.
I .
39 C

.

, . , 39 C,
. 34 C
.
, 34 C
. RF


39 C

34 C.
, RF .
.

35

36

10

10.4 (), () (): -

()

C, N , -

. -

C, N -

DNA.

DNA -

. ,

, -

. -

. -

DNA

, -

10.9 . 3-TCAATGGACTAGCAT-5

( 5-TACGATCAGGTAACT-3).
. 3-GAGTTCTTAAGGT-5
( 5-TGGAATTCTTGAGAA-3).

/ -

10.10 - -

. , -

Drosophila ( -

. , -

http://www.flybase.org) -

- -

. , -

10.12 (G) = (C) () = (),

(G + A) = (C + T) (G + T) = (A + C). -

, (), () () 1.

10.15 DNA , () = ()

(G) = (C). 80 ,

80 . -

110 G, 110

C. , (110 +
110 + 80 + 80) = 380 , 190

10 DNA:

10.2 . .

10.16

. .

() () (G) (C), DNA -

. .

. (DNA IIIS -

DNA.

IIS

10.17 GC

).

, .

, GC

, -

, -

. GC ,
. , GC
DNA, .

GC,

. 3 -

: () 69 C, () 73 C, () 78 C, () 82 C

4.

() 84 C.

10.22 . -

. :

. 200.000 .
. 10
360. ,
100.000/10 = 10.000 .
. 0,34 nm

10.29 .

() . , 100.000 0,34 nm = 3,4 10 nm = 34 m.

10.24

5-GUUA-3 (0,30 0,25 0,25 0,20) =

. ,

0,00375. 10

S. cerevisae, -

10 -

CEN.

. 1, 2, 3

4, 2, 3, 4 5

... , -

0,00375 10 = 3.750.

, DNA .

10.25 . CGAGG 2

GCTCC

3. ,

.
. .

37

38

10

). , -

. -

: -

. .

. .

.
( ) .

,
.
. -

. DNA -

(,

RNA ).

, 70-

80% .

.
.
, .

. . DNA

10 nm, .

. ,

30 nm

, -

DNA

. ,

E. coli, 100

, -

DNA.

DNA 400 kb.

DNA, -

10.31 . -

10.32 16

( ), -

DNA CEN. -

DNA

(CDE, Centromere DNA Elements). CDEII,

76-86 bp >90% ,

CDEI,

. -

RTCACRTG ( R = A G),

CDEIII, 25 bp

, -

. CDE

11 DNA

11.2 :

. -

DNA = L, 14-14 = LL.

15

-15 DNA = ,

15

-14

, CDEI -

. 1: HL, 2: 1/2 HL, 1/2 LL, 3: 1/4

1 (Cbf1,

HL, 3/4 LL, 4: 1/8 HL, 7/8 LL, 6: 1/32 HL, 31/32 LL,

Centromere-binding factor 1), CDEII

8: 1/128 HL, 127/128 LL.

. 1: 1/2 HH, 1/2 LL, 2: 1/4 HH, 3/4 LL,

CDEIII

3: 1/8 HH, 7/8 LL, 4: 1/16 HH, 15/16 LL, 6: 1/64

Cbf3. Cbf3

HH, 63/64 LL, 8: 1/256 HH, 255/256 LL.

11.4 . DNA

10.34 DNA .

. ,

10.36 .

3 , -

5 3 -

. , DNA

DNA -

3 5 5 3,

DNA

DNA.

. DNA -

. -

. ,

. , -

). -

, 36 kb, -

. DNA

, -

DNA : -

. ,

DNA

(5 3).

Alu .

GC. ,


.

11.5 DNA in vitro DNA

, dATP, dGTP, dCTP dTTP, DNA .
11.6
Kornberg -

39

40

11

DNA in vivo

E. coli. -

polA1 polAex1. -

polA1 99% -

E. coli.

DNA ,

, DNA

( -

42 C, polAex1,

, -

, -

5 3.

, -

. -

DNA .

, -

11.9 -

, B D , C -

11.17 DNA

11.14 DNA

3-

E. coli. DNA

5-

DNA,

E. coli, DNA

DNA

(.

5 3 ( -

11.7). -

), 3

5 ( ). -

() -

DNA .

, -

11.19 . DNA -

ter

. -

cos

DNA

E. coli -

, DNA

, -

DNA

Okazaki.

. A

DNA -

DNA, ,

11.15

. ter -

() 12 -

. E. coli,

. -

DNA . , ,

, -

. -

E. coli,

11.21 DNA -

, -

. DNA

11.20 . 13

DNA 5 3.

() () (G) (C),

DNA. -

3 , -

, DNA -

. -

. -

, -

DNA -

E. coli, DNA

DNA.

13, -

3 5 -

. DNA

. , DNA -

. DNA

. -

. DNA

, -

, -

5 3, -

RNA

, -

5 Okazaki. DNA -

, V V

DNA.

13.

DNA DNA -

: Pol /, Pol Pol . -

DNA

41

42

11

10

RNA Pol

DNA.

30 -

, DNA -

. RNA/DNA -

() BUdR

Pol Pol

() .

. -

, DNA -

. DNA -

BUdR ( ) -

DNA -

DNA BUdR

DNA.

11.23 4

).
.

G2, 30
S 4,5

DNA

. -

DNA BUdR

- -

( ).

, 4,5

11.28 -

RNA. RNA -

. RNA

S,

, -

, ,

4,5 .

, -

11.26 . - -

DNA BUdR.

RNA .

BUdR ( )

11.31

DNA,

Giemsa

DNA BUdR -

, DNA ,

BUdR, BUdR
( BudR).

a1 a2+ a1+ a2,

2:2,

Holliday
a1 a2, . -

a1 a3+ a1+ a3 a2 a3+ a2+ a3

12

12.4 PKU,

U.

DNA. a1 a3 a1 a3

, -

2:2 a3 : a3 3:1 a1 : a1.

a1 a1 .

a2 a3 a2 a3, -

12.6

a2 ,

a3 a3 :

a3 3:1,
a3 a3+.
11.34 ry 206 ry 209 , ry206 ry+ kar ry+ ace kar+ ry+ ace+.
ry
ry+
. ry206 kar ry206 ace ry+
ry kar + ry 209 ace+,
209. ry209 kar+ ry 209 ace + ry+

ry kar ry206 ace,
206 (. 11.18).
kar ace (kar+ ry+ ace,
kar ry+ ace+)
:
Holliday
ry206 ry209 ,
(. 11.17).

