1

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. . . . . .

2- ,

3- 1

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

1994

4 28.070 75 576.32/36 : ., ., ., ., ., . : 3- . 2- ., . 75 . . 1. . .-.: , 1994.-517 ., . ISBN 5-03-001985-5 ( - ) . . 3- . 1- (.: , 1986-1987). . . 1 , , , , , . , , , .

1903010000 023 46 92 29 056(01) 94

28.070

ISBN 5-03-001986-3 (.) 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 , . ., . ., . ., . ., . ., 1994

5

. , . , . , . . 19861987 . . , . , , 1989 ., , . , , ; , , , , , . , . , : (. 10) (. 21). , . , 7, , , , . 8, , . . . , , . . - , , , , , , . , , . . .

6

50 , . , . - , , , - . . , , , . , . . , , , . , . , , . , , ; . , : , , , . , , , . , , , , . . , . - , - , . , , . , , , . . . . ( ) 8, ( ), ( , -) ( ) 13. , . , , . , , , . Garland Publishing, , , , , , , . . , , , , .

7

. , , . . . , , . , , , . , , . , , . , , , . , . , . , . . , , - : . , , , , . , . III . , , , , . , , . , . , , . , , , , . , , , . , , , . , . , - , , . . , , , , . , , -

8 . . , . ( ), ( ), ( ), ( ), ( ) ( ) . , , . . , , , , , . 15 16. - . , , , . , - . , , , . , , .

9

. . I . I 4, . , , , , . II ; , . III , . 4 , , . . . , . . , . . .1 . (. 5-14) , . . , 5-4 , 4 5 . , , , . . . ______________________________________________________ 1 ., . . . . . . . .

10I

. 1.1. ( ). ( Tony Brain Science Photo Library.)

11

. 1.. ( ). ( Herskowitz Schabatach.)

12

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

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

1.1. - 1.1.1. , , . , ? ? , - , . , , . , , (.. ) . - . , , 2, 4, NH3 2, , . , (. 1-1). , , (HCN) (), (. 1-2). , : , , . , , , . , : . , . , , - , .

. 1-1. , . , 4, NH3 2. . U- .

1.1.2. , , , . , . , . , , () -

. 1-2. , , . 1-1. , , .

14 () . , - 20 , . , , . , . , (. 1-3). , . , , , . , , ( , poly U), , ( , poly A). , . Poly U poly , (. 1-4).

. 1-3. ( , A, U, G, ) . .

. 1-4. ( G , - U) (). (). , , . , , , (U), (), () (G). - U G - , , , . , , U . . , , . (. 1-5). , . , .

15

. 1-5. ( ). 1 . 2 . , . , . , , , , , . , , , . , , , .

1.1.3. . , . . , , . , . , . , (-) . , . GGGG . - , , (. 1-6). , . , , . , : , .

16

. 1-6. () . - . , . - , .

1.1.4. . . , , , . , , (, ). , , , , . , . , , -

17

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

1.1.5. , , 3,5-4 . - . , -

18 ( , ). , . () , , -, , . , , . . , , , , , . , , . ? , , ? , . , . . , , . , , . . . , , , . , -, (. 1-7, ). - , ( ) 50 . . 5 , , 60% . . , , . ( ), . , . - - : . , -, . , -

. 1-8. (Escherichia coli) () 4 (5). ( ) , , -, . , , . .

19 , . , , . : , , , (. 1-8).

1.1.6. , , . , , , , . , , , . , , . , (), , . , , (. 1-9).

. 1-9. , . , ( . 1-7), , , . - (, , ), . , . , -, , - : ( ), - ( ). , , , - . , , (. 1-10). : . , , , ; . 5 ( ). , . , , . , . , , , : .

20

. 1-10. . () , , . (), , .

1.1.7. , , . , . , . , ( , , , ), , 3,5-4 . . , . , , - (. 1-11). 0,3 , 750 . , , , . , , , , . , -, , , . -

21

. 1-11. Spiroplasma citrii , . (, , ): , , . , , . . , , , . , , . , , , ( . 3) : . , , . : ( ), ( ). . , . , , (. 1-12). , , , .

. 1-12. , , . 3,5 , . -

22 , , , , . . , -, , , . , .

1.2. - [6] , , . , , , . , . . 1,5 . ( , ) , .

1.2.1. , [7] - , . - (. 1-13). , , , , , , , . (. . 1-8, ). . , . 20 , , 5 . ( ) , 11 . . , , , . , . : - , , , -

. 1-13. , .

23

. 1-14. ( ). . , , , , (. 1-14). , - , , , , . 2 N2. , .

1.2.2. , , , . , , . , , . , , . , , , , : , - . , . , , . . 1-15. , , , - , . , , , , (. . ).

24

. 1-15. , . (, , , ), . () . , , . . , , . , . , . . , . , , , - , . . , , . .

. 1-16. ( ) . , , . , , . ( . L. Sogin, H.J. Elwood, J. H. Gunderson, Proc. Natl. Acad. Sci. USA, 83, 1383-1387, 1986.)

