. , .

- .-. . .

- . . .

2002

FROM CELLS TO ATOMS

AN ILLUSTRATED INTRODUCTION TO
MOLECULAR BIOLOGY
ANTHONY R. REES
MA, DPhil
Lecturer in Molecular Biophysics and
4itor in Biochemistry, Christ Church,
University of Oxford

MICHAEL J. E. STERNBERG
MA, MSc, DPhill yal
Society University Research Fellow
Department of Crystallography,
irkbeck College, University of London

Foreword by
SIR DAVID PHILLIPS
FRS
Professor of Molecular Biophysics
University of Oxford

Blackwell Scientific Publications

Oxford London Edinburgh Boston
Palo Alto Melbourn

 577.2
28.070
54

54

., .
: :
. . .: , 2002. 142 , .
ISBN 5-03-003521-4
.
44 ,
, .
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577. 2
28.070

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26.12.01 . 84 18'/|. .
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.-. . 16,44. . 4/9797. 5000 . . 5914.

,
. 107996, -6, , 1- ., 2.



.
143200, . , . , 93.

ISBN 5-03-003521-4 (.)

1984 by Blackwell Scientific Publications ISBN
0-632-00888-1 (.)
, , 1988, 2002


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- .
(...)
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,
.

Part of Fig. 4.1 from Branton D., Klug A. (1975). J. Mol.

Biol., 92,
559-564, various plates and figures. Part of Fig. 4.1
from Marvin D. et al. (1975), Nature (Lond.), 243,
19-23, fig. 5. Part of Fig. 5.1 from KJug A. (1960).
Advances in Virus Research,
vol. 7, p. 274.
Part of Fig. 5.1 from Wilson I. et al. (1981). Nature, 289,
336. Fig. 6.1, 8.1 and 11.1 I. Geis from Dickerson
R.E., Geis I.
(1969). The Structure and Action of
Proteins,
Benjamin/Cummings, Menlo Park, California. Fig. 9.1
I. Geis from Cantor C.R., Schimmel P.R. (1980)
Biophysical Chemistry, Part 1, fig. 2.23a, b, W.H.
Freeman,
San Francisco. Fig. 10.1 J.S. Richardson in Anfinsen
C.B. et al. (eds.) (1981).
Advances in Protein Chemistry, vol. 34, figs. 73,
74, 77,
Academic Press, New York. Fig. 10.2 courtesy A.C.T.
North from Imoto T. et al. (1972), fig. 2;
in Boyer P.D. (ed.). The Enzymes, vol. 7, 3rd edn.,
Academic
Press, New York.

Fig. 14.2 adapted from Johnson L.N. et al. (1968).

Structure Function and Evolution of Proteins.
Brookhaven Stymposia in Biology, No. 2, Fig. 2.
Fig. 15.1. after Perutz M.F. (1978). Hemoglobin structure
and respiration transport, Scientific American, 239, No.
6, 80; and Baldwin J., Chothia (1979). J. of
Molecular Biology, 129, 175-220, figs. 1.2.
Fig. 18.1 from Metzler D.E. (1977). Biochemistry - the
Chemical Reactions of Living Cells, fig. 2.23a,
Academic Press, New York.
Fig. 19.1 from Holbrook S.R. et al. (1978). J. of Molecular
Biology, 123, 631-660, fig. 4.
Part of Fig. 26.1 from Worcel A., Burgi E. (1972). J. of
Molecular Biology, 71, 127-147, fig. 12.
Part of Fig. 26.1 and 26.2 from Kornberg A. (1980).
DNA. Replication, figs. 9.5 and 9.11, W.H. Freeman,
San Francisco.
Part of Fig. 34. 1 from Henderson R. et al (1975). Nature
(Lond.), 257,28-31.
Fig. 37.1 after Stryer L. (1981). Biochemistry, 2nd edn., fig.
34.8, W.H. Freeman, San Francisco; and Cohen
(1975). The protein switch of muscle contraction,
Scientific American, 233, No. 5, 39.
Fig. 39.1 part from Taylor D.L. et al. (1982).
Philosophical Transactions of the Royal Society of
London, Series B, 299, 185-197, fig. 4; and part from
DePamphilis M.L., Adler J. (1971). J. of Bacteriology,
105, 395, fig. 25.
Fig. 40.1 from Amzel L.M. et al. (1974). Proceedings of the
National Academy of Sciences of the USA, 71, 14271430, fig. 4.
Part of Fig. 41.1 from Schrader W.T. et al. (1981). In:
Grcep Roy (ed.). Recent Progression in Hormone
Research, vol. 37, p. 620, fig. 27, Academic Press, New
York.
Fig. 43.1 from Quigley G.J. et al. (1980). Proceedings of
the National Academy of Sciences of the USA, 77,
7204-7208, figs 4, 6.

Parts of table 44.1 and worked example from Price N.C.,

Dwek R.A. (1979). Principles and Problems in Physical
Biochemistry for Biochemists, 2nd edn., Oxford
University Press, Oxford.