, -

,

. ,
,
.
(
), ( ), (
).
.
,



. .

, .
, , ,
Tay-Sachs .
,

43

44

12

. -

. -

, -

, -

, -

-. -

, -

-.

. -

. ,

, ,

, -

. ,

DNA

. -

, ,
.

12.10 F1 F1 a +/a b +/b c +/c

d +/d a +/a b +/b c +/c d +/d.
.

12.7 -

F2, a /a, b /b /

c /c. ,

.
, : = 1
.
( a +/ b +/ c +/)

DNA
. ,
,

3/4 3/4 3/4 = 27/64. -

12.15 4

1 27/64 = 37/64.

. -

. -

a / b / c / d /d.

E. coli

3/4 3/4 3/4 1/4 = 27/256.

12.12 . -

) -

a , b c -

17 C 42 C

. -

1 -

. F2
a /a b /b c /c. -

17 C 42 C.

1/4 1/4 = 1/64.

1/4

. -

1, ,

d, F2

. F2

. 17 C, -

d /d / / / ( -

17 C, -

) d / a /a b /b c /c

, ,

( ). -

1 1) + (3/4 1/4 1/4 1/4) = 67/256.

, 42 C,
-

12.14 . 1/2 w /w bw /bw st /st , -

1/2 w /Y bw /bw st /st ,

1,

, -

(1/4 1
+

. w /w

se /se bw /bw w /Y se/se

bw/bw ,
.
. w /w + v /v + bw /bw, w +/Y v /v + bw/bw,

.
. 1/4 w +/w w +/Y bw /bw + st /st +,
, 1/4 w +/w w +/Y bw /bw
st /st +, , 1/4 w +/w w +/Y bw /bw + st /st,
, 1/4 w +/w w +/Y bw /bw
st/st,
3-
.
+

. E. coli

42 C, ,
, . , , 17 C
,
, . , 1, 17 C 42 C.
2,

45

46

12

. . -

42 C 17 C. -

c .

d .

12.18

12.16

. , -

( -

),

. -

. -

. ,

1 4
3 4
.

,
.


( )
. , ( ) ,


. , D E

4.
(D E)

4 -

(
)
,
. a ,
. c ,
a b d. d, b

. , -

a c d b.

12.17 .
c d + c + d.
. c /c + d /d +
PD ( , 4

cd + 4 c +d, ) NPD ( , 4 c +d + 4 cd, 4 4 ), c d .

, 1,25-

( D), -

. 4

().

, 4

12.22 p + 50%

p +/p +. p + -

12.19

( p +/ c +/c +

d e ,

), -

50% -

e. ,

(.. 25%

, c d,

. , -

100%

. ,

12..

12.23 . -

, -

. ,

12.20 . ,

50%

GSS ,

(-

/+)

GSS. ,

. -

Lesch-Nyhan ,

, .

. 1-

GSS -

25-

25--

. -

, . -

47

48

12

12.

50:50
+/+

()

+/+

+/+

p+/p+

100

100

p /p

()

()

()

20

60

p /p

50

p /p

300

200

p /p

p /p

50

. 1,

. ,

9% GSS, -

, , 5--

2, 50% -

GSS,

. , -

GSS 9%

GSS.

, 5--

. -

. GSS

50% ,

( -

GSS -

). -

5-.

( ),

, -

GSS.

( ,

. ( -

GSS -

14.)

GSS -

.)

. GSS

. HbC HbS

GSS -

HbC ,

HbS .

GSS -

, -

. ,

. , HbC/HbS

GSS. -

HbC

GSS, (

HbS.

) .

12.31 . , -

, -

12.000 -

1 2.000 . -

GSS

12.27 . 12.11, Hb

17.000 1 90.000 .

Norfolk -

, HbS

. -

. , -

. Guthrie -

, -

. , -

B. subtilis

-2-.

, -

B. subtilis,

. , -

. , -

HbS/+

49

50

12

12.33 .

DNA

. -

, -

, Guthrie

. -

, -

. -

. DNA

(..

. , -

). -

DNA .

DNA -

. ,

. -

, -

. Lesch-Nyhan

DNA -

, -

Chvez . -

Tay-Sachs -

. -

. , -

DNA -

. RNA

- ,

. -

tRNA. DNA

. Lieberman

, -

, -

( )

Tay-Sachs. ,

Chvez ,

mRNA,

. DNA -

Lesch-

Nyhan. -

, RNA -

( mRNA -

), -

( rRNA -

. ,

tRNA

),

, . -

RNA ( snRNA

).

DNA, RNA

DNA RNA -

. ,

5 3. -

DNA -

Lesch-Nyhan.

-. , DNA -

13 :

3-OH -

13.1 DNA RNA

RNA .

RNA -

, -

. DNA ,

DNA . ,

RNA . -

RNA -

DNA -

DNA , RNA -

51

52

13

13.3 , RNA -

RNA 5

3,

DNA 3 5. , -

RNA

. E. coli -

RNA

. , -

, -

. -

RNA : RNA I,

II III. RNA I 28S,

RNA

18S 5,8S rRNA .

RNA II hnRNA (

mRNA) snRNA

. RNA III

13.7 . . 5-AG-3

tRNA,

5S rRNA snRNA.

RNA -

10 35 -

E. coli, -

RNA

70. , ,

( -

. 35

).

(TTGACA)

8 5 -

35 bp

10 (TATAAT)

( 35),

32

5 .

10 bp

( 10), -

5-AG-3 -

17 bp DNA -

44 5

RNA

. .

DNA,

. (3 5) .

. (5 3) .

. -

13.9 . E. coli -

. -

. tRNA -

, -

. -

. -

35 10 -

RNA -

. -

RNA.

13.18 , -

18S, 28S 5,8S rRNA

rDNA pre-rRNA,

( ),

( ) -

, -

T4 ( ).

rRNA (. 13.19).

70

32

54

23

. -

5S RNA -

RNA , -

5S rRNA, -

5S rRNA , -

rRNA

. -

pre-rRNA Tetrahymena -

, ,

- 28S rRNA

N2, )

( I).

28S rRNA --

13.10 RNA I -

o pre-rRNA (. 13.14).

18S, 5,8S 28S

rRNA. RNA II -

RNA -

II -

mRNA.