25 - , . , : . , , , . - . , , , . , , , . , . , . . , (, , ), (. 1-16). (. 1-17). , .

. 1-17. , ? , , . , 5-10 , . 75 ( , NAD-) . , . . , , . , 10%, - 17%. , , , , . ( W. M. Brown, . . Prager, A. Wang, . . Wilson, J. Mol. Biol., 18; 225-239, 1982.)

261.2.3 2 N2 , , . , ? , ( 2 N2) . , . ( , , ) . 2 , 2 . - - . , , . , - . - ( ). , 2 N2 , , , - , .. . . , , , 2 N2. , 2S, ( ) . , , , 2. . , , ( ). . 2 N2 , , , , , -, 1 . . , , , : , , .

271.2.4. . , , . , , 21%. , , , , ( ). , , . . , , . 2 2. . , , , , . , + . + . , .

1.2.5. , ? , , , . , , . , , , . , , . , , . - ( 1-1), . ( ). , , (. 1-18). , , , - , . . --

28

1-1. .

29

30

. 1-18. . . ( Daniel S. Friend.)

. 1-19. . . , , , (). (). ( J. Burgess.)

. 1-20. . , , . ( Daniel S. Friend.) (. 1-19 1-20). , , . - , , , . . , . , .

1.2.6. , : , , , . , , .

31 , . , , , , , . . . , , (, ) . , , , , , , . , Pelomyxa palustris, , , . , Pelomyxa , . . , , . , . , + ( ), . , . ( , ), , , , (. ).

1.2.7. , -, . , (. 1-20). , , . , , - . -

. 1-21. , , ( Cyanophora paradoxa). . ( Jeremy D. Pickett-Heaps.)

32

. 1-22. . - , (. 1-21). 1-1. 1-10 , , 10 100 , , , .

, , , - , , ( )

33 . 1-22 . , , , . , , . , , , . , , (. 1-1). , , . , , .

1.2.8. , , 1000 . , 1000 . , , . . , - , - . , . - . , (

. 1-23. , (). , , , . . ( George Palade.)

34

. 1-24. . . (. 1-1). , , . ( Daniel S. Friend.) ) . (. 1-23), , , . , (. 1-24), . , , , . , - (). () , ( ). . . , , . . ? , , . : . , , , . (). - , , . , , . , : , - , - . .

1.2.9. , , . , , . . (. 1-25), , , , . , , , () , , - , .

35

. 1-25. , , , . ( John Heuser.) , . , - , . , , . , .

1.2.10. , (. 1-26). , (. . 1-16) : , . , , , , , , . , , . , , . Didinium - . 150 , , (. 1-27). Didinium . ( , Paramecium), . Didinium Paramecium , , , . , - , . , , , .

36

. 1-26. , . , 10 . , , , , - , - , - , - . ( . A. Sleigh, The Biology of Protozoa. London: Edward Arnold, 1973.) Didinium, , : , , , . , .

1.2.11. , - . , -

. 1-27. . , . - , . Didinium () , . , Didinium , Paramecium. ( D. Barlow.)

37 . , , , , . - ( 1-2). , , . . ? , . , . , . . , , () , , . , , , . , . , . , , , . , , - - ( 1-2). . , , .

1.2.12. , . , 1000 , , 10 , (. 1-28). , -, (, 1 % ) . 99% ? , . , - , . , () - ,

38

1-2. .

39

40

. 1-28. , , . , , , , . , , , , . ( . . Macgregor, Philos. Trans. R. Soc. Lond. (Biol.), 283, 309-318, 1978.) . , , , , . . , , - . , , . - , , , , .

1.2.13. , . , - , (. 1-29). (. 1-30). ( ) . , : ( ) . . , , , .

41

. 1-29. , , , , .

. 1-30. . - , - . , . , , . , , . , : 1) ( ) 2) ( ). , , . , ( , , , ), : , - . , , . , , (), , . , . : ( ) . , -. , , , , - , . , . , . , . , , ( ) , . , . - , .

1.3. - , , , . , , . ? -, , , . , , ( -

42 ) ( ). , . , . , . , , , , , , . , , , - , .

1.3.1. , . , . . , , , , . , , , ( ). , : , (. 1-31), , .

. 1-31. ( ) , (Chondromyces crocatus). , , . ( P. L. Grilione, J. Pangborn, J. Bacteriol., 124, 1558-1565, 1975.)

43

. 1-32. , . , . ( , ) - , , . , , , , . , Chlamydomonas, , . , . ( Gonium) , 4, 8, 16 32 . , , . , . . - Volvox; 50 000 , . Volvox , (. 1-32). Volvox - , . , , .

1.3.2. Volvox , . , . . , , . , Volvox : . , , . , , , . , . , , . . , , .

441.3.3. , - . . Volvox, , , . - , ( ), . , . ( ), () . , ( ), , , , . , : . , . , . , , : , . .

1.3.4. , , . , . - , , - . , , , , . : - - . , (. 1-33). , , . , . , . , .

