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- .-. . .

- . . .

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

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



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.-. . 16,44. . 4/9797. 5000 . . 5914.

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



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143200, . , . , 93.

ISBN 5-03-003521-4 (.)


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


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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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NH2 ( N-),
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= -57, = -47.
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-, - . -, -,
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. 9.2. -.


= 60, = 140. (. 11).
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624 ,
0,9 2,4 .
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310, -. 310 3 ,
3
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(, Glu, Met, Ala Leu) -

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-, (Gly Pro)
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-
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>N- .
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Glu .
- -.

10.



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.

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Ala, Val, He, Leu, Met, Phe . ,

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.


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10 . 1050,
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1037 (~1030 ). ,
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11.

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() , .


.
GlyXY, X Y
. (Pro) X,
(Hyp) Y. :
GLy ProHypGlyProMetGlyProHypGlyLeuAla
, .
16 N- 25 -
. , , , .

,

0,29 ,
.
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, GlX2Y3G4,
N -. .
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0,15 ) -

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(G1 G4 , ). 1000 ,

300 .
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.

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,
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.

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- 0,35, 0,57 .


GlyAlaGlyAlaGlySer.
Ala Ser
-,
. -
, . ,
,

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


, .

, , , Arg,
Asp Glu. - , .
,
, , . ,
-, -. , - ,
30-
.
0,51 , -, 0,54 . , ,
,
(coiled-coil) , . 11
,
. - ,
- . -.
-.
, (. 36)
-,
-.

12.

,
(. . ).
1010
.
- ( S),
()
().

,
( )
- (ES)
.
.
,

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S- , , , , ,
. ,
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.


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(. ).

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90- XIX .
: ()
().

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,
.

.
S (
). G0 S

.
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() , , G0.
,

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, .
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G
G: , ,
.

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1.
.
2. ,
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(S) .
S-
.
3.
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.
4.
,

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,
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.

S-, -

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S .
,
- -

.
.

,
.

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).

-,
, . -


,

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

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.
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, .

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), - ,
(. . ).

13.3
, .
, -: v |S]
S- ,
, -
(. 13.1). . ,
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( ), , ( ).

,
,
. , . (R-, . relax
), (-, .
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'- , , -

().





, , ,

.

, -: .

, ,
. ,

. , , ,
.
() ,
,


. , ,
, ; -

. , . , .
,


,
.

,
.
, ,
. ,

, ,
: )
) .

31.

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


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), (1 ->4) (1 > 6)
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.

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2

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1935 . .

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35.



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4 .
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10 ), , . , .

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40.


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25 000) 220.
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. L-
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V- L- -.
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(LI, L2 L3 . 40.1) 6 . V - 3 (HI, H2 ).
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.
.
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VL CL. -
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3).
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.
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-
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.

lgG
.
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, (. . 10.1).
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VL, a CH1 CL. Fc- -
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, ,
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,
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Y- .

.
VH- VL-
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,
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,
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,
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.
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IgG, .
Clq ( 1)
. 18 . 3 . ,
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(. 11).
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IgG
, IgG
Clq (. 40.2).


(>10 8 ) .

.
,
( 10 8 ), .

-
. V- -
. , . V ,
( . joining ). VL- , 100
V- 5 J-,

V/J- 10 .
, 100 - 5 - 10 = 5- 103
VL-. V- , V, J D ( . diversity ). 100 V-,
5 J- 50 D-, V/J- J/D-co 10 .
, 100 5 50 10 10 =
= 2,5 106 -. 5 103 VL-
VHVL-nap,
,
5- 103 - 2,5 - 10 6 = 1010.
. 5 IgG, IgA,
IgM, IgD igE
. 5
, a, JJ., 5 8.
- L- - (IgM IgG 4, -).

41.

, .
: , , (. 6000), (. .
) , ,
(. 172). -,
.
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. ,
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.
,
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: ,
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.
(
) , . ,


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, , -

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,
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3',5'- ( AMP, ). ,
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2) , , , . ,
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, , , , ,
.

,
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(,
)
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. , , , . ,

. -

, ,
.
( ,
cGMP) (, 2+);
.
AMP
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2~4 -), R
. R ,
. ,

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( ).

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- ,
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.

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)
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-

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, .

42.

,

. XIX .
, , . , 1928 . 1935 . , , .

, .
, .

,

( ), ( ).
. 42.1 ; . , .
42.1




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

(. 24). ,
.
- , Penicillium notation.
: -. R . , L- D- (. 6)
,
(RCO). N
(- ) .
,
,
.

( G), R
.
1957 .,
,
, ,
. . . , 1965 .,
, -


.
(. . 42.1 42.2) D-ala ( ).
- ().
D- ,
(. . 42.2;
-
. 42.1). , D-ala, D-ala -,
-,
. , D--- (.
. 42.1). , -, :
, , - ,
- (. 42.3),
.

,

,
(). , . 30 000, , - .

43.

-
,
;

(,
Streptomyces). ,
, , ,
,
, .
- ,
, in vivo, -, , , . , , , ,
,
.

, , ,
, . ( , ,
).
A, ,
.
,
,
,
;
.
. ,
, D,

; , ,
.
- ,

(. . 43.1).
, ,
, , ,
.
, ,
,
-, -.
, -
, ,

. , , ,


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

-, ,

, ,
,
, -

. ,
. .
, , , . : , . ,
, , , .

- (- , , -
, . . ,

, .

44.


s ; ,
, . , ,
S:

: .
.
.

. 44.1, , , s ; , ,
s, .


, . 7.
1 . (
-
, )
, (, ),
, ,
, .
1- ,
. , ( U) ,

,

(, ,

). (), (1) :

a PV - , .
,
,
, .
2- . 1-
,
.
,
,

,
,
,
,
.

, ,
1- , , ,

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

,
.
, (S); S
.
S > 0. (
),

.
, ,
, . 1-
2- , , : - () ( ,
. . TS)) , .

. ,
,
(G),
, G < , (
); G = 0 ,
;
G > 0,
.
(Go - G
,
,
. . ,
;
1 -1. Go
.
' = []/[] ( [] [] -
),
, G
.
, G (5) (4).

. Go
,
; ,
+,
1 -1. , [+] = 1 = 10, .
. = 0.
, = 7 ,
G, G', ,
+, -
,
10-7 -1, =
7. ,


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

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. ., . .:
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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
---- 39
-2 4, 18, 27

4 14, 26, 27
7, 17
-5 27
2, 3
-6 27
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

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