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2Molecular Biology of the Cell SECOND EDITION Bruce Alberts, Dennis Bray, Julian Lewis, Martin Raff, Keith Roberts, James D. Watson

Garland Publishing, Inc. New York London

3. . . . . c . 2- , 3- 2 . . . . , . . . . , . . . . , . , . . . . - . ..

1994

4 28.070 75 576.32/36 : ., ., ., ., . . . : 3- . 2- . . 75 . . 2.: . .- .: , 1993.-539 ., . ISBN 5-03-001987-1 ( - ) . . 3- . 1- (.: , 1986-1987). . . 2 , , . , , , . 1903010000-024 7-93-235 041 (01)-94 28.070

ISBN 5-03-001987-1 (.) ISBN 5-03-001985-5 ISBN 0-8240-3695-6 (.) 1989 by Bruce Alberts, Dennis Bray, Julian Lewis, Martin Raff, Keith Roberts and James D. Watson , .., . ., . ., . . 1993

5

8 , , . . , . , , (). , , ; , . , , , . . , , . , . 10 (1010) 10000 , - , . , . , . , . , () , , .

8.1. . , , .

8.1.1. [1] . ,

6

. 8-1. . , , , , , , , . . , . , , , , . , , . 8-1. . . . - . . , , , . (). . , . . , ; , . , , . , , , . , . , (

7 8-1. () 1) 1) 54 22 9 6 6 1 1 1 1 1700 1

1 400 300 200

1) , . , , .

) , . , , , , . , , , 8-2.

1) 2

1) 5

35 60 16 10-5 ). , , (. 7.1.8).

363

12.3.8 2 + [15] 2+ 1947 ., , 2+ . , 2+ , . 2+ (. 12-20), , - . , 2+ , ; 2+ - , (. 19.3.2). , 2+ ; -. ().12-14

12.3.9 (InsP3) 2 [16] () 1953 ., , -

. 12-20. , Ca2+ , . - , . 2+ .

364

. 12-21. () 10% . (PIP 12)- (). , 12, 10% 1% . () - . , , , . PI-1- (PIP) 1-6 (12), , , (. 12-21). 12 , PI, 12. , 12, . , 25 (. 12-4). , , , , , . , G- ( Gp), . 2 : - 12-4. , .

-

PDGF

365

. 12-22. 12 (InsP3), 2 + - . - - , (. ).2 +

(. 12-22). . , . (InsP3)- , 2+ (. 12-22). lnsP3 ( ) , 2 + . , InsP3 . : 1) 2 + , , , (, , 2+ , ); 2) InsP3 ( ) . InsP3: , , -1,3,4,5 (InsP4), , , . 2+ , 10 .

12.3.10. , 12, [17] InsP3, 12, 2+ , 12 - . : , (. . 12-8), , , -

366

, -. , , -, 2 + . - - - , 2 + , 2 + . , , - , , 2 + , InsP3. - , . -, 2 + , -, . , -, -, +/+- , (. 6.4.10); . - , , (. 19.5). - . , , - (. . 10-1). , , , - - ( ) , . , y . InsP3 , 23187 , 2+ (. 6.4.19).

. 12-23. - . , G- (Gp), - , 12 InsP3 . . ( 2+ ) -. , -- , ; . InsP3 2+-, , - .

367

- , - (. 12-23). , . , , -, . - , , , . . - , , , , . , -, . , , .

12.3.11. ras- G-, -, . , - , , ( - ). . . , , - -, (. 13.4.2). , , ras- ( , -rat sarcoma). G-, GTP. ras-, ras-, ras- ( ) . , , GTP- , G- (. 12.3.4). ras-, . , ras- - -. , , - . ras--

368

, , Gp-, , ras-, : Gp G-- , (. . 12-3). , ras-, , G . , G-, .

12.3.12. G- , [19] , G-, . , G- (Gs Gi) , (Gp) . G-, , , cGMP (. 12.4.5); -. G- , , G- . , Gi- , . ( , , - , -. . 6.4.17.) G-, , . : - Gi, , 65% . , , , G- , . , , . - (. 12-24), , . , . , , , ( , , , G- - . . 6.27) , (. 10.3.2). , - - .

. 12-24. - . - (. 15.5.2). , , G-, . ( R. A. Dixon et al., Nature 321: 75-79, 1986.)

369

12.3.13. - , [20] G-, , . -, - - , . -. , , (, PDGF - platelet-derived growth factor, . . 13.3.4) (, EGP - epidermal growth factor), (. . 12-25). , , , 0,1% . - , . . ? , -. EGF , . , . , . , , , , . , , , . PDGF , -, , (PI-, . . 12-21). , , PDGF , Gp1. , - (. 6.5.7). , -, . , _____________ 1 , , 12.-. .

370

- (. 12.5). , , .