. RNA II

snRNA. RNA III

20 30 RNA,

5S rRNA, tRNA

5 -

RNA.

m7Gppp 5-5 -

, 18S, 5,8S, 28S 5S

. (A),

rRNA -

, 3

. mRNA -

(A),

53

54

13

. , -

mRNA,

mRNA

. mRNA -

- , -

. -

, .

13.21

2, , -

, -

. ,

, -

13.22 1 (5 m7G) + 100 ( 1) + 50

. -

( 2) + 25 ( 3) + 200 ( (A)) =

376 .

13.23

5-GU-3

. U1

U1 snRNA

, U1 snRNA

5 -

GC TA -

mRNA. -

U1 snRNA,

, -

tub mRNA

. -

U1 snRNA

mRNA tub/tub. -

5 . ,

tub mRNA,

(), -

tub + ,

, -

2 -

( , -

, -

. -

3 ), -

mRNA. ,

tub ,

tub + -

. ,

tub/tub +

mRNA.

mRNA -

, -

. ,

, ,

. tub -

(. 14.2).

. tub/tub,

, -

. -

, -

13.26 . 3.

. -

. 1, 2, 3, 4.

. 3.

. 1, 2.

. 1 ( tRNA

rRNA ).

. 4.

. 1.

,
. ,

14 :

14.1 .

14.3 . -

, -

(.

12.10). ,

14.4 . ,

, ,

. -

. -

. ,

- - -

. -

.
-

. -

. -

, -

. -

55

56

14

Shine-Dalgarno

mRNA -

, (), -

. , 23S rRNA

14.6 -

rRNA -

. , 3 16S rRNA

23S rRNA. -

mRNA. rRNA E. coli -

Shine-Dalgarno, -

50S,

mRNA (RBS, Ribo-

rRNA 1.

some Binding Site). -

14.9 H 14.7 14.1

16S rRNA / mRNA

. -

3 16S rRNA

tRNA

Ile
Phe
Tyr
His
Gln
Asn
Lys
Asp
Glu
Cys
Trp
Met

1
1
1
1
1
1
1
1
1
1
1
2

Val
Pro
Thr
Ala
Gly
Leu
Arg
Ser

2
2
2
2
2
3
3
3

3 1 tRNA ()
2 1 tRNA ()

1
, 1 tRNA 1 tRNA

4 : 2 1 tRNA ()

6 : 2 1 tRNA ()

32

61

P (CCC) = 0,6 0,6 0,6 = 0,216 21,6% Pro

P (CCA) = 0,6 0,6 0,4 = 0,144 14,4% Pro
( 36% Pro)
P (CAC) = 0,6 0,4 0,6 = 0,144 14,4% His
P (CAA) = 0,6 0,4 0,4 = 0,096 9,64% Gln

14.10 , ,
tRNA
. , , -

. 4 G : 1 C

2 = 8 ,

14.14 mRNA

: GGC, GCG, GCC, CGG, CGC, CCC

CCG. 80% G 20% C:

7--G 3 (A).
.
eIF-4F 40S, MettRNA. Met eIF. ,


(A) eIF-4G,
eIF-4F -

P (GGG) = 0,8 0,8 0,8 = 0,512 51,2% Gly

P (GGC) = 0,8 0,8 0,2 = 0,128 12,8% Gly
( 64% Gly)
P (GCG) = 0,8 0,2 0,8 = 0,128 12,8% Ala
P (GCC) = 0,8 0,2 0,2 = 0,032 3,2% Ala
( 16% Ala)
P (CGG) = 0,2 0,8 0,8 = 0,128 12,8% Arg
P (CGC) = 0,2 0,8 0,2 = 0,032 3,2% Arg
( 16% Arg)
P (CCC) = 0,2 0,2 0,2 = 0,008 0,8% Pro
P (CCG) = 0,2 0,2 0,8 = 0,032 3,2% Pro
( 4% Pro)

.
14.17 -

. 1 : 3 U : 1 C -

33 = 27 . -

() -

, UAA, .

. -

20% , 60% U 20% C,

0,6 0,2

0,2 = 0,024 2,4%. 26 (97,6%) -

. 4 : 6 C

() () -

2 = 8 ,

: AAA, AAC, ACC, ACA, CCC, ACA, CAC

. -

CAA. 40% 60% C:

P (AAA) = 0,4 0,4 0,4 = 0,064 6,4% Lys

P (AAC) = 0,4 0,4 0,6 = 0,096 9,6% Asn
P (ACC) = 0,4 0,6 0,6 = 0,144 14,4% Thr
P (ACA) = 0,4 0,6 0,4 = 0,096 9,6% Thr
( 24% Thr)

27 ,
. . :

57

58

14

(0,8/0,976)% = 0,82% Lys

Lys

3,28%

(3,2/0,976)% = 3,28% Asn

Asp

3,28%

(12,0/0,976)% = 12,3 % Ile

Glu

3,28%

(9,6/0,976)% = 9,84% Tyr

Cys

3,28%

19,2/0,976)% = 19,67% Leu

Ile

3/61 = 4,92%

(28,8/0,976)% = 29,5% Phe

Val

4/61 = 6,56%

(4,0/0,976)% = 4,1% Thr

Pro

6,56%

(0,8/0,976)% = 0,82% Gln

Thr

6,56%

(3,2/0,976)% = 3,28% His

Ala

6,56%

(4,0/0,976)% = 4,1% Pro

Gly

6,56%

(12,0/0,976)% = 12,3% Ser

Leu

6/61 = 9,84%

Arg

9,84%

Ser

9,84%

14.18 20

, 20 .

. A : 1 U : 1 G : 1 C

4 = 64 :

1/64. -

mRNA,

20 .

3/64,
1/61.

. 5

25 = 32

. 3

33 = 27

. 2

52 = 25

. (

14.20 . 3-TAC AAA ATA AAA ATA AAA ATA AAA

14.7), -

ATA...-5. ( fMet Met -

.)
. 5-ATG TTT TAT TTT TAT TTT TAT TTT

TAT...-3.

Trp

1/61 = 1,64%

Met

1,64%

Phe

2/61 = 3,28%

Try

3,28%

His

3,28%

Gln

3,28%

Asn

3,28%

.
tRNA Phe 3--5, tRNA Tyr 3-AUA-5.
14.22 RNA : Lys
(AAA, AAG), Arg (AGG, AGA), Glu (GAG, GAA)
Gly (GGA, GGG). ,

. RNA -

14.26 , , ,

: Lys (AAA), Asn (AAU), Ile (AUA, AUU),

. -

Tyr (UAU), Leu (UUA), Phe (UUU) -

(UAA). -

. ,

(1/4 3/4 3/4) = 9/64 = 0,14 14%. ,

( -

RNA

):

. A 14%, -

Met-Phe-Ser-Asn-Tyr- ... -Met-GlyTrp-Val

(1/0,14 = 7,14).