45

. 1-33. . () , , () . (. . ), . , . , , ( ), , . , , . - , , . , , . . , . , . , (. 1-33, 1-34 1-35). , . , , ,

. 1-34. . , . ( Amata Horubruch.)

46

. 1-35. , , , . , , , . - , . , . , , , , , .

1.3.5. , , . , . , , . , , , . , . , , , . - , , , , . , , . , , . , . , , . , . , , , , .

1.3.6. - , , . , .. , . , . , , , , , ,

47 . , , , , . , . , .

1.3.7. , , , . - ? , , . . , , , - . ,

. 1-36. , , . () , () . ( . Haeckel, Anthropogeni, oder Entwickelungsgeschichte des Menschen. Leipzig, Engelmann, 1874.)

48 , , . , , (. 1-36). . , . , , . , , . - . , . , , , . - , . - , . , , . , , , .

1.3.8. , . . , . , , , , . , . . . . ( ) . . , , . , , . , () : - , - . - , . , -

49, ( , ), . , , . , , , . .

1.3.9. 200 , , , . 200 , , , . 1-2 . , . , , . ( 1-1) . - , : , . (. 1-37). 1. - . , . , . 2. , . : , , . , . , , , . , , - .

. 1-37. : , , .

50

. 1-38. ( ) - - . ( J. yles, D.F. Bainton, Cell, 24, 905-914, 1981. Cell Press.) 3. , , , , , .

1.3.10. . . : - , - . . - . , , . . , : . , , (. 1-38). : , , , . , , .. . - . , , , , . , (-) - , . , , , , , - . , . , , , . , , , ( ), ,

51 , . , , , .

1.3.11. [17, 18] - , , . , , . - . , , . , , . , , , . , , . , Gonyaulax (. 1-39) 24- , , . , . , , . , , , , , , , . .

. 1-39. . Gonyaulax polyedra, . 22 58 , 18 . . ( ) Gonyaulax. ( - D. Njus, V.D. Gooch, J. W. Hastings, Cell Biophys., 3, 223-231, 1981; - John Dolge.)

52

. 1-40. . , , , , . , ; , . (, AMP, ). , , , , . , , , , . , , , . , , . , ? . , - - , (. 1-40). , , . , .

1.3.12. . , , , , , . , ,

53

. 1-41. , . , , . ( Zoltan Gabor.) (. 1-41). , , (. 1-42). , . , : , , .. . . , , . , . , .

54

. 1-42. . (Drosophila). , , . , -. . , . - ( ). - . , , , . , . . , . ( - Rudi Turner and Antony Mahowald; - N. Stausfeld, Atlas of an Insect Brain. New York, Springer, 1976.) . - - . - , , , . . , , , . -

55

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

56

. 1-43. . , . ( , , .) , . , . , , , .

- , .

57 , , . , . . , , (. 1-43). . - , . - . , . () .

Bendall D. S. ed. Evolution from Molecules to Men. Cambridge, UK. Cambridge University Press, 1983. Curtis H. Biology, 4th ed. New York. Worth, 1983. Darnell J. E., Lodish H. F., Baltimore D. Molecular Cell Biology, Chapter 25. New York. W.H. Freeman, 1986. Darwin C. On the Origin of Species. London. Murray, 1859. Reprinted, New York, Penguin, 1984. Evolution. Sci. Am. 239(3), 1978. (An entire issue devoted to the topic.) Keeton W. ., Gould J. W., Gould C. G. Biological Science, 4th ed. New York, Norton, 1986. Maynard Smith J. The Theory of Evolution, 3rd ed. New York, Penguin, 1975. Watson J. D., Hopkins N. H., Roberts J. W., Steitz J. A., Weiner A. M. Molecular Biology of the Gene, 4th ed. Chapter 28. Menlo Park CA. Benjamin-Cummings, 1987. Wilson E. B. The Cell in Development and Heredity, 3rd ed. New York: Macmillan, 1925. (Reprinted, New York: Garland, 1987.) 1. Ferris J.P., Usher D.A. Origins of life. In: Biochemistry, 2nd ed. (G. Zubay ed.), pp. 1120-1151. New York: Macmillan, 1988. Miller S. M., Orgel L. E. The Origins of Life on the Earth. Englewood Clifis, NJ. Prentice-Hall. 1974. Schopf J. W., Hayes J. M., Walter M. R. Evolution of earth's earliest ecosystems: recent progress and unsolved problems. In: Earth's Earliest Biosphere: Its Origin and Evolution (J. W. Schopf ed.), pp. 361-384. Princeton, NJ. Princeton University Press, 1983. 2. Eigen M., Gardiner W., Schuster P., Winkler-Oswatitsch R. The origin of genetic information. Sci. Am., 244(4), 88-118, 1981. 3. Cech T.R. RNA as an enzyme. Sci. Am. 255(5), 64-75, 1986. 4. Alberts . . The function of the hereditary materials: biological catalyses reflect the cell's evolutionary history. Am. Zool. 26, 781 - 796, 1986. Darnell J. E., Doolittle W. F. Speculations on the early course of evolution. Proc. Natl. Acad. Sci. USA, 83, 1271 - 1275, 1986. Orgel L.E. RNA catalysis and the origin of life. J. Theor. Biol. 123, 127-149, 1986. 5. Rogers M. et al. Construction of the mycoplasma evolutionary tree from 5S rRNA sequence data. Proc. Natl. Acad. Sci. USA, 82, 1160-1164, 1985. 6. Vidal G. The oldest eukaryotic cells. Sci. Am., 250(2), 48-57, 1984. 7. Clarke P. H. Enzymes in bacterial populations. In: Biochemical Evolution (H. Gutfreund ed.), pp. 116-149. Cambridge UK. Cambridge University Press, 1981.