12.3.14. - [21] 1979 . , - , 60 v-src (. 13.4.2). , ( 1982 .), EGF. , erbB EGF. EGF- , , , . , , , , , ( ) . . , . . . sis, , PDGF, erbA - . . 13, , . - , - , . , , , , G-, . 70 , , , , (. 12-25). , , - , - , . , - 2+- , , .

371

. 12-25. , . ( ) 250 ; . - -, ( ) . , - - - , . PDGF, -, - , (. 18.6.20). - (. . 12-27) (. . 12-31), , .

, . , , - , . - ,

372

- . , G-, , , GTP- (G-). G- , GTP. , . - , - (12), : 1) (InsP3), 2+ ; 2) , , . , , G-, . G- , InsP3 , , , 2 + .

12.4. AMP [22] 2+-, , , . , .12-21

12.4.1. AMP - [23] , - (-). -. , -, , N- . . - . - ; , ( , ). , , ( . .), , . , , -1- . -1- -6-, , . .

373

. 12-26. , ( ) . - , , ( ) , . - ( ), ( ). , - , . -, - ( , 1959 .). ( ) , (. 12-26). -, , - , , . , , , . . (. 12.1.2), , , . ( 30 ), , . , , -, . - - , , . , . -; - (. 12-27). - , , , , .

374

. 12-27. - (-). , . . , , . (. . 12.4.8). - - , . - , , , , -.

12.4.2. [24] , , , . , AMP. . AMP- , , , - , . . - -

. 12-28. AMP , , . -, -, .

375

. 12-29. - ( ). , -. 2+- , 12 , 2 + . - , N- . ( Y. S. Babu et al., Nature 315: 37-40, 1985.) - . (. 12-28). , -, , , , - .12-22

12.4.3. [25] 2 + 10-7 , , 5 10- , , , 2+ () 106 /. , Mg2+ 10-3 , 1000- 2+ Mg2+. 2+- . ; . 11 (. 11.1.12). , - - - . 107 , 1% . 2 + , , . 150 , 2 + ; (. 12-29). - AMP, 2+- , . ( , . ),

. 12-30. , 2+ , . -, 2+ - : 2 + (. . 12.4.8).

376

2 + - , , (. 1230). , 2 +-, . 2+/- (-), . - - - . -, 2+/- ( - II); 2+ . , - 2+ , 2+/- .

12.4.4. 2+ [26] 2+ . -, 2+ . , 2+/ , ,- - - ; - . -, - -. -, - - , , , AMP, 2 + . 2 + , . , (. . 12-26). , : AMP, . , , , , . - ; 2+ --; - - ; - (. 12-31). 2 +-, , . 2 + , (. 11.1.11), , -, . , 2 ' -, , . , , ; , , - 2 + - -

. 12-31. . -; - - AMP, ()- 2 + . .

377

, . -, . , . , , 100 , . , , - , - , -, -. , , - , . , , , .12-16

12.4.5. cGMP [27] - , . (cGMP, . 12-32), 10 . , cGMP (G-), (. . 12-25), GMP , , . , cGMP GTP, , cGMP . , , , . cGMP , cGMP Na+ . cGMP . ; G- , . - (T) cGMP-, cGMP, , cGMP, , , . , ( . . 19.6.7). , G, , . ; - . ( G5-) ; ,

. 12-32. GMP.

378

, . , , , G-, . , , . , ?

12.4.6. [28] , , . . 12-33 , , , , , S-, . . . (10-9 ) , 2+ , (10~6 ) . , , -. , , , , (. 12.4.1). . , 2 + , , . , ; , .12-25

12.4.7. - , [29] - (, AMP 2+), - - , . , , X Y, . X - 10 X; , 100 . Y 10 , .. 100 , Y 10 .

379

. 12-33. G- , . . GS- (. . 12.3.5).

10 , , 1 Y 900 (10-100 100), - 90. , - , - , , (. 12-34). , , , . , , 2 , 10 . , .

380

. 12-34. () () 10 ( ). , ( ), , - . - .

12.4.8. [30] () . , , , (. 12.2.1). - , , , . . , - . , , . , , , . ? , - - ( ). , , -, - . ,

381

. 12-35. . . , - . , . , , , , ( ). ( . R. Mulvihill, R. D. Palmiter, J. Biol. Chem. 252: 2060-2068, 1977, .) (. 12-35). - ; , 2+ (. . 12-30), 10 50- . , - -, , (. 12-36 12-37). , , . , , , (. 12.4.1, 12.4.2). - . . , - , Na + . , , ; , . . , .

. 12-36. , , - -. (1, 2, 8 16) .

382

. 12-37. , . . . , , . , (. 12-38). , , , . . . , . 12-38, , , , , , , , : , . 2+/- (. 12.4.3), , , 2+ . 2+/ . , . , .

