:

:
IV - 4

, 2008.

IV - 4

____________________________________________________________________________3
______________________________________________4
______________________________________________________________________________5
_________________________________________________________________________________5
____________________________________________________6
_______________________________________________________________________7
____________________________________________________8
____________________________________________________8

________________________________________________________________9
_______________________________________________10

___________________________________________________________________________11
________________________________________________11
___________________________________________________________________12

__________________________________________________________________12
_____________________________________________________________13

__________________________________________________________________15
____________________________________________________________________16
__________________________________________________________________17
___________________________________________________________17
_____________________________________________________________18

, ____________________18
____________________________19
________________________________________20
_______________________________________________________________________21

IV - 4

( )
, .
,

. .

() , . ,
,
.

IV - 4

, ,
. ,
,
. , .
, .

,
.
. 20
.
.
, ,
.
.
,
. , ,
. ,
, .
, . 1838.
,
1958.
X-.

.

, . .
, . AUG
. ,
64.
() , -. .

.
. [6]
.
. -
. N- C-.

IV - 4

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

( )
.
.


.

.
.
, ,

.
, .
.

,
. ,
, .
440 , 220 .
,
. ,

. , . ,
.
( ) 2 ( ,
).
.
5

IV - 4

. , .
5'( ) 3'( ).
5' 3',
3' 5'.
. (),
(), () ():

.
: ()
() , , ( ).
() () ( ),
. 36,
(- -).
.
3,4 nm (). ,
1953. .
- ( ) ( ).
() , .
.
, . ,
, 1
.

IV - 4

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

.

.

. , .
.
.
,
, .

, .
, .
7

IV - 4

,
. ,
.
,
.
. , 2.5 .
.
, .
. ,
.
: H1, H2A, H2B, H3 H4.
. (
H2A, H2B, H3, H4), .
, 166 (- -).
H1, ,
( )
.
.
.

.

IV - 4

. ,
, . , ,

. ,
.

.
.
.
.
, .
(, ) ,
. - ,
2 1 .

.
.
.
() .
,
(. semi= ; konservativan =).

(Escherichia coli).
(N15) .
N14
. N15 N14
.
,
5'-> 3' . ,

.
. 1000-2000 ,
10 .
, ()
.
( Y).
: , .
, ,
.
,
.

IV - 4

oriC . -
oriC - (--).

.
, , .
. ,
75 100 .
.
,
. , ,
,
.
,
(- -).

. .
, ,
. ,
.
, 3' 5'
5' 3' ().
,
5' 3' 3' 5'.
.

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

3' 5' ,
(
)
. ,
, ,
. ,
,
(- -).
.
10

IV - 4

,
.
.
850
, 150
.

( )
, , ,
.

.
, .
,
(= ; =).
.

, :
* , -
* - -
* -.
()
, . .

. - ()
( ). -
, - .
.
: -
.

. .
- -
- .

11

IV - 4

()
.
4 : , , .
() , ,
( ).
,
. ,
, ,
.
, (
, ),
, .
,
.
. ,
, , .
( ),
.

. .
,
, .
() (G) .
(C) ()
. (U)
.
, , 64
43 = 64. , 5'- UUUAAACCC -3',
UUU, AAA CCC, .
, .

12

IV - 4

64 .

C
A
G

UUU (Phe/F)
UUC (Phe/F)
UUA (Leu/L)
UUG (Leu/L) , Start
UCU (Ser/S)
UCC (Ser/S)
UCA (Ser/S)
UCG (Ser/S)
UAU (Tyr/Y)
UAC (Tyr/Y)
UAA Ochre (Stop)
UAG Amber (Stop)
UGU (Cys/C)
UGC (Cys/C)
UGA Opal (Stop)
UGG (Trp/W)
C
CUU (Leu/L)
CUC (Leu/L)
CUA (Leu/L)
CUG (Leu/L) , Start
CCU (Pro/P)
CCC (Pro/P)
CCA (Pro/P)
CCG (Pro/P)
CAU (His/H)
CAC (His/H)
CAA (Gln/Q)
CAG (Gln/Q)

13

IV - 4

CGU (Arg/R)
CGC (Arg/R)
CGA (Arg/R)
CGG (Arg/R)
A
AUU (Ile/I) , Start2
AUC (Ile/I)
AUA (Ile/I)
AUG (Met/M) , Start1
ACU (Thr/T)
ACC (Thr/T)
ACA (Thr/T)
ACG (Thr/T)
AAU (Asn/N)
AAC (Asn/N)
AAA (Lys/K)
AAG (Lys/K)
AGU (Ser/S)
AGC (Ser/S)
AGA (Arg/R)
AGG (Arg/R)
G
GUU (Val/V)
GUC (Val/V)
GUA (Val/V)
GUG (Val/V) , Start2
GCU (Ala/A)
GCC (Ala/A)
GCA (Ala/A)
GCG (Ala/A)
GAU (Asp/D)
GAC (Asp/D)
GAA (Glu/E)
GAG (Glu/E)
GGU (Gly/G)
14

IV - 4

GGC (Gly/G)
GGA (Gly/G)
GGG (Gly/G)
1 AUG -
.
2 .

()
,
. ,
5' 3' . ()
(
), ,
:
, :
*U
*GC

,
,
.
.
- .
-, -,
-. , -,
.
(
):
* : 3' CCCG 5'
* : 5' UUUGGGC 3'

15

IV - 4

,
. (-)
.
. ,
:
*
*
*
, ,
, .
, , .

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

,
. ,
, .

-, -
. 98 % ,
. , N-
, . f-. f-,
, .
, f- .
,
- -8 -13 .

16

IV - 4

50S 30S ,
. (-)
-.
.

,
. ,
, - ,
.

, , -
. - 5' 3' --
. ,
.

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

( ).
, :
*
* .
:
* ( -)
* (-, -).

17

IV - 4

.
.
:
* ,
* ,
* -
.


.

1955-1963
4 Escherichia coli
:
1. - .
2. -
3. -
, :
-,
-,
e.coli,

.
e.coli ,

.
, (z,y,a). .
.
.

18

IV - 4

, ,..

,
.
,
. .
.

.
,,, , .
,. .
. ==
+ .

. .
. () ,
.
. , ,
.

19

IV - 4

. :
*
, ,
, .
* :
o
o
o

,
.
( )
.
.
,
() .
:
* ;
* ;
*
, ,
.

() .
. .
:
*
; ()
*
* .
;
*
;
* () ()
;
* ,
;
* .

20

IV - 4

, ,
:
1. ,
.
2.
3. ,
, .
4.
.
5. ( , ,
)

. ,

.
.

.

,
. ,
, .

21

IV - 4

1) . , . , 3. 4.
,
2) www.google .com ( )

22

IV - 4

:
: ______________
: ______________
:
______________

__________ (__)
__________ (__)
__________ (__)

:
______________

23