58Doolittle W.F. Archaebacteria coming of age. Trends Genet., 1, 268-269, 1985. Wese C. R. Bacterial evolution. Microbiol. Rev., 5, 221-271, 1987. 8. Wilson A. The molecular basis of evolution. Sci. Am., 253(4), 164-173, 1985. 9. Dickerson R. E. Cytochrome and the evolution of energy metabolism. Sci. Am., 242(3), 136-153, 1980. 10. Cavalier-Smith T. The origin of eukaryotic and archaebacterial cells. Ann.N.Y. Acad. Sci. 503, 17-54, 1987. Margulis L. Symbiosis in Cell Evolution. New York, W. H. Freeman, 1981. 11. Vossbrinck C.R., Maddox J.V., Friedman S., Debrunner-Vossbrinck B. A., Woese C. R., Ribosomal RNA sequence suggest microsporidia are extremely ancient eukarvotes. Nature, 326, 411-414, 1987. Yang D., Oyaizu Y., Oyaizu H., Olsen G. J., Woese C. R. Mitochondrial origins. Proc. Natl. Acad. Sci. USA, 82, 4443-4447. 1985. 12. Sleigh M. A. The Biology of Protozoa. London, Edward Arnold, 1973. Sogin M. I., Elwood H. J., Gunderson J. H. Evolutionary diversity of eukaryotic small-subunit rRNA genes. Proc. Natl. Acad. Sci. USA, 83, 1383-1387, 1986. 13. Buchsbaum R. Animals Without Backbones, 2nd ed. Chicago. University of Chicago Press, 1976. Field K. G. et al. Molecular phytogeny of the animal kingdom. Science, 239, 748-753, 1988. Margulis L., Schwartz K. V. Five Kingdoms: An illustrated Guide to the Phyla of Life on Earth, 2nd ed. New York. W.H. Freeman, 1987. Shapiro J. A. Bacteria as multicellular organisms. Sci. Am., 258(6), 82-89, 1988. Valentine J. W. The evolution of multicellular plants and animals. Sci. Am., 239(3), 140-158, 1978. 14. Bode P. M., Bode H. R. Patterning in Hydra. In: Pattern Formation (G. M. Malacinski, S.V. Bryant eds.), pp. 213-244. New York, Macmillan, 1984. 15. Raff R. A., Kaufman T. C. Embryos, Genes, and Evolution. New York. Macmillan, 1983. 16. Maynard Smith J. The Evolution of Sex. Cambridge, UK. Cambridge University Press, 1978. Michod R. E., Levin B. R. eds. The Evolution of Sex: An Examination of Current Ideas. Sunderland, MA. Sinauer, 1988. 17. Winfree A. T. The Timing of Biological Clocks. New York. W. H. Freeman, 1987. 18. Bullock . ., Orkand R., Grinnell A. Introduction to Nervous Systems. San Francisco. Freeman, 1977.

59

2. , , , , . , 150 (Wohler), (vital force), . , , - . , , . , : , . , . , - , . , . , , . . , : , . . 3 (), .

2.1. 2.1.1. - [1] , (, , N, , , S) 99% . , , (. 2-1). ? , , - . 70% , . , .

60

. 2-1. , ( ), . . , , 50% , . , , , ( 2-1). , , , , , . ; , . . . , . , (, N ), , , ( 2-2). . , . , ; , (. 2-2). , , ( ) . , , .

612.1.2. [2] , , (3), (), () (NH2), . , , . 2-2. . 100 1000, 30 . , , , . , (. ). , ( ) . , , , , . , . ( . 3) , , . , : , , . , . , , , , (. 2-1).

2.1.3. [3] - - (2)n, n - . , , 6126 (. 2-3). . 2-3, , . , , , , , . . -, , . , , . -, , ( 2-3). -

62

2-1. .

63

64

2-2. , .

65

66

. 2-2. , . - . 2-1. , % (, ) 70 1 1 0,4 0,4 1 0,2 26

1 20 250 100 100 50 ~300 ~3000

(, . .) , . , - , . , , 11 (. 2-4), (6126), , . - ; (, ) , , . . (. 2.3.2) 2 2.

. 2-3. , - . . ( ) , (). . (p-D-). - (); , . . 2-3.

67

68

69

. 2-4. , D-. , . , , , , , , . , .

. 2-5. . ( ) . . : 6126 + 2 + 62 + 62 + . , . NADH (. . 2.3.1). , ( , - ), . , . , (, ), .