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

. 12-38. . , , .

383

; 2+/ -, 2+ /- (a2 + -). -, 2+ , . AMP 2 + .

12.5. - , . , -, - , . , (), . ( ). . ; , .12-28

12.5.1. - [31] , -, , (. 6.5.7). () , , . , -. , . , : , , EGF, EGF 10 , EGF 1 . ; . , , : (. 6.5.10).

384

. .12-29 12-30

12.5.2. [32] . , (.. ) (. 12.3.14). - . , ; , - . (. . 12-24), , . , -, , S-. , - (. 12.3.12), : , , G- . , , , (. 19.6.8).

12.5.3. , G- [33] , , , G-. , PGE1, GS-, PGE1, , GS- - ( , , - , ). GS- , 8-, ( GS-) . GS- PGE1 , , . G- . - (. 12-39),

. 12-39. , . , ? , - , . 1975 . , . , . 4 , ( ). , , .

385

. 12-40. - . , ( ) G-. , G-, , ( , . 12.5.4). , .

. 12-41. Salmonella typhimurium, () (). 5 . - . (. A. Rubik, D.E. Koshland, Proc. Natl. Acad. Sci. USA 75: 2820-2824, 1978.) , . , , ( ). - . Gi (. 12.3.6), . , , , , - . , . , , (, , ..). , , Gi-. -, , . 12-40.

12.5.4. [34] , . , , . , . . 14.3.2 Dictyostelium discoideum . 11.6.4 . , - . (), , () (. 12-41). , -

386

. 12-42. , . coli. , . ( 100 /) (). . . , ( . L. de Pamphilis, J. Adler, J. Bacteriol. 105: 384-395; 396-407, 1971, .) - - E.coli Salmonella typhimurium. ; , . , , (. 11.3.2). - . , . , , (. 12-42). , . , . (. 12-43). , : (. 12-44, \ . , , . , , , ( ). , (. 12-44, ). , , .

. 12-43. . coli . (), , . () , .

387

. 12-44. . () , . () , . . , (- ). - . , , , ( ) , , , . , : , - , . , , , .12-31

12.5.5. [35] . , -, . (. ), - (. 12-45).

. 12-45. . , , 13 . , . ( A. F. Russo, D. . Koshland, Jr., Science 220: 1016-1020, 1983. Copyright 1983 by the A A AS.)

388

. 12-46. . 1 2 - , 3 4. , , ; . ( ). . , . , , (. 6.4.13). , ( ), , (. 12-46). (- ), . , , , .

12.5.6. [35] , . , , , . , ( ) : 1) , , , ; 2) , , - , (. 12-47). (), -

389

. 12-47. ( ) . , ( , ) . , . , , . , , . , . . , - , . (. 12-48). ; , . (. 12-48). , , .

12.5.7. [36] , . , - CheA, CheW, CheY CheZ. CheY CheZ , , . CheY , . , CheY , . CheZ , CheY, , . CheA , , CheW CheY CheZ , .

. 12-48. . .

390

. 12-49. , , , . , CheA. CheA CheY. CheY- , . , CheA CheY, . CheZ CheY-, CheY. 10 , (. . 12-34). CheA CheW, . in vitro , CheA- , , CheY. - CheY CheZ, , , CheY in vivo. -, CheY ( ), CheA, ( ), CheZ. , CheA CheY , CheY ; , , CheA- CheY, CheY (. 12-49). . CheA , (. . 1248), . -, , , , . , .

, , , , , . : 1) , ; 2) ; 3) ( G-) . , , - : .

391

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394

13 . , , . , . , (, , ), . . , , . : , . . ( ) , . , , - . . , , , , . ; . , , . , (. 13.5.2) ; .

13.1. . ,

395

, ; , . , , . , ( ). . , . , - , . , .13-3 13-4 13.1.1. , [1] - , S- . - , G1 ( , gap-); , G2, -. , G1, S G2 90% . ,

. 13-1. . , () (), . G1, , . S- ; , ( ). 2, , .. . . ( , , - ); , , . , . 24- , : 8 , - , G1.

396

. 13-2. , 3-. . 4.5.2. ( ) , 3- , , - S. ( James Cleaver.) - 16-24 , 1 2 . . 13-1; . 1950- . . 3- - , , 3 , in vitro (. 13-2). 3- . , (, , S-), . , -, 3--, , , , . , , , . , , , 30% . , 3-, , S- (. 13-3). 5% ( , ), . , S- . , , 3-, , , ; , . G2. . . 13-4. , - (, ) (, ). , G1, ( ) , ( ). , G1 G0 (. . 13.3.8).13-5

13.1.2. in vitro [1, 2], , . . , (. 4.1.5) , , , ; -

397

. 13-3. , , ( ). , 0,7 G1 1,4 . , ( ) , .