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16 N- 25 -
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,

0,35 , , Ala Ser, 0,57 .

, .

, , , Arg,
Asp Glu. - , .
,
, , . ,
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30-
.
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,
(coiled-coil) , . 11
,
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, (. 36)
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1010
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,
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S (
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G: , ,
.

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1.
.
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.

S-, -

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S .
,
- -

.
.

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13.

,

-. ,
S
1. -
S S 2 3 :

, ,
,
. v
[S] :

Vmax - , , ,

KM , , , -

. KM ( , 3 2),
.
v [S] .
-
.
- , .

1/v 1/[S], KM/Vmax, 1/v

1/Vmx.

Vmax KM

, .
(. 13.2).

-,
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,

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Vmax . ,
,
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(. 13.1). . ,
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tense - ). R- , -
, R- ( ).
.
- R.

.
v [S] S-
(. ).
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.

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, R-KOH, .

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

14. :

 , , .
, ,
.

,
N- (NAG) N- (NAM). , - 1 -,
1 - 4- (. 31).
.
, 129 (. 10).

, D .
, D
, ,
, , , (. 14.3)
35
52.

( 0,3 )
, D . Glu 35
,

(. .

. 14.2. .
(NAG-NAM)3 . NH- - . . rf 62
rp 63 , .
.

, . . ,
. , , , , ,
( ,
, , D, F) (. 14.2).

. 14.3.

). Asp52, , ,
(. .
).

.
1. - Glu 35
, ,- ,
D.

, ,
F,

.
2. , D-,
,
.

.
-.

Asp 52.
3. (),
,
-,

. - -

(+) Glu 35,

.
4. .


1.
NAM,
D.
2. Glu 35,
.
3. Glu 35,
.
4.
-
Asp 52.

15. :

- ,
.
. - -
( , 2) -

( , 2).
.
,
.

.

-, , -

15. :

- ,
.
. - -
( , 2) -

( , 2).
.
,
.

.

-, , -

 (. 16). 140150 .
,
.
- -
.

-.
- , ,
, , F, G , 10
25 (. 10).
. -, D,
, . -
,
.
-
. .
,
-. , HisF8 -
- F, ValFG5 - F G, 2
- , - .
, ,
, . ,
.
His.
(HisF8)
. .

(HisE7)

HisF8
-

.
,
.
,
,
,
.
(, - 2 ValFG5 -), , (,
LysC5 -
- -).
, . .
.
,
- F- (. 15.1).
1 1
2- 2 -.
, .
,
. 15.3.

( ).
-

,
.
S- ; , . . ,

.
S- , .
100% 75%. ,
, ,
.

, (. 13). , ,
,
.
, .

, .
, , - (tense-state), R-
(relaxed-state).
T- R-
,
R. -

-.

.
- R-.

. R-,
,
. ,

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

16.

, . , .

.
, , , F10 - (- .

,
. ,
.
,
F8 His, .

,
.
-

 ; , F6 . ,
, , . ,
,
, .

, ,

.
, . , - - 155
(. 16.1).


.

( )
,
, , -
/- 47,
- /- 24.

(. 27).
,
, .
.
, .
, , . . , - .
.
.

. ,
, . ,
, . -

.
,


.
, ,
, . - - ,
, ,
, - - , .
.
.

, -. , - , - . , , .
,
-, . ,
. ,
, ,
, , , .

,
1%
,
.
( . 16.2),
,

 , , -, . IV , ,


. IV (1% 600 . ).
, . , , (1%
1 . ). , 105 , , , .


.
,
IV. , ,
. ,
, ,
, , 1200 .
, 100 . .
,
-
.

.
,
. , , , ,

. Glu Val
. -
. Glu

Val.
, , .
, - Val,
.

- -

.
, ,
,

,
(. 15).

,

, , ,
. ( ),
, .

17.

 -
() ()
,
. .

, () .
.

(). , .
,
,
: -D- -D-2-.
,
() 2'
. , ,

, ,

, .
- .
( ) (G)
(), () (U).
A, G, U, A, G, .
-N- , 1- N1-
N9- .
,
5'- . -5'--, -5'-- -5'- , XDP
, X - .
, .
( 7) . , , XDP
, .
, (
5'-) .
17.1. -
, .
(ATP) .
(2-4),
Pj.
(.
7):

17.1.

()
()
(G)
(U)

(AMP)
()
(GMP)
()

()

(dAMP)

()

(dCMP)

(dGMP)

(G)
()

(dTMP)

ATP, ADP AMP (Mg2+ 2+).

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

 (. , . 6). , ,
,
5'- 3'- .
, 5'- 3'- .
. ,
5'- , ACG; , , : pApCpG.
() d (): dAdCdG. , ; , .
, , :

18.