2.1.4. - , (. 2-5), : , () , , , () . ,

70

2-4. , , .

71

72 . , , , , ( 2-4). , , ( ) . (). , ( 2-4); , . . , , -, , . - . , , -- , , , . , . , - , . , , , . , , , . . , ( 2-4). , , , .

. 2-6. , 7. - , , . .

2.1.5. - , , . , , (. 2-6). - , (. 2-7). 20 , - ( 2-5). 20 , , . , , 20 , - , .

. 2-7. . , . . .

73

2-5. 20 , .

74

75

. 2-8. . +, , , . , , + , ( ). , ( ) . , . : , . , , . 7 . , 5 20 (. 2-8), , ( 2-5). , , , .

2.1.6. [5] ( , + ) ( ), . , , . (), () (U) , ; (G) () - , . , ( 2-6).

76

2-6. , .

77

78

. 2-9. (). () () .

. 2-10. , . - . G-A-T-C. . (. 2-9) , , . ; , , , . , , , . , AMP (cAMP) - . . 1, . - , , 3'- 5'- (. 2-10). , , . , , ; A, U, G . , ( -2 ), ; , , G . . ( ) - G U - . .

79 - . , , . , , . , . , , , - . , , . , , .

2.2. [6] . , . , , , , . , , , : , .. , , . , . , , , , , . , , . . , , . ? , : , , .

2.2.1. [7] , , , (. 2-11). , , . , ,

80

. 2-11. : , , , . . , . ( ) , , . , . . , . , . , - . , .. . , , , , , . , XIX , , , (, ), . , . , , , , , . , ; . . . 2-11 , , , , . - , . , , , . , . , , . , , , .

2.2.2. [8] () , - . . , -

81

. 2-12. . , . . 7; , . ( ) ; , , NADPH. ( ) NADPH , (. 2-12). ( ) + 2 + 2 + + 2, . . . , . , 2 , , .

2.2.3. , , , .. , - . , , , . - , , , . . ,

. 2-13. . , . , . , 2 .

82 , . , , ; 2, , , . , - , (. 2-13). , , .

2.2.4. [9] . , 2, - 2. , , , , 2 2. , , , . , . , , . ; , . , (, ) - . , Fe2+ , , Fe3+ , , 1-. , , . , , , , , , . , , , (. 2-14). ( , , ) 2 2, . .

. 2-14. . , ( , ).

832.2.5. [10] 2, - 2, , : , (. 2-15), . . . , , , , . ( ), , (. 2-16). , . . (), , () . , , , 1014 . , .

. 2-15. , . X Y. , X , , .

. 2-16. , , . ( X) ; . ( ) . , , , - . 1, ( X Y) .

2.2.6. , , [11] , . , , , , ( ),

84

. 2-17. , . . 2 2, . . ; . , , , , . . . . . . 2-17 . , , (. 2-17, ). , (. 2-17, ). , , . , - - - . (), , , , , . , , , . , (. 2-17), (. 2-17, ); , - , - , .

2.2.7. [12] ? , , (i) (. 2.4.1). , , , , . , , . , ,

85 (ADP PI); . , , , . , , , , , . . , , , , . . , , .

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

2.3. [6, 13]2.3.1. , , , , , , . , , (. 2-18). , . 1 : , , . , , , . 2

86

. 2-18. , . , . . , , , (-) (. 2-19). - . 3 - 2 20. . ,

87

. 2-19. - (-). , 2 (. . 2-18), (). , , 2 2, i + ADP -> . , , , , . , , . , , 1 . , ADP .

2.3.2. 2 , . , , , . ,

88

. 2-20. . . 4 , , . 5 6 NADH (. . 2-21). (. 2-20). , : 1) 1-4 -3- ( , ); 2) 5 6 -3- , ADP ; 3) 7, 8 9 , , ADP, 1. ( ), 5 6. , . - , , - ,

89

. 2-21. 5 6 : NADH (. . 2-20). 5 -3- -3-. ( -: ) -3- - NAD+ (. . 2-22). , . - , . . , 6 ADP, , . (. 2-21). , ADP NAD + NADH (. 2-22). , . , , NADPH , , . -3- (. . 7.3.4). 3 , , 2 2. (. . , ) (, ), , . , , 2 ( ), ( ), . , , , , 5 , NAD + , (. . 7.4.1).

2.3.3. [14] . , . ( ) , , , -

90

. 2-22. NADH NAD+ - . . NADH NAD + . NAD ; , ( ) (AMP). NAD + , ( ), (-, ~), NADH. , . - . . NAD+ NADH. , , . - NAD + , NADH, + (. . 7-18). . , , , -, . , ( 3 . 2-18), . ( 3) ( ) : .

2.3.4. [15] - , -. , , , - , . . 2-23, - , , ( ). ,

91

. 2-23. . , , . 2, - NAD+ FAD. , (*). . 7-14. 2, - . ( , , , 2, , .) 2, , ( ) . , , . (- ) , , . ( , a GTP, ADP + GTR + GDP.) , , - ( -) ; NAD+ NADH, (FAD) - FADH2. , , ; ( ) . , 2 , , . -

92

. 2-24. . (, , ) , . , + . + ; (. . 7-35). , 2. , , ( ) -, NADH. - , , -, . , , , , , - . , , 2-. , , , , .