. 13-4. , . 3-, ; . . , , , . S ( ), G2, G1 . , G2, ; , G2 + , , , .. , G2, S , ( ) G,. 3, 7, 22 11 .

398

. 13-5. , - , , - (. . 4.3.1). , . . : (1 ), .. G1, (2 ), . . G2 , , S. , G1 , G2 , . , G1 , G2 + M. - . , , , , - 3-. ; (. 13-5). , . . , ; , , . , . , .

. 13-6. . , , . . - .

399

. 13-7. . , , ( ). , . , -, . - , (. 13-6). , , , G1 . : , , , .

13.1.3. [3] , . (. 13-7). , -, , , , , . (. 13-8) , . , - . , . , , , . , , G1, , . , , , . , ( ), (. . 13-8). , , , S-; , , . ? , , , . . S- .

13.1.4. - S- [4] (. 4.3.5) , . . S- , G1 , G1 (. 13-9, ). ,

400

. 13-8. , G1 S, . G1 S- . 1000 , ( ) . , . (P. Coffino, V. . Groppi, Adv. Cyclic Nucleotide Res. 14: 399-410, 1981.) , G1 - ( ), . -, , S-, . S-, , G1 S . S- G2? - . G2 Gl (. 13-9,), G1 ; G2 S-, (. 13-9, ). , S- ( - ) G2 , .

13.1.5. [4, 5] . 9 (. 9.3.6), S- , -

. 13-9. , .

401

, . G2 S- (. 13-9, ) G2 S- . , G1. - , , S- , . , , , S- ( S-), . S- , S-, . -, , , - . , . . 9 (. 9.3.1), : , , , . , , , . , - S-, . , S-, . , . , G1S , (. 13.3.3).13-6

13.1.6. - , [4, 6], S- G2, . , , . S G2 , , . , - , . S-; , G2 (, ) . , ;

402

, , ( SOS-. . 5.2.9). , -. , : ( ), , . , .13-7

13.1.7. - (MPF) [4,7] , -, - . G2 , . . - (1, S G2), -, , ( G1 S) (. 13-9, 13-10). -, - - (M-phase-promoting factor, MPF), . , , , MPF, , .

. 13-10. PtK . . PtK 1, . . PtK S, . . PtK G2, , , . (. Sperling, P. Rao, Humangenetik 23: 235-258, 1974.)

403

13.1.8. - [8] , , . , ; , , . 1) S-, S- ; 2) - (MPF), - ; 3) - - (M-phase-delaying factor, , S-), S- , MPF. , . ( , , ) , , , . . , ; , - ; - S- ; ; G1. : G1 , (. 13.5.7). , , (. 18.2.1, 19.8.2), , , .

13.1.9. , , [9] , , Xenopus. Xenopus, , . , ( ), . , . G2 ( , G2-. . 15.2.7). , - (. 15.3.3). :

404

. 13-11. 7 Xenopus 12 , S G1 G2. 4096 (212) . . , , (. 13-11). , , . , 90 , 11 30- , 7 4096 (212) . , , . , S , a Gl G2 , .

13.1.10. - (MPF) [10] , . , , . - (MPF, ), . Xenopus, -. , G2 -. ( MPF mutaration promoting factor, ; . . 15.3.6). MPF () (. 13-12). , MPF, (. 13-13).

. 13-12. - (MPF, ) , Xenopus. G2 MPF; MPF; S MPF.

405

. 13-13. MPF Xenopus. MPF . ( ) . MPF : , , , , , Xenopus -. Xenopus MPF. , ; - , , -, . MPF, , : MPF Xenopus, MPF (. 15.3.6). , MPF - , . MPF , (. ); -, . -, , , ; MPF- , , H1, , , (. 9.2.2); , , , MPF .

13.1.11. MPF [8, 10, 11] MPF, 30 , -

406

. 13-14. , pus. ; , , , . , . . (. , P. Tydeman, M. Kirschner, . Natl. Acad. Sci USA 77: 462-466, 1980.) , . , , , - (. 13-14). . , , . , . , , ( , ) MPF. , us - MPF, , , F -. , ; , , MPF. us S- , -, . , (, , , ), : , -, MPF . , Xenopus . , G1, S-, -, . , Xenopus . , , , , . - , pus, . , MPF, , .

407

. 13-15. MPF , . , 5% , . , -. , . , . Xenopus, , MPF, G2 . , MPF - , - -; MPF (. 13-15). , , ; , .

, Gl ( ), S ( ), G2 ( ) (). S- - [ S- - (MPF ) ]. S S- , - , . - - - , , , . , , Xenopus, . , -, MPF .