,
.
, , - - 1944 ., ,
.
(, S), , ,
( R), . ,
, S-,
R-
, R- . , .
2, 1952 ., -

.
, -
. coll.
32 ( ) 35S ( ) . coll. ,
, 32. , , , , .

1953 .
, , 1
; - . 1

. . .

 . , ,
.

,
- . -
,
. ,
,
. ,
1`-
AT- CG-nap
1,085 . AT- GC- -
.

. ,
, ,

. ,
- , , .
()
(. 20, 21).

, , .
50- ,
. ,

,
, .

.
,
0,34
36 .
, , 10
(360/36 = 10), 3,4
(10 0,34 ).
20 .
. , , . .
,
1950 1953 .,
.
(.
18.2) (), - .
3,4 , . . ,
10- 0,34 , . . . - (. ).

. 18.2.
. .

.
. , ,
.

,

-
.
, ,
. ,
,
.

. 1 -,
- Z-. - - , . -
20
.
11 . -
-. Z 12
. Z
. . -,
1

, -

. . .

 -, Z-
1.

()
,

(. 26
28). -:

.

.
(, ) , .
.

. II,
() ,
46 (. 30). ,
. , R1 . coli , :

, ,
, , ,
: ,
5'- '-, .
2- ,
.
5`-
' Z-,
. . .

32

4 .
, .
,
,
G ,
. ,
, ,
5'-
, . ,
G 1, 5, 7 19
20 , ,
, 5, 7 19 ,
G.

.
,
, (
).
, , , . 19.3.
.

(. 20, 21).
- I (. 20, 21).
, .
( 2'3'-),
.
. ,
.
- . . .
, -, .

19.

. 19.1.

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

().

(. 5).
() .
.

( )
.

.
-

(. ). (. 24) , .
(S)-
, (. 44). , . . coll
. 19.1.
()

.
, -

19.1. . coll

, %

, S

80

23
6
5

15
5

4-26

. ,

1 000 000
500 000
35 000
25 000
25 000-1000 000

3000
1500
100
75
75-3000

'- .

,
. ,
Phe .
. 75 ,

. = 25000
4S.
;
PHKAla

1965
.

5'-; G. '-
' () .

, , G U,
,

,
.
, h

( ), (D)
, Y. ,

( 3) ;
-

m 2
,
.

, -

. ,

, ,
.

(,
D,
) (D,
).

(,
PHKPhe).
PHKPhe
.

. - .

, -

,
,
.
,
- .
- 11
, 3,1
.

0,28 (3,1 /11);

33
(360/11). ~20
. -
-- (. 18).

-.

2' ,
.
.
.
,
. -
,

; D .
10 .

.

. ,
,

,
, ,

-.

2'-

.
- ,
.

-
, ,
,
(.
19.2).

. ,
,

,
.
, -,
- .

.

,

.



100 (
1965 .; . ). .
.
. 5' '- 324.
, . - ,
, , (. 18).
,
, ,
. 18, -
.
.
, . . ( ),

-. ( ,
; . . 30.) , ,
- 3'GGGUCA. 5'-GCCAGT. (. 18),
, . 19.3.

20.

, , , . ;

. 4.


, .
. ,
, . .
. 20.1.

1958 . , .
,
( I5N), ,
, ()
(14N).
. , ,
. , I5N, 14N,
.
,
G ,
- . ,
, , .
, , -

. 20.2.

, .

() (, . coli)
.
: . coli
1700 1 ,
40 .
, ,
50 1 .

, . . , .
, , 65 106 , . (. 20.2).
- ,

.
.
.

.

. (. 21). .
, . coli, , . -,
.

. . coli
, , .
, -.
; , ,
174.

-. ,
,
. ( . 20.2)
,
'- () . 5'-
,
.

21.

E.coli - ,
. . coli
,
.
. , .
- ,
.
- '- , , , 5' > 3'. , 5' >
'- . : 1) -, -, ;
2) (ATP, CTP,
GTP ) 3) '-- , . . 21.2.

- I, II III, polI, polII polIII. polI ( 1958 .), , ,

polIII. PolI

(. ), . 5' > '-

, > 5', . . > 5'-. Poll pollII 5' > '- .

,
. .
.
, ,
( ). - 5' 3', , ;
. , ,
, (), . .
, ,
, , .
1000 , 100200
.
- . -
'--, , .
,
,
(). -, '--, ( 1060 ), .
-
, '-- ;
- III. , ,
, - . -
poll.
- 5' , 5'- -

, . - - I, > 5'-.
,
5' > '- . , - , .

( ) ,
.

, 109
. ,
'-

.
polIII
, 3'5'-
, ,
.
, pollII
.

,
. . , rep-,
. , ,
.
: , , ,
, - ,
.

,
,
.
,

,
,
.
,
, ,
(. 26). - . coli

.
, ( ; . 22) -
.

22.

. - ,
- , . , +- . .