2.3.5. [9, 16] ; . NADH FADH2 , , 2. , , . ; . NADH FADH2 , . . , - ( + ). , , . , , , . NADH FADH2 -, , ( ), . - , -. , . , , . , , ,

93 (. 2-24). . ; (-) ADP . , . - . 7.

2.3.6. , , . . - , 2/3 . , . , . , . . , , , . , . . , . , , . . , . 20 , , (. 2-25): . , . - ; . , , : , . , ( ),

. 2-25. , .

94 - , , . , , , . 2 2. , , . , .

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

2.4. [17] . , . , , : . , . ( ) .

2.4.1. [18] , . , , . , : . , , , , , -

95. , . , . , , . , ; , G. , , G, , ( 2-7). , ; , G . , ,

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

G, G . . , G . . : ,

G +1 13 /. (, 6 1023 .) , G 12 /. , , , , , .

2.4.2. , - - ( ), : - + -

- + 2.

( ), -, . , , -

96 , . , , , , "" . . , , , , . ? . , , : . ( , ), , , . , ,

, (, (. 2-11} " "). , . : , , , , , . , : , " ". , , ( ). (, ) , . , , ( ). , , , , . , ( o ) ( h), = - h. , , . , . ,

, ( V . (), . ( \/) , , (5) . , . (H) -

, ( = + PV). , ( ), , , . , , , "", , ( ), "". , - h = . , , :

, 1000 , "" . , , . "" . : , (. . "" ), , "" "", ( 10298). , "" "" 50:50, , , , . , " . "", , : " ". 2-7. .

97, S ( ). , , , , , . (S). : ( S) ,

,

; R (R =2 -1 -1), a S (. .). ( "" ) ( "", "") , . , = 1, = 1000! (500! 500!) = 10298. , , , "" "", R ln(10289), 1370 . . (6 1023 ). , (/ > 1), , S (. . , , . , , , . , , , , , AS ( SK ) , (h), () :

S > 0), .

, G , . , : ? ( ) , , , , - , . , , ( SOK< 01, S>0), S ( S = S + S), 0. , . (""), , . , , " " ( ), . , , (), (), (V), () (S) . () . (G) , . V, , G ( + PV), , S . . (G G ) G , , , . ( G) ,

- S. ,

= = -h

h/T

S), a S S. .

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

, :

G , , , -, , .

10 . , G = , . , ; .

G , ,

G

.

98 , . , - - -, , (. 2-17). G (. 2-26), , . - - -, , - , -, -. - -3 ( -1. - + 2. - + -

), :

--

+ ADP,

- + i.

- , : - + - - ADP + i. , , , ( ). (. 2-27). G ADP 11 13 / (. . 7.1.10). -

11 13 /. G . . , - .

. 2-26. , ,

. 2-27. , . ( -- ), . . , .

99

. 2-28. : , . , , . 2-26. , , , . , , , (+ ).

. G = 13 /. , AMP PPj () (. 2-28). ; 26 /. ? - - -. - -1. - +

G,

, , + 10 /, , ,

-

. :

-

-

+ AMP,

- + PPi, 2. - + -3. PPi + 2 2i. : - + - - + 2 AMP + 2i. , , - , (, 2). (

100

. 2-29. ( ) , . (, ) . - , , , ; . , . 3) - , ( , 2). , . ( ) . 2-29.

2.4.3. , . , . , - , , (. 2-2). - , . - - (-), . ,

101 2-2. , 1) NADH, NADPH + (-) S- ____________________________________________ 1) , . - , . , . . (. 2-19). , , , . - , (. 230). , -, , , . ( ). - , - .

. 2-30. . - ( -). -. , - (HCO3-), , . .

1022.4.4. , . , , , , , , . NADH (. . 2-21). NADH , ( -) . NADH NADPH . , , - . , (. 2-31). , , , - NADPH ( + ) (+ + - = 2). NADH, -, NADPH, , , (. . 2-22). , NADH, NADPH - , , , . NADH NADPH : NADPH , (. 2-32). . , NADPH . NADH, , , , NADPH . -

. 2-31. . - NADPH ( + ) .

. 2-32. NADPH, NADH (. 2-22) , ( ) .

103, NADH NADPH .

2.4.5. , , ( ), . . , ( ) . , . 2-33. , , . - . (. . 2.4.2),

. 2-33. , . (). , . - ().

104

. 2-34. . , . : , , (. 2-29). , , . , , , (. 2-34). , , . . , .

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

2.5. [19]2.5.1. - , , , . 2-35, , . 2-35. , . . 500 ( ). . 10000 , - . ( ) -. ().

105

106 . 0,1 , , . , ; , . AMP ( , , , , ; - . B . , , , , . . . . , , . , . . , .

2.5.2. [20] , . . , ; , , (. 2-36). , , , , , . , . 2-37, , . ; , , , , ,

. 2-36. . Z , , .