Met-Phe-Ser-Asn

...-Met-Gly-Trp-Val (

RNA

AUG.)

14.24 5-GAU-3

: Met-Phe-Ser-Asn-Tyr- ... -Met-Gly-

5-AUC-3,
Ile. 5-CAU-3

Trp-Val
:

5-AUG-3,
Met. tRNA -

Met-Phe-Ser-Lys-Tyr- ... -Met-GlyTrp-Val

Met-Phe-Ser-Asn-Ser- ... ( -

tRNA.Met

5-AUG-3, -

. -

, , Ile

Met. -

, ...Trp-Gly-

tRNA.Met,

Gly-Cys...)

: Met-Phe-Ser-Asn-Tyr-...-Met-Gly-

Met. , -

Trp-Val-Trp... ( -

, tRNA

.)

AUG -

14.30 . mRNA -

, 5

Met-Val-Ser-Ser-Pro-Ile-Gly-Ala-Ala-Ile-Ser

().

Met-Val-Ser-Ser-Pro-Met-Gly-Ala-Ala-Ile-Ser

Met-Val-Ser-Ser-Pro-Ile-Gly-Ala-Ala-Met-Ser

1.480

Met-Val-Ser-Ser-Pro-Met-Gly-Ala-Ala-Met-Ser

1.480 3 = 4.440 . 5

59

60

15

3 -

15.6 .

6.500 4.440 = 2.060 ,

5-CUG-3,

32% mRNA.

tRNA 5-CAG-3. -

. F508

tRNA 5-GUG-3,

5-CAC-3.

DNA.

CG

GC.

. Leu.

mRNA

. Val.

mRNA

. Leu.

. -

15.8

, -

. -

. , -

5BU

DNA .

DNA

, -

C DNA .

, TA CG.

, -

lacZ-1 -

, -

14.31 -

. -

. D

, -

(.

12.8). lacZ-2 , -

15 DNA,
DNA

, .

15.1 .

15.3 . ( -

).

15.4 . - -

. -

, -

15.9 . .
. , mRNA UGG
UAG, ,
5-AUG-ACC-CAU-UAG--3.
15.11 . UAG: Gln (CAG), Lys (AAG), Glu (GAG),
Leu (UUG), Ser (UCG), Trp (UGG), Tyr (UAC,
UAU), (UAA).
. UAA: Lys (AAA), Gln (CAA), Glu (GAA),
Leu (UUA), Ser (UCA), Tyr (UAC, UAU), -

15.23 - -

(UGA, UAG).

G -

. UGA: Arg (AGA, CGA), Gly (GGA), Leu

, C

(UUA), Ser (UCA), Cys (UGC, UGU), Trp (UGG),

(UAA).

. C,

15.13

U C:

15.18 . 5

15.27 C -

. -

U. U .

CG

G.

GC :

.

(U C). Pro CCN, ()
Ser UC, Leu CUN
Phe UUY. (

.
C U CG TA.

5 3,
.)
DNA, 5-C

15.21 dGTP

Pro

UCN Ser, -

3-AGN-5.

61

62

15

C
Pro DNA, CUN
Leu, 3-GAN-5.

C DNA,
(UUN mRNA),
3--5. Phe , C (U
mRNA)
3--5 3-G-5.
C U

,
(U mRNA) Phe DNA ,
3--5.
, .
15.28
()
.

ara-1: BU ,
, ara-1
. BU CG GC,
CG
. ara-1, , ,
TA CG CG .

ara-2: BU, AP HA, .

CG
, TA CG.
ara-3: ,
CG. , CG CG
.
.
15.35 15.24,

Ds (
),
C
. Ds C. Ds
C Ac ,
.
.
Ds.
C,
C Ac,

Ds. , ,
C.
B D, D
Ac. , C, A D B D
Mendel .

15.37

northern DNA RNA, -

RNA,

, -

Ty),

RNA. -

RNA.

16.4

DNA -

RNA -

DNA A, T, C G -

DNA ( -

, -

).

(-, -, G-C C-G)

16 DNA

1/4. (1/4)2.

16.2

, n -

(1/4)n. -

DNA

, -

. -

1/4.096 = (1/4)6,

(1) DNA .
(2) :

.
16.7 ,

DNA -

Southern

E. coli

northern.

, -

DNA RNA -

16.12

(3) (dT)

()

cDNA mRNA.

(4)
- PCR.

cDNA

, ,

. -

. ,

Southern

cDNA -

63

64

16

. cDNA

-, -

. -

, cDNA -

AUG. , ,

. -

, -

, -

cDNA

. , -

cDNA 5

Shine-Dalgarno,

. , -

Kozak.

16.13 cDNA. -

--

cDNA -

mRNA . -

pre-mRNA,

16.16 -

. cDNA ,

1. DNA.

2. DNA

16.14 pUC19 . -

Shine-Dalgarno

( Sau 3A) -

lacZ.

cDNA -

(5-10 kb).

pUC19, -

3. 5-

10 kb

lacZ. ,

( -

cDNA 5

).

4.

pUC19 ,

BamHI,
Sau 3A. Bam HI
Sau 3A.

,
cDNA,

5. DNA ( , pUC19), .

.
.

6. E. coli.

, ..

7.

( pUC19

) X-gal, -

. -

( -

).

BAC -

DNA ( 200

16.22 -

300 kb).

16.17 = ln(1 P)/ln(1 f ),

. , -

N , , f DNA
ln . , p = 0,90, f = (2 105)/(3 109) = 34.538.
16.20 . 16.17, = [ln (1
p)]/[ln(1 f )]. , p = 0,95. (i), f = (7
103)/(3 109). (ii), f = (1 106)/(3 109).
(i)
1,28 106 (ii) 8,99 103.
.
,
. ,



. ,
1 b
7 kb.
, ,

, ura3

URA3. ,

.
URA3.
URA3 ( ),

. ,



,
.
,

.
16.24 ,
DNA

65

66

16

. -

, 5-ATG TT(T

G) TA(T C) TGG ATG AT(T, C A) GG(A, G, T

pUC19 EcoRI

C) T A(T C)-3. 96 -

10 kb.