- - ,
. : , , '
. (2')
. () 500 000. , -, -

-, (
GTP) ,

,

.
- ,
40 .
G +- . ( 3' - 5' +-)
10 .
+- ,

 , ,
.
TATPuATPu, ( G). ,
.
40 -,
, , - .
,

. ,
-
- (2'), . ,
-
. - -,
, , -.

. . 22.2
'-.

.
.

.
, . . . 5'- '-.
-,
20% , . , 5'- 3', ,
Q . - ,
.
. '-
,
. (. 19),

(, , G U).

.
. . coli (16S, 23S 5S)
(. 24)
.
. Q;
, , , .

( ) , .
GC-, -. GC- . , +- ,
- -
.
.

,
--

- -. I
,
, 18S, 5,8S 28S. II , , . III 5S-pPHK.
, ,
, . ,
. ,
, ,
, G.

-,

 (). 5'- '- .

.

, ,

() (. 27).

,
.
,
GU GA AG - .

5'- , . '- poly(A)

150-200 .

, .

,

,
18S-, 5,8S- 28S- (. 24),
. ,

.

23.

. 23.1.
-
.
, . .
, . ,
20 ;
, 4 4 = 16 , 4 4 4 = 64 .
. .
ABCDEFGHI , ABC,
1, DEF 2
. . , ABC 1,
CDE - 2 . .
,
(. ). -

, . . ,
.

5'- '- , +- ,
5' -* 3'. 5'- N- . , N- -.
, (. 23.1).
. 23.1 ( 5'-) , , (
'-) .
64 61
(, CUA Leu),
(.). -, . ,

AUG Met GUG Val , N (. 25).

, Met Val.
23.1

Arg, Leu, Ser

4
3

Ala, Gly, Pro, Thr, Val

He (.)

Asn, Asp, Cys, Gin, Glu, His, Lys,

Phe, Tyr
Met, Trp

, . . (. . 23.1).
, Phe , UUU
UUC. , , -. , , ,
, , ,
, . , ( U) ,
( G) .
(, UCG AGU
Ser).

. , , ,
.

(I),
U, , (.
23.2).

. 23.2.

23.2

U,
G
U

U
G

. 23.2.
64 .
.

. U

(Phe, Leu, He, Met Val).
,

(, His, Gin, Asn, Lys, Asp
Glu).

( ).
-,
.
,

,
(. 27).
, -, - - G4 (.
23.1).

.
.
,
,

. , .
(-) ,
.
, (. 16)
Glu Val (3- . GAA GUA, . .
U .

()
, .
- .
,
,
-

 - . ,
, ,
(1981 .)

. , UGA , , AGA
AGC Arg, .
60-
, .
. , ,
, , -- (. 24).
poly(U) (. . UUUUUUU...), . , poly(U) poly(Phe)
(. . Phe, Phe, Phe...); , UUU
Phe. ,
Pro, Lys.
-

, ,
, .
, ; , , , 2U 1G, Cys, .
,
.
,
. ,

- (. 24). ,
UGU
- Cys. , UGU
Cys. , UAA, UAG UGA, -.
,
.

24.

. 24.1.

,

.
; . 25.

, -(1)-
, ()-.

5' > 3'.

()
(. 19).


: -.
. -

. (, , )(. 25).

'- -. -,
-.
. :

- () 2'- '-- (. 24.2). -

. 24.2.

: ,

, . , ; ,
,
, . . .
, ,
.

. ,

. 24.3.

- . . coli 70S,
, , 80S. , ,
70S, .

in vitro Mg2+.
708- 50S- , SOS- - 60S- 408.

(. 24.3) . :

2:1 1:1.

.
,


,
(. 19).


Arg+ Lys+. , -

 ,
.


in vitro. ,


.
. . 24.4 708-
. coli. 20
30 . (80S)

, 1,15
( 23 35 ).
, , . (. 24.4), . ,
-
, {) .
, ,
-,
(. 25).
. ,

,

(. 24.1).
,
,
,
, .
5'- '-.
.
,
,

.
;
.

25.


. colL

AUG,
(Met) [ GUC -
(Val)].
Met, Val. :
AUG GUC,
? Met ( Val) (. ).
( fMet) Met, . (), Met
. , (
AUG,
AUG.

30S-cy6, , - - fMet, . 508-,
705- .
GTP GDP Pj.
, (IF1, IF2 IF3).

.
: 1) , 2) 3) (. 25.1).
- . ,

, EF-Tu,
EF1, GTP.
TPHK-EFI GTP , GTP
GDP . EF1 GDP,
, , -. EF1
, EF-Ts, EF2,
GDP , EF1 EF2. ,
,
. 25.2.

. 25.2.
,
, ,
-. 508-
, ,
. 25.3. , ' (. 25.3).
,
, EF-G(EF3), GPT.
, ,
, - , ,
'-. ,

.
. . ,
UAA, UGA UAG.
, -

 . , ,
, . ,
, . . 30S- 505-.