107

. 2-37. , , . , . ( ), . . . 3.

2.5.3. [21] , , . , , . , , , . . , , , ( 1, 3 9 . 2-20) .

108

. 2-38. , , . ( ) ( ), . , 1, 3 9 . , .

109

. 2-39. , , , NADH. , , . . , (), - , (. 2-38). , , GTP. . (. 2-38) , , , . , , . . 3 , . -6- ; . AMP, ADP , , . , , ; , ( ) ( ). , () ( -1,6- -6-), . , , , - , . , ADP, ( + -6- ADP + -1,6-), - . , . , (. 2-39). . , . , , , .

2.5.4. [22]

110 . , , ( ) . , ( ) (, ) . , , . , . , , , -. - , (, ), .

2.5.5. , [23] ( ). , , . , , , , , , , . , . , . , - , (. 2-40), a . , , . . (. 2-41). , . , , , NADH 2 . , , . , , , , , , -

. 2-40. - - , . -.

111

. 2-41. . , - . . , . . - . , , , . , , , - . , , , , , . . , , , (. 2-42). , , .

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

. 2-42. . , . - - .

112 Herriott J.; Jacobson G; Marmur J.; Parsom W. Papers in Biochemistry. Reading, MA: Addison-Wesley, 1984. *Lehninger A.L. Principles of Biochemistry. New York: Worth, 1982. Stryer L. Biochemistry, 3rd ed. New York: W.H. Freeman, 1988. Wood W..; Wilson J.H.; Benbow R.M.; Hood I.E. Biochemistry: A Problems Approach, 2nd ed. Menlo Park. CA: Benjamin-Cummings, 1981. 1. Henderson L.J. The Fitness of the Environment. Boston: Beacon, 1927; reprinted 1958. ( .) 2. Ingroham J. L.; Maaloe .; Neidhardt F. C. Growth of the Bacterial Cell. Sunderland, MA: Sinauer, 1983. 3. Roehrig K. L. Carbohydrate Biochemistry and Metabolism. Westport, CT: AUI Publishing, 1984. Sharon N. Carbohydrates. Sci. Am., 243(5): 90-116, 1980. 4. Robertson R. N. The Lively Membranes. Cambridge, U. K.; Cambridge University Press, 1983. 5. Saenger W. Principles of Nucleic Acid Structure. New York: Springer, 1984. 6. Hess .; Markus M. Order and chaos in biochemistry. Trends Biochem. Sci. 12,: 45-48, 1987. Lehninger A.L. Bioenergetics: The Molecular Basis of Biological Energy Transformations. 2nd ed. Menlo Park, A: Benjamin-Commings, 1971. **Schrodinger E. What is Life? Mind and Matter. Cambridge, U. K.: Cambridge University Press, 1969. 7. Dickerson R.E. Molecular Thermodynamics. Menlo Park. Ca.; Benjamin, 1969. Klotz I. M. Energy Changes in Biochemical Reactions. New York: Academic Press, 1967. 8. Raven P. H.; Evert R. H.; Eihhorn S. E. Biology of Plants, 4th ed. New York: Worth, 1981. 9. Racker E. From Pasteur to Mitchell: a hundred years of bioenergetics. Fed. Proc. 39: 210-215, 1980. 10. Fothergill-Gilmore L.A. The evolution of the glicolitic pathway. Trends Biochem. Sci. 11: 47-51, 1986. Shulman R.G. NMR Spectroscopy of living cells. Sci. Am. 248(1): 86-93, 1983. 11. Lipmann F. Wanderings of a Biochemist. New York: Wiley, 1971. 12. Schlenk F. The ancestry, birth and adolescence of ATP. Trends Biochem. Sci. 12: 367-368, 1987. 13. McGilvery R. W. Biochemistry: A Functional Approach, 3rd ed. Philadelphia: Saunders, 1983. Racker E. A. New Look at Mechanisms in Bioenergetics. New York: Academic Press, 1976. 14. Kornberg H.L. Tricarboxylic acid cycles. Bioessays 7: 236-238, 1987. 15. Krebs H. A. The history of the tricarboxylic acid cycle. Perspect. Biol. Med. 14: 154-170, 1970. Krebs H. A.; Martin A. Reminiscences and Reflections. Oxford, U. K.: Clarendon Press; New York: Oxford University Press, 1981. 16. Pullmann M. E.; at al. Partial resolution of the ensymes catalyzing oxidative phosphorylation. J. Biol. Chem. 235: 3322-3329, 1960. 17. Prigogine I.; Stengers I. Order out Chaos: Man's New Dialogue with Nature. New York: Bantam Books, 1984. 18. Eisenberg D.; Crothers D. Physical Chemistry with Applications to the Life Sciences. Menlo Park, A: Benjamin-Cummings, 1979. Reich J. G.; Selkov E. E. Energy Metabolism of the Cell - A Theoretical Treatise. New York: Academic Press, 1981. 19. Martin B.R. Metabolic Regulation: A Molecular Approach. Oxford, U.K.; Blackwell Scientific, 1987. ***Newsholme E.A.; Start C. Regulation in Metabolism. New York: Wiley, 1973. 20. Pardee A. B. Molecular basis of biological regulation: origins of feedback inhibition and allostery. Bioessays 2: 37-40, 1985. 21. Hess B. Oscillating reactions. Trends Biochem. Sci. 2: 193-195, 1977. 22. Cohen P. Control of Enzyme Activity. 2nd ed. London: Chapman and Hall, 1983. * . . 3- . -.: . 1985. ** . . . 2-. ., , 1972. *** ., . . .: , 1977.