. ,

, -

DNA (10 kb) DNA

cDNA (. 16.11).

(2,69 kb).

16.27

. -

EcoRI 10 kb, -

10 kb.

. -

16.30 DNA -

( ), -

1.900 bp

2.100 bp

2.500 bp .

16.31 . 2,0 kb,

, 2.500 bp 2.100 bp

DNA

1.900 bp,

- 200 bp

- 600 bp .

EcoRI.
.

, , DNA .

16.29 . DNA

,
EcoRI
. EcoRI,
DNA

EcoRI, DNA
EcoRI. , ,

.
. ,
.
. (), .

EcoRI ,
.

,
EcoRI.

.

. EcoRI ,
EcoRI.
EcoRI 3,0 kb 14 .
14
. ,
.
EcoRI ,
. , ,
3,0 kb.
. TDF
DNA ,
.
16.32 . Not I 8 bp, Bam HI 6 bp. 8 bp 16 6 bp
(. 16.6).
. Not I cDNA
47 kb. ,
.
. Bam HI 10,5 kb, 8,2 kb
4,1 kb
cDNA. , .
. RNA
28,9 kb.
cDNA, DNA .
. BAC BamHI

Not I.
: 47 kb Not I.
,
.
BamHI

.
16.34
, ,
pre-mRNA
mRNA ,
. ,
. , DNA
northern
mRNA .
mRNA ,
, . mRNA

, . DNA, ,
RT-PCR cDNA
mRNA .
. ,

. ,

67

68

16

DNA, PCR -

Southern

DNA cDNA,

cDNA. , -

DNA.

16.35 .

/ -

Southern cDNA

2,0 kb, 1,4 kb 1,3 kb -

, mRNA -

cDNA

. northern

. 7,8 kb,

7,4 kb, 6,1 kb 3,6 kb

pre-

cDNA -

mRNA -

. , -

. -

, 0,8 kb

0,8 kb, , ,
.
. cDNA
mRNA .
northern pre-mRNA
(
) . ,
.

(

16.36 -

Eco RI-Hin dIII Bam HI),

( cDNA

DNA 3

. -

Eco RI-Hin dIII 0,5 kb, -

, -

).

. -

Southern

. -

, cDNA -

. -

. ,
. ,
.
, ,

16.42 -

DNA,

17.6 SNP

PCR. PCR, DNA -

Taq -

, SNP

, -

RFLP. STR

, -

VNTR -

( 2 6 bp

STR 7 bp

, -

VNTR), SNP.

17.7 -

, , -

STR

PCR,

. ,

PCR , -

. , -

STR .

PCR, -

, -

17
DNA

.
STR -

17.3 -

( -

, -

. -

),

69

70

17

DNA. ,

x .

STR -

, 1,0 kb

DNA.

x -

17.8 . -

,
x.

10 . EcoRI
:
,
.

RFLP.

EcoRI -

, -

( ) -

: 5,0 kb, 3,1 kb, 1,9 kb, 4,0 kb 2,1 kb. -

, 5

5,0 kb, -

2.

, x.

EcoRI 5,0
kb. ,

5,0 kb
1 2. , , EcoRI.
,
.

1
2
3
4
5
6
7
8
9
10

, , / , ,
, / ,
x, / x,
x, , / x, ,
, / , ,
x, / , ,
, , / x, ,
, / x,
, / x, ,
x, / x, ,

. 1 -

, , / , , ,

, ,

4. -

Southern,
, : 3,1
1,9 kb. 6
, , / , x.

,
5,0 kb. 1,9 kb
4,
3,1 kb.

, ,

Tsp 45I,

, x 4,0 kb. ,
, , / , , , , / , x
1:1.
( 3,1 /
1,9 kb 4,0 / 3,1 / 1,9 kb).
17.9

.
, .
. , .

.

, . , . (



DNA .)
17.10 . PCR-RFLP:
DNA
PCR 200 bp
4. PCR,
Tsp 45I

. Tsp 45I
120 bp 80 bp. -

200 bp.
. PCR-RFLP,
DNA 120 bp
80 bp.
200 bp.
200 bp, 120 bp 80 bp.
. RT-PCR cDNA
PCR
Tsp 45I. , mRNA
. , cDNA (dT). ,
4 cDNA PCR, Tsp 45I
. 200 bp
, .
120 80 bp,
. ,
cDNA RT-PCR DNA.
17.14 James Susan Scott .
DNA
DNA
(.. , ,
). ,
DNA
Larson.
Bobby Larson. , -

71

72

17

) (2) -

DNA .

DNA .

. ( ) = (0,112 0,036

0,081 0,195) = 6,369 105. 1

15.701
, .

. ( ) = (0,112 0,036

0,081 0,195 0,062 0,075 0,158 0,065

0,067 0,085 0,089 0,028 0,039) = 1,683

1015. 1 5,94 1014

17.15 . PCR

() DNA- ,

17.20 ,

, -

DNA-. ,

. -

VNTR -

() DNA,

DNA

. -

( bp),

Southern. -

. ,

( DNA -

) .

RFLP

STR -

VNTR .

. PCR -

Southern

: (1)

STR

DNA

. -

in situ . -

, -

.
17.23 .

fru (
cDNA)
BD Gal4p.

D
Gal4p cDNA
Drosophila. ,

(. 17.15). ,
AD BD fru

cDNA . DNA cDNA
.
17.25 RNA

mRNA
(1) northern (
mRNA ,

cDNA (2) RT-PCR.
18
18.3 .
25% , , G, C,
6 bp (1/4)6 = 1/4.096
8 bp (1/4)8
=1/65.536. ,

Apa I, Hin dIII, Sac I Ssp I

4.096 bp Srf I Not I
65.536 bp.
. i.

: (1) (..
) / (2)
(..
-
G-C).
ii.

: (1) / (2) .
iii. Mycobacterium
tuberculosis
25% (, , C G),
6 bp 16
8 bp.
,

.
Mycobacterium tuberculosis
G-C
- / , 5--3, 5--3, 5--3 / 5--3
.
Sac I -

73

74

17

5-AG-3 5-CT-3,

Hin dIII, .

DNA PCR. -

PCR

, -

STS , -

. -

. -

STS -

. -

. , -

PCR -

4 b Caenorhabditis

elegans, Hin dIII

1.333 , Srf I 4
.
1.333 ,

. Not I 15 .