. i K\i!( VIMIUIM
,
, , -

.
,
Met, fMet.
,
AUG: ,
Met, .

. , (80S).
. , , 708-
. fMet.

26.

.26.1.

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

4,6 106 ( 4600 , ...).
1 , , . coli 2 ,
(),
.
, ,
, ,
;
, .
40 , 40
000 , . coli.
700 , . coli.
, , , ,

.

, .
400 () 100 000 ... ().
,
(1,74 ) ,
.
-, , ,
.
(, , ; . 5) ,
-, .
, , 1 ... . 26.1
.
,
10 000 ,
(. . )
.
,
, .

26.1

(
)

SV40 ( )

5,0 103

4 ()
. coli ()

2,0- 105

200

4,6- 106

4600

2 , 8 - 109

100 000-500 0001

,
. 4 , .
,
; .
(. 4). , 4,
.
()

(). , , ,
,
(), .
26.2 .

, ,


.


,
. 1

. . .

 ,
. ,
, , ,
, .
26.3. , , , (, SV40).

,
, ; ,
, , . coli.

(.
)
,
,
12 .

.
,
, ,
, , ,
.
,
.


(),

.
,
',
.
, ,
.
, , . ,
, 10 (, . coli).
56 .
200 , -

,
. , ,
. 26.1. ,
. . (/) 11 300 21 000.
: HI, 2, 2, 4.
2, 2, 4.

,
. HI ,
;
(
) . HI, -,
, ,
, ,
, -

.

2030 ,

(. 29).


.
6 .


40
. ,

.
5 00010 000 ,
,
100-200 [40 100(200) = 5 000(10 000)]. , ,
,
.

,

.
.

27.

-
, - . 100000 (
).
, 10 . , , -

- (. ). 56 , 0,1% .
2000
3000 ,
.

 : , () . , , , .
,

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

.
( )
- , ,
, .
()
, .
,
. , . coli
,
( )
. . 27.1
: , .
, .
, .

,
, .
lac- (. 28),
(Z, Y )
,
. , , - (I), , ,

lac-. , ,
-

, ,
(. 28). .

.
100 .
( ) ,

, .

10, .
.

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

. , .
,

,

, .
- (. 5).
( 2, 4, 5 6),
- ( 174).

, ,
- ,
,
.
,

.
(. . , . coli) 174
1977 .
, , ( , ) ,
(
). ,
,
, . ,
, , -

 fMet.
,

.
. ,

, , ,
.
, . ,

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

. 5S-PHK
, , . .
, . 5,8S-, 18S- 28S-pPHK ,
.

()
.
.
,
1020 , .
- , , . . , , .
1977 . Drosophila melanogaster.
; ,
8 , .
-
. -
() ,

. .
, ,
. ,
. . 27.2;
-.

(. 22).

. 27.2.
( 10 200
), . .
.

,
;
,
,

.
, ,
SV40,
( 1488 1601 - ,
), :
VP1 VP2. ,

( )
,
,
. SV40
. ,
TL (
: L = large = ),
. (,) 246
, (2) 1879. 345
TL ,

.

28.

,
. , , ,
, , , .

, , ,
.

- , ,
. -


.

1961 . ,
, .
( )
(- ,
, 100 . .
: (,
) . , ,
Z, Y
lac-. ,
, .
- ( I) lac-
Z, Y .
I .
, I
(I--), ,
Z, Y
, . .
. I
, I,
Z, Y
. , I - , .
, . lac-
lac-.
, .
() ,
. - ,
. ,
, .
2- ,
. 28.2,
lac -.
() -
. . ()
, ,
, -

.

,
,

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

( ).
, ,
. ,
.
.

(trp-), ,
.
, ,
.
- ().
( 1 . 28.1).
,
. , trp
-
.
, (trpE) ( 2 . 28.1).
, , . trp-
90%

 140 trp-.

( ). , ,
. , , ; ,
. , -,
(. ). (
) , ,
, lac. .


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

,
,
. ,
, -
, ,
. , ,
, ,
-. , ,
.

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

,

,
. , ,
, -
,
.
, . .
5'- ( ,
) poly () '-- (.
22),
, .
,

. ,
, , .
,
. ,
.

;

. ,
,
,

-,

,

.
.
,

(. 41).

29.

 ,
,
.
. ,
,
, .
P

,
,
. ,
. - , ;
.


( 40 ). ,
,
,
. ,
, (20220 ).
. 29.1
. ,
.

,
. ,
,
,
.
,
.
,
.

.
- ,
, . , ,
. ,
,
,
-.
,
. , , 20 .

,
24 .

,


.
: Go, G 1, S G2.
G2, .
() , , , , .
, ,
.
Go . . , Go, , . .
.
:
. G o
,
,
(, AMP).
, Go,

, G o G1.
G1 ,
,
S. G|
( ). , ,
G,
. -
. 8 ,
. .
G1
S-.
S ,
;
. S 68 .
,
, (3-). . , S-, 3 - ,
, .
, S-.
S-, , , D, -

 -,
(. 43). , .
G2.
G2 ,

, . . G2
. G2
,
. G2
26
. G2
. G2
.