13.2. [12] . , , 1/1000 , . Xenopus- , . , -

408

. 13-16. . () G1, S, G2 . , , : . ( ). G1 S, , S, G2 . . ( ), . , , . : Saccharomyces cerevisiae, , Schizosaccharomyces pombe. , : , , (. 13-16). , , , . . , . , (. . 15);

. 13-17. (Saccharomyces cerevisiae) (Schizosaccharomyces pombe). , , . . , , . , () G1, . , ; , . - , , , .

409

(. 13-17; . . 10.3.2). ; ( ) . ; , .13-10

13.2.1. , , [12, 13] , , .

. 13-18. , (cdc) . , , , , ( S). , , , cdc ( ). , , (, ), , .

410

. , . , , () , () . , -. , , , , . , , , . (cdccell-division-cycle) , (. 13-18). , , cdc; , . 40 50 cdc. . , cdc, S-, , - , . , , cdc, . .13-11

13.2.2. cdc , cdc , cdc. , , , . , , , , -, . , cdc. , , (. 1319). , , , cdc, - , . -, , , , , .

411

. 13-19. cdc . () . ( G1). A.. ; , ( ) , , ( ). cdc, . , cdc8 . , ( ), ( ) ( ) . . cdc, , . [ L. . Hartwell, J. Cell Biol. 77: 627-637, 1978; J. R. Prmgle, L.H. Hartwell. In: The Molecular Biology of the Yeast Saccharomyces (J. N. Stern et al., eds.), pp. 97-142. Cold Spring Harbor Laboratory, 1981.] , (. 16.5.2).

13.2.3. , , , . ; , , , (. 13-20). ? , - , . , , , . G:, , , G2. G1 , S- , G1 , S-, . (Start). ( ) .

13.2.4. [14] , , G1 , , , ; G1 . . -

412

. 13-20. , , . . , , , . . : , , . ( .) , , (. 13-20, ). G1 , ; , , G1 , , . , , , , - . , Amoeba proteus , , , , . , , , : , (. . 13-40 13-41). , - ( ) ( , , ). , (, - ) - ; , . ; . , - . cdc , , , . , , , . - (cdc28) (cdc2) . , : , - .

413

, G1 , , , . . , , .

13.2.5. , - (MPF) [15] - , , . ( ) cdc. . 13-21, cdc . cdc28 cdc2 ; . cdc2 cdc28 , : cdc2 cdc28 . , cdc2 , , , , , cdc2/28. -, , , , , , . cdc2/28, , . , , , MPF . , , cdc -cdc 13, cdc2, (. 13.1.11). , MPF , , ; cdc2/28 - - - , MPF - G1.

. 13-21. cdc . , cdc, , . cdc2 , cdc28 . , , cdc .

414

, , , .

- , . , , (cdc), . , , , ( ) , . cdc, , . ( cdc2 cdc28-y ) , MPF; (cdc!3) .

13.3. , , , , . . , - , . , , . , , ; . , , , . . . 17 21 , , . 16 , .

13.3.1. [16] , 1013, . ; , , , ,

415

. 13-22. . , . , . . 20 ( ) . : , , . ( . S. Pptten, R. Schofield, L G. Lajtha, Biochim. Biophys. Acta 560: 281-299, 1979.) , (. 13-22). - : , . S-; . , , S- , ( 12 24 ) , . ( G0) , . , , ; ; , . , , (. 13-23). . ,

416

. 13-23. . 3- S ( ); . S , - ; - . , G0, . 40- , , S , S- . , ; 36- , . ( . Harding, J. R. Reddan, N.J. Unakar, M. Bagchi, Int. Rev. Cytol. 31: 215-300, 1971.) , . , .

13.3.2. , , , G1 - [17] in vitro (. 4.3.4), . , ; - - . (, ) , , - , . - , , , : Gt , . G1, S, G2 . G1 (R), , . ; , , . , , G1 , .

417

. 13-24. , in vitro. , .

13.3.3. , -, [18] , , . , (. 13-24). , , G1. -, GJ (. 13-25), , , R . , ; R , , - , , . . , , , S- (. 13.1.5). , . , : , , , .13-15 13-16

13.3.4. [19, 20], , , . , , ; . , , ( 10-9 - 10-11 ). . , . , , , PDGF (plateled-derived growth factor). , , -

. 13-25. , - G1 , , , (R). , . , , - - R. , - (. ).

418

. 13-26. (PDGF) . PDGF , , , . , . - , PDGF . , , , . (. 17.5) , ( , ) PDGF. . -, PDGF , (. 13-26). , PDGF, ( ). , (. 13-1); . , . , , , , - . ( 30), - , . , , , , , , ; , , . , . , , , (. 16.2.3). , , , , .

419

13.1.