. : E. coli
Apa , Srf I Not I, M. tuberculosis
Hin dII Ssp I, A. thaliana C. elegans
Not I S. cerevisiae, D. melanogaster,
M. musculus . sapiens Srf I Not I.
18.4 . ~30 ,
() (CAG)N. 32, .
.


.
. , (
DNA ) PCR.

STS STS (.. ).

. 960 8.203 = 7.874.880 PCR.

. (i)

, 96
12 () + 8
(), 20 .
10 200 . (ii)
8.203 STS
200 8.203 = 1.640.600

PCR. (iii) , F, 6. (iv) ,

.
. STS
YAC
YAC . YAC

.
18..
.
: 2631 63 210
6892 5192 719 6193 991 3097,
4630 552.
. .

YAC . -

18.A
YAC

(2631

63)

(210

6892)

1071

(5192

719) (6193

991)

714

(3087

4630

522)

713

II-6A
IV-C3
X-H9
VII-E12
I-C8

STS. -

10

: 1, 1,

STS, -

1 1, 1, 1,

PCR Southern -

1 1 ., 96

STS .

. , -

= 1,22 1010 .

18.7 . 500 (13.543.099 + 10.894.467)

. -

96 + 80 = 176 (176

2 kb

8.203 = 1.443.728 PCR) -

200

, -

18.5 , . FISH

BAC -

. -

BAC -

5 kb, -

10 kb

FISH

. -

10 kb

2 kb -

. -

BAC -

. ,

STS

BAC. -

C -

75

76

18

18.8 -

70

35 (TTGACA) 10 (TATAAT)

. -

, ORF,

, -

(UAAGGAGG) -

. -

AUG ( GUG

. -

, -

(UAG, UAA, UGA), ORF.

Shine-Dalgarno

. -

. -

, -

. -

mRNA. -

. ,

, -

18.9

DNA, -

GU 5

, G 3

18.12 . ORF

CURAY

, 18-38

3 .

ORF -

. -

. -

, -

. , -

. -

, -

, -

ORF

. -

. ,

ORF

18.14 . cDNA -

. :

DNA -

(1) -

(..

-, -

, , -

), cDNA

. -

, (2) -

cDNA -

(..

).

. cDNA

, .

18.15

(.. )

. -

cDNA

-, -

. -

( -

) -

EST -

, -

EST -

. -

EST

, /

77

78

18

18.18 DNA

DNA -

DNA. -

.
, , DNA.
, ,

, -

DNA

,
DNA -

, -

. , -

(-

( + =

) , -

DNA

. -

.
, , EST.
,

(knockout) -

.
18.19
, , DNA

lod.

.
,

BAC .

SNP.

18.21 .

V . ,

V, -

. -

, -

. , -

V, -

19.9 -

lacI + lacO c lac + lac + lacY /lacI +

lacO + lac + lac lacY + ( lacI +,

lacI
lacI ).
19.11 19..
19.14 CAP,
cAMP,
RNA Plac. RNA CAP-cAMP
( ). ,
CAP
lac RNA

.
19.16 . DN
in vitro

DNA. ,

DNA
,
. , DN,
(
) .
, DNA
.
DN , CAP-cAMP.

.
,

, .
. ,
.
19

19.2 lac trp
.
-
DNA. Lac, lac,
lac.

TrpR, trpO
trp.
19.5
lacI lacO c.
19.7 .
, .
. ,


-. , ,
-, .

79

80

19

19.A

I P O Z Y
I + P + O + Z Y+
I + P + O + Z + Y
I P + O + Z + Y+
I s P + O + Z + Y+
I + P + O c Z + Y+
I s P + O c Z + Y+
I + P + O c Z + Y
I d P + O + Z + Y+
I P + O + Z + Y+

.
.
.
.
.
.
.
.
.

I + P + O + Z Y
I P + O + Z + Y+

I + P + O + Z Y
I s P + O + Z + Y

I + P + O + Z Y+
I + P + O c Z Y+

I + P + O + Z + Y
I P + O c Z + Y

I + P + O + Z Y+
I s P + O + Z + Y+

I + P + O c Z + Y+
I d P + O + Z + Y

I + P + O + Z Y+
I + P O c Z + Y

I + P + O + Z Y+
I + P O + Z + Y

I + P + O c Z Y+
I P O + Z + Y+

I + P + O + Z Y
I P + O + Z + Y

I + P O + Z Y+

+
+
+
+

+
+

+
+

+
+
+
+

+
+

+
+

. . 19.14.

. i. -

, -

, -

-,

ii. 12

iii. 69

10, -

CAP. CAP-cAMP

CAP, RNA

.
trp ,

.
iv. +28 -

Shine-Dalgarno. , ,

. -

--

Trp-tRNATrp,

, ,

Trp -

-.

v. +9 -

, -

. , -

mRNA. , -

cis- ),

trp -

( ).

Trp,

. -

19.19 trp ,

Trp-tRNATrp.

. -

, trp

. -

Trp-tRNA . -

- Trp

19.20 1:

(. 19.17). -

, -

Trp --

, RNA RNA -

. -

, ,

. , -

Trp -

, -

, ,

Trp

81

82

20

2: trp
, 2 1.
3: trp
trp (.
19.15).
,

trp trp, . , , 3

.
,

( ).
4: Trp-tRNATrp

trp. , Trp-tRNATrp , trp
. ,
, 4 ,
.
,
4 .
5: 5, 3 4 ( 19.17)
,
.
4.

19.24 cI
. cI

, .
19.26

-
-
UV

Cro DNA.

p
.

cII
.

pQ
.

P RM RNA
.

20
20.2 ,


. 5 trans-

( RNA-

), 3

, -

. , -

, -

. -

20.5 . 4 kb

. , -

DN I, -

DNA

20.9 rRNA

DN I. DN I

rRNA. ,

2 kb, -

rRNA -

4 kb EcoRI.

3 kb

, -

DN I -

mRNA. mRNA -

3 kb

EcoRI -

. , -

20.11

DN I

, 4 kb 3 kb,

DN I -

. , -

. -

(1) ,

DN I

EcoRI -

3 kb 4 kb. -

, (2) -

, -

mRNA , (3)

mRNA

20.7 . -

, (4)

, DNA

mRNA , (5)

mRNA

, (6)

mRNA -

. -

, (7)

( ) .

83

84

20

mRNA

mRNA

. , -

20.16 . X, -

CGG -

mRNA.