, , . .
,
, , , . (. 39), ,
. , -

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

.
. , ,
, . .
,
, . , .
.
Go G1, .

30. :

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

(. ) , .

, ,
,
. . , -

,
, - (),
, , , .
, .. ,
, -, .
, . , ,
,
,
, - , (. 27), . .
() ()
, -

 . , (3- ,
. , , .
.
, ( , ,

25 000 , 25 ...), -.
- ,
, ( - ), - .
, ( 10
) -;
, , oligo(dT). poly(dA),
'-
(. 22).
().

-
.
,
-.
(), . , 5'- ,
() . , , ,

.
() - , - I.
, ,
,
. ;
S1.
, ,
, -. .
- ,
,
. -

: .
.
,
- (. 26).
. ,
.
, (+-), -, .
, .
.
- (. 18) -
,
(
- );
. ,
,
.
-
,
.
, pBR322,
,
;
,
;
, .
,

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


. ,
. , , .
,
, .

3'- , , -

().

, , ,

.

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- ().
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43.

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).
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44.

s ; ,
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: .
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.

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s, .

, . 7.
1 . (
-
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). (), (1) :

a PV - , .
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.
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),

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2- , , : - () ( ,
. . TS)) , .

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,
(G),
, G < , (
); G = 0 ,
;
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,
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;
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.
' = []/[] ( [] [] -
),
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. Go
,
; ,
+,
1 -1. , [+] = 1 = 10, .
. = 0.
, = 7 ,
G, G', ,
+, -
,
10-7 -1, =
7. ,


, ,
,
, I, II .
., (
).

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, - 28
, , N- (NAG) 14, 31,35
N- (NAM) 14, 35

36
32, 33
2
41
39 ,
() 17
42, 43
4
(ADP) . - 2, 26-30
35

(AMP) . - 2, 35
4, 5

() 2, 14, 29, 35, 42

20, 21, 29
24
18, 30
--
1,2
38
22
---- 21
2325
---- 25
2, 34
37
Pfl 4
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-2 4, 18, 27

4 14, 26, 27
7, 17
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2, 3
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17
Bacillus subtilis 35
41
. coli, 35
4, 5
------- 27
(1) 31
------- 18
1, 3, 6, 36
------- 20, 21
39
------- 2
13
------- 22
12
------- 2325
1214 37,
------- 26
39 - 39 D
Micrococcus lysodeikticus 35
29, 43 37 (Ala)
Staphylococcus aureus 35
6
13, 25 1 , 1 8
1 , 2 , 39 8 - 2-5, 23-25, 28, 29, 41
31 31 -
6-16, 34, 37,40,41
2325 - 23 41
6
------- 40
6
------- 14
30
---- 15
- 10
41
34
30
34
29
37
(-, 34, 36
)37
6, 9, 11, 36
29, 30, 35, 42, 43
26, 36
40 19, 23-25
G- 41
.
rep- . ,
(Arg) 6 (Asn) 6

(Asp) 6
- 22
1, 7
,
21
21

21,26
2,3
43

()6

15
38
26
3, 38 (Val) 6
7
7
--- 8-10, 40
------ 18, 19
( ) 30
16 39
4 - G 4 2 3
SV40 26, 27
-174 4 , 2 0 , 2 7
() 4, 5
4
5
5
5
1, 4
4, 5
() 5
- 1, 4
23, 27
4, 5
1 , 4 , 5
4, 5
4, 5, 20
- 1, 5
4, 5, 26
1, 4, 5
32, 33, 40
, 38
7
7
---- 19, 23
--9-11, 15, 34,40
------- 18, 19
------- 31
------- 20, 21
------- 22
1, 2
2, 39
---- 2, 39
- 1, 1 8 , 2 6
1 , 1 1
36, 37, 39
36, 37
31
11
9 - 1 8 41
23
- 28
28
33

 33
2, 31, 32
4
- 18, 19
15, 16 5
10, 15, 16, 23
15
30
- 28
1, 4, 5, 27
1, 3, 27
-- 1, 2. 27
26
29
30
27
23, 27
27
28

()22
2
31
(Hyp) 11
32-34
31
32-34
34, 38
17,23

() 6
7, 10, 18,
19,40
40
21
(His) 6
16, 26
31 ,41
14, 31, 35
3, 34, 40
33, 34
3, 34
34
3234, 38
(Gly) 6
10, 15, 16
(Gin) 6
(Glu) 6
2, 31, 38
31
33
D- 31
() 3
30
5, 28-30, 32, 33, 37,38,41
1, 3
(G) 17
(GTP) 17, 25
43
2, 3, 37, 39
.

13

15, 16
-D-2- 17

.