(PDGF)

, 125 - 160 53

(. 13.3.4) (. 16.3.7) (. 12.3.13) PDGF ,

(, EGF)

I II 70 73 (IGF-I IGF-II) (TGF-P)

112 ( ) , - (. 16.2.3 17.7.1) 140 , , - 146 , (. 17.3.7) (. 17.6.1), Xenopus (. 16.2.3)

(, FGF)

-2 (IL-2) (, NEG)

153

- (. 18.6.11)

1 1 8 (. 19.7.10)

. . 17-2 . 17.5.8 (IL-3; GM-CSF, M-CSF; GCSF, )

13.3.5. [20, 21] , PDGF, . , , . . , , (. . 13-23) , ; . . , , , , , . -, , . ,

420

. 13-27. , , , . . , . , . , (. 13-27). , , , . , , . , , , , , , , , , . , , , - . , . , PDGF 10 -10 ( 3 ). 105 PDGF, . , , PDGF 150 . , , PDGF, , (. 6.5.12). , . , , - .13-14

13.3.6. [22] - , . . , , ( ). , , , . . , . (, ), , , , , (. 13-28). ( ) (. 11.2.8). , - -

421

. 13-28. . , () ; . 3- 1-2 , S. . , - , , . . , , , , , . ( . O'Neill, P. Jordan, G. Ireland, Cell 44: 489-496, 1986. Copyright Cell Press.) . , , , , , .

13.3.7. . , , ; , , , - , -. , , , , , . , , . , PDGF (. 13.4.6). , , . - , ; , (. 13.4.7).

422

13.3.8. , , - G0 [17, 23] , G1 . , . , , , . S , G1 . , . , , . . 3,5 . ( ) G1 , 8- ( ) . , G1 . . , ; , , , S, G2 , . . , , . , : 1) 2) , , .. . , , ; , , G0. , . G0 . , . 17.

13.3.9. [24] G0, , ; G1 , G0 . G0

423

. 13-29. ; . , G0. ( 10s). [. . Boycott. In: Essay on the Nervous System (B. Bellairs and E. G. Gray eds.). Oxford, U. K. Clarendon Press, 1974] , GJ, . G0 , : , , . . , , . , - , G0, , (. 13-29).13-17

13.3.10. G0 : [25] . , , , , - , . , , , 50 ; , , , . , 40- , 40 , 80 - 30 . . . . , , {. 16.3.4), , , , , 50 . , - ; , , . , , . , , ( ) . , - , . ,

424

. 13-30. . , , , . , , , . , , (); 8 , 8 (). , . , -. , . ( J. R. Smith, R.G. Whitney, Science 207: 8284, 1980.) . , . , , . , 50 , , 150 , . . , , ; . , - , , . , , . , , 50 , . , PDGF , . , . , , , - . , -, , , , (. 13-30). , , , . , - -

425

, . , , , .

, . , , . , . , G0 . G0 G1, S, G2 . , , , . , .

13.4. [26] , , . , ( ), . , , , , . - , . . , , , . , , , , . . 21. , , , .

426

13.4.1. , ( ), , ? , . , , . 1970- ., , . . , . , . () , (} . , , . . . , , . , , , , . 1-2. , _______________________________________________________________________________________________________________ 1. , . . . 2. . ;

. . . , 3. . . . ( , ) . ( )

. __________________________________________________________________________________________

427

. 13-31. , . (. 13-2): (. 13.3.5) (. 13-31); , ; , , ; ; ; , , .13-20

13.4.2. [28] , ; , . , . - , . , -. . 5-75. , , ( , . , ' , , . , ( ), . 60000. : 34, 39 ( ) (. 13-32). , ( ), . ,

428

. 13-32. , , , ( v-src) (, ). . (34 ), . . , , ( 39 ). ( G. Steven Martin.) , v-src. ( . onkos-, ), . ? - src -- (. 4.6.7), , , . src c-src . -, - - . , - , , , . , .13-21

13.4.3. , , [26, 29] , , . in vitro. , , . , , , (. 13-33). , , , , , . , . , -

429

. 13-33. , . , , , . , . Alu (. 10.5.11) , . - Alu , . , , . , , , . . 21. 50 (. 21.2.3, 21.2.4). , , . , , . , , , - . , , . . , . , , , . , , , , . , . , ; -, , . , , , .13-16 13-22

13.4.4. [26, 30] , () , . ,

430

. 13-34. . . . . 21-27. , c-sis, PDGF. , v-sis, , , PDGF, . - c-erbB, c-fms -rb - : -erbB (. 12.3.13), c-fins- , (M-CSF) ( , . 17.5.8), a c-erbA- (. 12.2). , , , ( ), , (. 12.3.14). , , , . , , . 13-34.