. Huntington,

. -

mRNA

CAG,

ChAT/VAChT
mRNA. , mRNA ,
mRNA. mRNA
, .
. ChAT VAChT:
mRNA . 3

mRNA . ,
AUG.
20.21 . .
. cDNA

mRNA. , 5 :
cDNA 4 5 cDNA 1, 2 3.
cDNA 1 5
cDNA 2 3. ,
cDNA 2 5
cDNA 3.

huntingtin, , .
.
CGG
FMR-1 .

. ( ,
X , , ,
.) , CAG
Huntington
, . ,
.
.

CGG . , huntingtin
36 .
20.20 . ,

21

. -

21.3 -

, .

, -

-.

. -

, DNA

. ,
-, . ,
, . ,

. , . , -,
,
, .



, ,
.
21.6 . Wilmut
,
. ,

,
(
G0 ).
-
,
.
,

. ,

. ,

.
. B .
, .
. .
,
, -
.


.
.

.

.
21.13 DNA .

85

86

21

Xic. X,
gfp .
. Xist .
, X gfp
,
Xic .
Xist RNA
,
H3.


Xic.
21.18 . X:A


X (sis-a, sis-b,
sis-c) (dpn).

. X:A 2:2,

,
.
Sxl PE.
X:A 1:2,
,
Sxl. , Sxl
PE X:A SXL
.
. PE
SXL X:A 2:2.

DNA .
.


DNA. RNA.


. RNA

.


.
RNA D
, 3H- RNA
.



.
21.17 . ,

gfp .
. . Xic, X Xic.

X:A 1:2

. , X:A

2:2 .

. X:A 2:2,

. -

SXL

dsx

mRNA Sxl, tra

DSX-,

dsx -

, DSX-F, -

PE,
SXL

pre-mRNA

Sxl, tra dsx. ,

X:A 2:2 -

. , SXL -

21.21 mRNA

msl-2,

. SXL, msl-2

21.24

, -

X -

. ,

(
)

( )

()

(

)

()

. X

. tra
TRA.
TRA
X:A 1:2,
, -

21.26 mRNA -

. TRA

X:A 2:2, -

dsx, DSX-F. DSX-F

mRNA -

. TRA,
DSX-M,

22

22.5 FeSV -

87

88

22

, -

). -

Southern

GTP, -

FeSV -

. cDNA fes -

1,2 kb -

DNA

, -

cDNA FeSV.

1,2 kb

cDNA -

- -

(1,0 kb),

- fes

(i), (iii)

HindIII,
. - fes , fes FeSV .
Hin dIII /
. -
fes
. DNA
3,4 kb

fes. ,
( 2,4 kb)
Hin dIII.
22.6
, , .
-,

.
22.7 . -
, - -

(viii) ,
. (ii) (vii)
,

. (iv), (v) (vi)
.
3
mRNA ,
. ,
.


-, .
22.8 .
,

.
. -
, ( -

), -

DNA

. ,

. -

. -,

EGF. EGF

, -

Grb2

SOS. SOS GDP Ras,

LTR .

G,

DNA -

GTP.

GTP-Ras Raf-1,

. ,

. ,

, DNA

Elk-1,

DNA - -

DNA.

G GTP,

G0 S .

DNA,

22.12

DNA - -

G0 S
.

22.14 ,

LTR . -

. ,

Southern

. -

DNA --

22.13

89

90

22

BRCA2 -

. -

. -

22.15 , -

, -

, -

. -

RB. -

. ,

RB/rb -

. , -

pRB.

RB. -

, -

, , -

. -

RB/rb . ,

.
22.18 .


. ,

DNA

DNA. ,
BRCA1

,
.

.
22.22 .


DNA
.
, .
, p53.

DNA,

p53

, -

DNA

WAF1. WAF1, p21,

G1.
DNA , p53 DNA WAF1,
BAX, BCL-2 . BCL-2,
.
22.25
. ,
,

. ,
.
,

.
22.26 .

.

.
-
,

.
.

,


.


.

- .
. EWS , ,

- . (
, , .)

91

92

23

. -

DNA

, -

. , -

23

23.1 . -

, -

PCR -

. -

. , -

PCR -

DNA

( ) DNA -

( ),

. -

, -

( PCR

) -

( -

. -

PCR,

).

. -

, -

, -

. -

O2,

, -

. ,

22.28 -

DNA -

. , -

DNA -

. 70S

23.6

50S 30S, 30S -

16S rRNA

DNA

RNA

, , -

RNA -

. -

( fMet).

mRNA rRNA -

23.4

tRNA. mRNA rRNA -

. -

tRNA -

. -

DNA . -

, mRNA

CsCl, -

( U UA

DNA

UAA

). ,

. -

mRNA,

DNA

5 .

23.8 .

, -

. -

2, 5 6 -

, -

, 1, 3,

) DNA, -

4 7 -

(. 23.5).

. -

cybid

. , -

, -

cybid

( , )

. -

DNA

DNA -

. , -

, -

DNA

. , -

. 23.4,

DNA -

DNA

93

94

23

. -

, .

cpDNA.

cybid -

(.. Chlamydomonas),

23.9

(wobble)

cpDNA -

tRNA

, .

. -

, -

cpDNA

tRNA, .

23.11 . -

23.12

, -

), -

(.. ) -

(.. ).

. -

23.15 . tudor -

( ),

tudor.

, -

(.. ). -

23.16 . -

[N ] -

. , -

[CMS ], [CMS ]

.
.

rf/rf [N ] Rf/Rf F1
[CMS ] Rf/rf.

. [CMS ] Rf/rf [N ]

rf/rf . [CMS ] Rf/rf

[CMS ] rf/rf .
23.17 . : [poky ]F [N ]+ . :
[poky ]F ( [fast-poky ])
[poky ]+ ( [poky ]). 1:1
F


[poky] ( ). [poky ]+ [poky ],
F [poky].
. : [N ]+ [poky ]F . : [N ]+ [N ]F. .
23.19 1/2 petite, 1/2 (grande).
23.24 l , l /l +/Y
l/+ () l /Y ( ). F1
(l /+ +/Y )
2:1 (1/4 l /+ , 1/4 +/+ , 1/4 +/Y
1/4 l /Y , ).
, F1
( )
.
23.27 Carlos.
Escobar Sanchez,
Carlos. Mendoza Carlos. -

DNA Carlos.
. RFLP .
Carlos
DNA
.
Escobar Mendoza
RFLP,
Carlos.
. Carlos
( Escobar Mendoza)
.
. Sanchez .
,
.
. Escobar Mendoza

RFLP ,
. RFLP ,
, Carlos

RFLP.
23.30 F1
dd. ,
d.
,

Dd.
Dd dd.
24
24.1
HardyWeinberg. p = f (C B), q = f (C P) r = f (C Y).