17, 18,21,22
17
36
3
(Dab) 35
(Dpm) 35
(D) 19
2/3/- 18
3
39
6, 10, 40
() 15
1 , 1 8
34, 38
38
38
38
; -, -, Z- 18
18
18
29, 43
-
() 19, 30
() 21, 30
26
27
1, 17
18
-- 18, 26
21
4, 5, 20, 21, 29, 30
27
26
18
27
26
19, 22, 28, 30
14, 26
26
- -
22
- . ,
-
-21
-4, 5, 18, 20, 21, 29,
30
10
(V-) 40
40
(-) 40
20, 29
20, 27
() 20
16
2, 39
34
34
() 2
32, 33
.

16
12
34,35
13
- 37
- 9
35
() 6
(1-, 1-)
37
1, 4, 5
40
(Ig, ) 10, 40
G (IgG),- 40
13

42,43
29
14
12, 13
, 28

12
25
20, 21,
29
2325, 28
22, 28
(1 )2 3
28, 30,41
29, 43
26, 29, 39
43
22, 27, 30, 40
4, 5, 18

(, )4, 5, 1 9 , 2 2 25,27,28,39
- 31
7
(+) 2, 3, 6, 14
(+) 36, 38
(Na+) 36, 38
4, 5,
26 4
- (-) 40
- 14
28
12, 13
20
23
31
9, 11
1 5 , 1 6
1, 2
1,3
- 41
- 28, 30
1,
2,4, 5, 26, 29, 32, 34, 35, 38, 39
-- 1,2,38,39
--- 1, 35, 29, 32, 34,
36-39,41
--- 1, 3-5, 29, 32, 34, 38, 39
- 2, 4, 5, 14, 29, 35, 42

----- 2
----- 3, 29, 31
3, 29
--- G (r , G r , S-, G 2 -, - 29
30
28
13
7
--- 6
----- 32, 33
--- 17
----- 31
23, 27
19,23-25
- 23
1, 8, 9, 11
40
18, 20, 21

6,8
- 8
8-11
- 18
- 32, 33
- 8
- 8
- 31
- 19
27
13, 15
28

24
----- 21
- 22
11
1-3, 19,
22, 24, 25, 27, 44
42
31
2
34
3
() 31
- 22, 28
13
- () 42
- 42
28
(lac-) 27,
28
2
32
(L-) 40
(Leu) 6
() 2 1 ,
30
(Lys) 6
4, 5, 35
3, 34, 38
4, 5
- 10, 14, 35
- 37
Z- 37
26
32

32
2
1-5, 32-34, 36, 38
34
3234
35
- (,-) 40
31, 35
27
2
3
.

2, 3, 34, 36, 38
36, 38
29
(Met) 6
36
3, 29, 39
.
3, 36, 37, 39
19, 22
15, 16
37, 39
37
32
- 23
3, 29, 39
43
1, 3, 23, 25, 34
() 13
13
32
1, 3, 36, 37, 39, 41
17, 31
2
16, 23
23
23
55
() 36, 37

() 37
11, 36, 37, 39

12
3234
- 36, 38
5
- 33
36
36
36
36
36
7
39
3234
(,) .

1, 36, 37
36
36
42

- 23
. ,

26
22
S- 13, 15

()19, 30
18
1-3
21
15, 16
32

() 36
27, 28
lac- 27, 28
5, 43
27
.

6
-- 17
18, 22, 28
32
30, 35, 42
17
4
25
6, 8, 25
-- 35, 42
35
1, 6
--- 1 , 1 7
22
12

2
()36
19
- 20
38
17, 23
32
2, 27, 30
- pBR322 30
3
3
-- 18
10
34
41
4
26
.
18
6, 8 1 1 , 24, 25
1, 14, 31, 34, 35,40,42
() 2, 24
33
37
15
,

13

 36
21
9, 10
(
) 22
41
28, 30
1, 4, 5
1, 2
(Pro) 6
22, 28

4, 5
39
()
39
- 811
-- 4, 5
--- 10, 15
-- 18
--- 10, 40
--- 35
--- 10, 14
--- 34
--- 37
--- 36
--- 31
--- 2, 3, 24
-- 19
--- 7, 10, 18
41
36
43
13, 14
39
-22
29

22,28
39
() 19
(u) 17, 23

5
20, 21
4, 5, 20, 21, 29, 30
---
4, 5, 20, 21, 29
.
28
3, 39
EcoRl 18
() 18, 30
5
36
36, 41
35
P-D- 17
22
.
17
17
1-5, 23-25
() 2, 3, 19, 22,
24,27
42
, 19
() 22
(,
) 4, 5, 19, 22-25, 27, 28, 30
17
19
() 2, 3, 19, 22, 24,
27
19
4, 5, 22
() 19, 22-25, 27
5
- 19
- - .