13.4.5. , . 12, , PDGF, - - , . , -

431

. 13-35. , , v-src , ; , src, c-src, , , , . src , . { Ira Pastan; M. . Willingham, G. Jay, I. Pastan, Cell 18: 125-134, 1979. Copyright Cell Press.) EGF M-CSF (. 13-34). , c-ras, , GTP , , G-, (. 12.3.11). ras, , , ; , , . , ras, , . , , 2+ AMP, , , , . , . , , . , , - -, . , cdc/28 , - /- , . , (, . 13-34); , c-jun, -1 (. 10.2.8, . ); - c-fos, , . , -, , , : - , G0, . , - , , . .

13.4.6. [32, 33] c-src, v-src . src - - . 60000 ( P60src), , (. 8.2.3). , , , (. 13-35). ,

432

. 13-36. src , , . . . - , src src- . . , ( ). src ( ). src , . -, src, , , , . (L. R. Rohrschneider, Proc. Natl. Acad. Sci. USA 77: 3514-3518, 1980.) src , . . , , (. 13-36). -, src , v-src ( ) ( ; . . 11.2.11), , , (. . 13-30). src. , v-src e (. 11.2.8). in vitro , ( ), ( ). , , v-src, , . , , - ; . , , . , . , v-srct , : ( , - . . 14.1.3) , ( ). ? PDGF, : , . , PDGF , v-src. , , PDGF, c-src, , src,

433

. 13-37. , , PDGF. PDGF ( ) c-src. , , , . , . , , src (v-src). -, src - c-abl cyes,- , c-src . , , . ( ), . , , . (. 13-37). v-src ( ) - , . , PDGF, , , v-src, G0 .

13.4.7. [33] . , : G0 , ( ). , , ( ),

434

, , . , - . , , -, , , (. 13-37). : (, ), , . , , , , , . . , , . 13-34, , . , (. 13.3.6), , , . , , ? , . , , v-src, , . , , : . , , , .

. 13-38. , . ( 1, 2 3) , G0 . , ( ), ( ). , , ( ) : , .

435

. 13-39. , . : 1) , , 2) , , . , , , ; , . , , . , , . (), ; , . () , , . () , , . , , (. 13-38) . , , - , , , . , , : 1) , (. 11.2.8); 2) , , ; 3)

436

, , . , . . 13-39.

13.4.8. [20, 34], , , , . , , , . , , , (. 13.3.10). , -, , . . , , Caenorhabditis elegans, , 959 ; , (. 16.3.3). , . , , . , , , ,

. 13-40. . , , , . , - , , . [G. Frankhauser, In: Analysis of Development (. H. Willier, P. A. Weiss, V. Hamburger, eds.), pp. 126-150. Philadelphia, Saunders, 1955.]

437

. 13-41. (. . 13-40). . . . ; , . (G. Frankhauser, Int. Rev. Cytol. 1: 165-193, 1952.) (. 13-40 13-41). , , , , . , . . , . , , , , , , , (. 16.4.9). . , , . : , , 12 , . , , , ( ) . , , (. 21.1.4). , , , , . , , , , , , , (. 5.6.10). , , .

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

438

. , , , , . , , , . 13-29

13.5. [35] - . , , , (. 13-42). -, , (MPF), , (. 13.2.5). -, , , , , , . . 1, ( HI). HI , (. 13.2.5), MPF (. 9.1.12) . , , , . - , : , . , -. , . ; , . , - , - , . , , , , . . ,

. 13-42. - G2 G1 . () ().

439

() ()', , , , . . , ; , , (. . 13.5.18). , , , . - . , , 105 .

13.5.1. - [35] . , . , . : ( ) . 13-1. - , , , - ; (. 13-43). . 1344 13-45 . , 131. , .

13.5.2. - [36] . 11 , - , (. 11.4.4). , , (-) . , , , , . -; , -, -

440

. 13-43. , . . , . (, , ) , ; , G2.

. 13-44. ( ) . ; . 100 . , , . ( ); , . , . . . , . , ; . ( . de Brabander.)

441

. 13-45. . Haemanthus (), (. 4.1.5). . . : (N). . : , (). , ( ) . . : , . . : , . . : , , . 3. : ; , (3). ( Andrew Bajer.)

. 13-46. . . ( ) ( ) ( ), . Gt . S . G2. , . , - . , , . , , , . .

442

G2 , , . , , . . S . , , . , , . , . .

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

13-1. .

443

, , . 1 /. . , . . .

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

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

444

. 13-47. ) . ( ), -. - , (. 11.4.3). , , , ( ), - . , , - , (. 11-2). I , 13-46, 1 . - S , . - , . , . , , , . . , . , , , MPF , (. 13.2.5). , , 20 ( 5 15 , . . 13-48). , , , - (. 11.4.3, 11-2), , ( in vitro). , , , ( ), , (. . 13-46). , , , , . , , , . , , - (. 13-47). , 40. , . , . 13-48. , , , (. 13-49).