95

96

24

: f (C BC B) + f (C BC P) + f (C BC Y) = p 2 + 2pq +

2pr = 236/500 = 0,473

: f (C PC P) + f (C PC Y) = q 2 + 2qr = 231/500 =
0,462
: f (C YC Y) = r 2 = 33/500 = 0,066

2pq = 2

0,4 0,6 = 0,48 = -

. , 0,48/[(2 0,16) + 0,48] = 0,48/0,80 =

60%
.
. q 2 = 1% = 0,01, q = 0,1, p = 0,9

, p, q
r p + q + r = 1:

2pq = 0,18 . ,

0,18/[0,18 + (2 0,01)] = 0,18/0,20 = 0,90 = 90%

r = 0,066, r = 0,066 = 0,26

24.5 2pq/q 2 = 8. , 2p = 8q. , 2(1 q) = 8q

2 = 10q, q = 0,2.

q. , q + 2qr = 0,462 r = 0,26,

q 2 + 2q(0,26) = 0,462.
q 2 + 0,52q 0,462 = 0
q:

24.8 . S s p q

, :

q=
q=

0,52 (0,52)2 4(1)( 0,462)

2(188)SS + 717Ss

2(3.146)
= 0,467

2(1)

p=

= 0,1737

6.292

717SS + 2(2.241)ss
2(3.146)

1.093

5.199

= 0,8263

6.292

q :

q 2 + 2qr = 0,462
r 2 = 0,066

, :

q 2 + 2qr + r 2 = 0,462 + 0,066

(q + r )2 = 0,528
q + r = 0,727
q = 0,727 r = 0,727 0,26 = 0,467
p + q + r = 1:

p = 1 (q + r ) = 1 (0,26 + 0,467) = 0,273

24.4 . 0,16 = 0,40 = 40% = : 1 0,4 = 0,6 = 60% = -

SS

188

Ss
ss

d 2/e

95

+93

91,0

717

903

186

38,3

2.241

2.148

+93

4,0

3.146

3.146

133,3

(df ),
:

p q (df = = 3 2 = 1). 2 133,3

: P < 0,0001. ,


Hardy-Weinberg.
24.11
,

24.30 . -

. -

Bam HI -

q,

. -

q.

4,1 kb

0,4 -

Bam HI

. -

(0,4) = 0,16

, -

. -

4,1 kb. -

. -

, -

2pq = 2(0,6)(0,4) =

Bam HI 6,7 kb.

0,48.

24.13 64/10.000 . -

0,0064 = q q = 0,08 =
2

. + -

2(6) + 38 = 50

24.16

2(56) + 38 = 150

q=

u+v
=

. , f(+) = 0,25 f() = 0,75.

6 107

. -

(6 107) + (6 108)

6 107

(6 107) + (0,6 107)

Hardy-Weinberg, (0,25)2
0,0625
4,1 kb. 6,25 -

= 0,91

6,6

, 6 .
(0,75)2 0,5625 6,7 kb.

, 0,0081 AA,

56,25 ,

0,1638 Aa 0,8281 aa.

56. , 2(0,25)(0,75) 0,375

24.20 px = mp + (1 + m)p

px = [20/(20 + 80)](0,50) + {1 [20/(20 +

80)]}(0,70) = 0,66
24.23 .
HardyWeinberg. , : 0,25 AA,
0,5 Aa 0,25 aa.
. q = 0,33.
. q = 0,66.
24.25 . q = 0,63.
. q = 0,64.
. q = 0,66.
24.27 q = u/s = (5 105)/0,8 = 0,0000625.

37,5 ,
38.
,
2 .
24.32 .
1 ( ).

p1, p 2, p 3,, pn,
p12 + p 22 + p 32 + pn2.
G1A,
(0,398)2 + (0,240)2 + (0,211)2
+ (0,086)2 + (0,036)2 + (0,016)2 + (0,007)2 + (0,006)2
= 0,270

97

98

25

1 0,270 = 0,730.

. -

0,741

G10X, 0,740 G10C 0,662

G10L. . , 2

Hardy-Weinberg. Hardy-Weinberg.
. .
G10X G10L, (G10X: X133, 135 X137, X141 G10L:
L155, L157, L161). ,
2 .

. Ne = (4 Nf Nm)/(Nf + Nm), Nm
.
24.34 . . ,

.
.

Hardy-Weinberg.


. ,


.

.
. ,
.
.
.
24.38 ,


, RFLP
DNA . (),

. (),

. , .
25
25.1

.
, 135

(45 3 -

), 135
. 9 135
( 6,67%

).

(.. -

(0,00% -

),

). -

DNA, -

, 98 135

, -

(72,59% -

). -

25.7 -

. , -

25.2 K = 3/4 ln[1 4/3(p)]

. , DNA -

K = 3/4 ln[1 4/3(0,12)]

K = 3/4 ln(1 0,16)
K = 3/4(0,17)
K = 0,13
25.3 r = K/2T
r = 0,13/[2(80.000.000)]
r = 8,1 1010
25.4

.
DNA,
. .
25.5 1 108 //.
2 108 //.
25.6 DNA ,

DNA .
25.8 ,
. ,
. ,


.
25.9 ,
.
25.10
.
25.11
.

99

100

25

RNA -

, -

. -

25.15 -

25.12 dHR = 6, dHD = 10, dRD = 11

dA1 = (dHR + dHD dRD)/2

dA2 = (dHR + dRD dHD)/2
dA1 = (6 + 10 11)/2
dA2 = (6 + 11 10)/2
dA1 = 2,5
dA2 = 3,5
dA1 dA2 ,

.
.
25.13
.


,
.
25.14 DNA
(
, ). ,

. ,
-

.
,

.
25.16

.

Nrooted = (2n 3)!/[2n 2(n 2)!]

Nrooted = 9!/(16 4!)
Nrooted = 362.880/384
Nrooted = 945
Nunrooted = (2n 5)!/[2n 3(n 3)!]
Nunrooted = 7!/(8 3!)
Nunrooted = 5.040/48
Nunrooted = 105
105
945 ,
.
25.19
,
.
25.20
.

25.21

. -

, -

, ,

. -

101