- ()
21,30
- 4, 5, 21, 22, 28

1820
39
40
23, 27
36
.
13
28

41
28
10
1
-- 1015
--- 4, 5
--- 10, 15, 16
-- 18
--- 10, 40
-- 10, 14
-- 19
1, 4, 5,
10-16, 18, 19,40

37
37
37
.
4, 5
(S), .

21, 26
11
1
(G) 7, 12, 44
23
18, 40
- 19
(Ser) 6
13
16, 23
,
13, 15
, 13
4, 5

36
24, 25
- 4, 5, 20, 21,29, 30
4, 5, 22
37, 39
29
34, 36, 38
36
12
- (-) 8-11
10, 40
31
--- 11
() () 8, 9
21, 22
6
--- 17
7, 10, 12, 15, 40
R- 13, 15
- 13, 15
1820
22
27
- 22

38
12,13
22
22
36
8,9, 11
-- 18
-- 19
--- 31
- 811
10, 15
--- 34
--- 11
9
() () 8, 9
32
19
6
32, 33,41
42
- . -
- 18,40
1214
11
()
26

3234
33, 34, 36
16
35
34
11
29
34
7
23
22, 28

 25
33
33
35, 42
42
42
3
() 17
28
() 6
37
37
3
21
24
- 20, 21
() 42
- 8
--- 32-34
( , ) 22, 27
19, 22, 27, 28, 30
25
4, 5, 19, 23-25, 28
35, 42
27
38
- 38
38
- 38
- 36, 38
() 19, 22-25,
27
--- 24, 25
-- (
) 19
(Thr) 6
10, 34
- - 1 8
-- 19
() 32
() 6
(trp)28
11
11, 36
37
3, 39
- 33
(-) 40
6, 3234, 38
.
.
- .

1-4, 26
- 5
(U) 17
- . -

( )
24,25
( ) 24, 25
J- 40
.
38
S- 29
(Phe) 6
1214
24, 25
----- 20, 21
----- 22
- 28
4, 5
14
13
16
38
39
16
10
2
34
- 19
(fVal) 23, 25
(fMet) 23, 25
32, 34
34, 36
32, 34
32, 33
17,21-23
3234
32
32
32
33
2, 3
Fab- 40
Fc- 40
31

18, 19
31
() 42
1, 3, 25, 34
33, 34
32, 34
22
31
26
3, 26
1-3, 18, 26, 27, 29, 30
26
11
2, 3, 31
() 36
3, 29

1, 19, 30,44
3, 26
33
33 42
AMP (cAMP) 28, 41
-GMP(cGMP)41
- 8
-- 32, 34
(Cys) 6
3
6
27
() 17
1-5, 28, 32, 36, 38, 39, 41
39
3, 39
16, 34
10, 15
36
11
poly(A) 22, 28, 30
16, 23
30
3, 34
2, 38
.
2, 3, 38
(1)
31 22, 27, 40
21
2
() 36
1, 24, 34,
35,37
7
38
25
22 () 7, 44


7, 10
(S) 7, 44
7, 10
40
33
33
1, 4, 5
1, 3
36
41
1, 3, 29, 34, 41
3
1
1-3, 28, 29, 34, 36, 41
3


................................................... 5
....................................................................... 6
........................................................ 7

I.
1.
2.
3.
4.
5.

.
( . . ) ................................... 10
. ( . . ) .................................................................... 13
. ( . .
) ............................................................................. 16
- . ( . . ) .................................................................... 19
- . ( . . ) .................................................................... 22

22. . ( . .
) ...........................................................................70
23. . ( . . )..73
24. . (
. . ).....................................................76
25. . (
. . ).....................................................79
26. . (
. . ).....................................................82
27. . ( . . )..85
28. . ( . .
) .......................................................................88
29. . ( . . )..91
30. : . (
. . )..........................................94

IV.
II.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
16.

. ( . .
) ............................................................................. 25
. ( . . )..28
. ( . .
) ............................................................................. 31
. ( . .
) ............................................................................. 33
. ( . .
) .............................................................................. 36
. (
. . ) ............................................................ 39
. (
. . ) .............................................................. 42
. (
. . ) ...................................................... 45
: . (
. . ) ............................................................. 48
: . (
. . ) ...................................................... 51
. ( . . ).......... 54

III.
17. .
( . . ) .................................... 57
18. . ( . . ) ...60
19. . ( . . ) ...63
20. . ( . .
) ........................................................................... 66
21. . ( . .
) ..................................................................... 68

31. . ( . .
) .....................................................................97
32. : 1. ( . . ) ...............100
33. : 2. ( . . ) ...............103
34. . ( . .
) ...................................................................106
35. . ( . .
) ...................................................................109

V.
36. . ( . . ).112
37. . ( . . ) ..................................................................115
38. . (
A. . ).....................................................118
39. . (
B. . ) ....................................................121
40. . ( . .
) ..........................................................................124
41. .
( . . ) ...................................127
42. . ( . .
) ..................................................................130
43. . (
. . ) ....................................................132
44. .
( . . )..................................134
......................................................................136
..................................................137