445

. 13-48. , , - , . . , , . , , . 50%- , ti,2, , , , . ( W. . Saxon et al., J. Cell Biol. 99: 2175-2187, 1984, by copyright permission of the Rockefeller Univ. Press.) (D2O) ( ), . , . , , , , . , , , , . , .13-25

13.5.3. [37] , , , , - , .

. 13-49. . : (), () (). ( . D. Salmon, R.R. Segall, J. Cell Biol. 86: 355-365, 1980. Repr. by copyright permission of the Rockefeller Univ. Press.)

446

. 13-50. , . , . . ( ), . (. 13-50). , , (. 13-51). , , : , 20 40, - , .. , . , , . - , ; . , Saccharomyces ccrevisiae 17 110 (. 13-52); , . , , , (), . , , , . , , ( , ), , . , , . ; (. 13-53), , - . ,

. 13-51. . (), , , , (S). , . , , ( ), . ( Bill Brinkley; - J.D. Pickett-Heaps, L. . Fowke, Aust. J. Biol. Sci. 23: 71-92, 1970. Repr. by permission of CSIRO.)

447

. 13-52. Saccharomyces cerevisiae. , ; , .

. 13-53. , . . . , G1, . . G2 . (S. L. Brenner, . . Brinkley, Cold Spring Harbor Symp. Quant. Biol. 46: 241-254, 1982.) , , , , , . ? . , (. 13-54). , : , . , , , - . , , -, , (. . 13-61).13-28

13.5.4. -, - , [38] , , . ,

. 13-54. , , , (). in vitro , (). 1 , , . , , ( ), , , ( ). ( Luise Evans.)

448

. , 1) , , 2) , . 10-20 . , . , , , , . , , ; , . , (. 15.2.7). - , . , (cap), - - , ' -. , , , . ( } . (. ), - , , , . , , - . -, , , , .

13.5.5. [39] . , , . -, , , . , . , , , -

449

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

13.5.6. , , [40] , ? , , , , . . , (. 13-55). , - . , , . 1356. , . , . , , ,

. 13-55. , , , , . , , .

450

. 13-56. . ( ), , . . , , ( ), . , . , . , , , . , , , . (. . 13-43), , . . , , , ; , , (. 9.2.3). , , . , , , . , , , , .

451

. 13-57. Haemanthus. () () . , . ( Andrew Bajer.)

13.5.7. [41] , - , , . , (. 13-57). , , , , . , 2+ . -, , (. 4.2.3), , , 2+ . -, . -, , - , . , , , , 2+ (. 13-58), 2+ (. 11.1.14).

. 13-58. ( ) ; . ( ; Peter Hepler, J. Cell Biol. 86: 490-499, 1980, by copyright permission of the Rockefeller Univ. Press.)

452

13.5.8. [42] , , , -, - , (. 13-59). , ; . - , . . , ( ), , ( ). . , , 1,5-2 . , , , , 15 , .

13.5.9. , [43] . 10 -5

. 13-59. , . . (, ). . (, ). , , , , (. . 13-46). , : 1} , 2) , , .

453

. 13-60. . . - , - -. , , . . , - (); . (). , 10000 , , . , - , , - . , . , , , , . , , , . , , , . , . , , . , , , (. 13-60), , D2O. , , , . . 13-61. , , - .

454

. 13-61. , : . . , ; , (. 10.4.9). . : , , , . , , , , (. . 13-60). . . , . 13-61, . , , , - . , , (, - . . 11.1.13), . , , ( , - , . . 13-60). , , , ; .

13.5.10. , , [44] , . , - -. , . (. 13-62), (. 13.5.18), , , . , , , .

455

. -62. , . . . . . ( Jeremy D. Pickett-Heaps.) . , , . , ( , ) , , . , , (. 11.3.7), , (. 10.4.9). . , . , (, , . . 13-56) , . , , . , - , . (. 13.5.13).

456

13.5.11. [45] , . -- , . : 1) , ; 2) - , , , (. 11.5.5); 3) , (. 8.3.1). . 1, -, (. 13.1.10), . , , . - , - , . , -, ( 1 ), . , , , , (. 13-63); ,

. 13-63. . . , , , , . , ; .

457

, . ( ) , , . , (. 9.4.19), , . , Xenopus, , ( ). , , , , , . , in vitro - , , (, MPF), . , , , . , , , , . , . , , ; , , .

13,5.12. [46] . 13-64 . , , - . , (- ), . , , , .

. 13-64. , , . , , . , , : . , a , . ( ), .

458

, (. . 13-48), , , / () / () (. . 13.1.10). MPF (. MPF . 13.1.11 . 13-15).13-30

13.5.13. , . , , , . . , Drosophila ; 6000 , . (. 16.5.2), , , . ,

. 13-65. . , . , . . . . . (H.W. Beams. R.G. Kessel, Am. Sci. 64: 279-290, 1976.)

459

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465

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466

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