113

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

3.1. [1] 10000 1 ., . . . 2 , (. 3-1). , , 75% ; (. 3-1). . 2, , , , (. . 2-33). . , , , ; , , , , , . (), , .

. 3-1. . , 60 .

114 3-1. , % 1) 2 (Na , K , Mg , , l . .) + + 2 + 2+ -

. coli 70 1 3 15 6 1 2 2 2-10-12 3

70 1 3 18 1,1 0,25 3 2 2 4-10-9 3

1 2000 1) , , - . , ; , ( ) ().

3.1.1. [2] , , . . . , , . : , . , ( 2-1). . 3-1. 30-300 , , (. 3-2) , 37 . , , -

115 , , . , (. 3-2). , , , . , , , , . , 3-1, ( ). ( ), . , , , , - . ; , , .

3.1.2. , [3] , , (. 2.4.5). , : , . , , 3 2, 2 - 1 . . , - , , . 3-3. (. 3-4). , , . . , , (), ( ). : , .

3.1.3. - [4] , . , , . , ,

116

3-1. , .

117

118 3-2. , / 1)

________________________________________________________

, 0,15 0,25 0,30 0,20

90 80 4 0,1

90 1 1 0,1

1) , . (/). - , 1000 1. - (), 0,24 . , . , .

, , , (. 3-5). , , . , 1 1 , 2 4 , 10 - 100 .. , - , . , , . , , 0,2 10 , . . , , , , (. 3-6).

. 3-2. , , .

119

. 3-5. . - . - . . 3-3. , . , , . (), () ( ) . - . , , .

. 3-4. - . , , , . , ,

. 3-6. , . , ; , . , , , . ( . . Bridgman and . S. Reese, J. Cell Biol. 99: 1655-1668, 1984. Rockefeller University Press.)

1203.1.4. , [5] , , . ( , ), , . ,

. 3-7. . / 2, . (3) . , , 2 , . [] [] ( ). , , - . 3 + , , , + . , , , , ; (), . , ; Kd, .

121 3-3. () [ ] + = (/) [A][A (/) 105 104 103 102 10 1 10-1 10-2 10-3 10-4 10-5 -7,1 -5,7 -4,3 -2,8 -1,4 0 1,4 2,8 4,3 5,7 7,1

+ , , , AG. 37

G0 ( 1,0 /). , , , . , , . , , . , . , - . , , . , , . , , (. 3-7). : , . G. (. 3-3), G . 103 1012 /, 4 17 /, 4 17 . , , , , , (. . 10.2.1). , , , (. . 1.3.1).

3.1.5. [6] . , ~ 1000 , 106 . , , . : 1) ( ), 2) () 3) . . , , , , . , , . -

122, -1 , 106 ; , 105 .. , . , , .

3.1.6. [7] : , , . , , , . , , (. . 2.4.1), . , , (. . 3-3). , . , , , . - . . . , , , . , . , . , , , . , , . , (. . 3.2.4).

, . . ; , , , .

123 , . .

3.2. [8]3.2.1. [9] , , , . , - , , . , . . , . , . . , , ? XIX , ( ) . , , , () , . 1944 . , , . 50- , , , . , ( ), , , .

3.2.2. , [10] , , . - , -, (), (), (G) (). , 5'- 3'- ( 2-6). , .

124

. 3-8. . () (). , , (. 3-2). ? ? . 50- , , . , , , - , . , . 1953 . , (. 3-8 3-2). , , , - . , . ( G, ) ( ) . , , G ( ) . , , , , (, 13 36%), : G - . , , G , . , .

1253.2.3. [11] , -. , . , -, . ', , , ', ' . , ', . , . - , G, - , . , , , . , , 4n. , (3109 ). (. 39), 500000 ! , . . 3-10. - 3' . ; (. . 5.3). . . (. 3-10). - , . , (. 3-11).

126

127

128

. 3-9.

- ( ,

). ( ), . , , .

3.2.4. [12] - . , ( 109 ). , , , G. - , .

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

129

. 3-11. . . . , , . , , , . 40- , , , . , , , . . ; . . .

3.2.5. [13] . : , . , , . , - , ; 50- , (. 3-12), , . , . , , , . , , , . , . - - , :

. 3-12. . - , , , . , 2-5. (-S-S-) . ( ), , (. . 3-48).

130 , , . , . , , .

3.2.6. - [14] , , - , , - , . , , : 1) 2) () (U) (. . 3-6). , , . ; . , . - . . . -, - -. , , . , . , , , (. 3-13). -,

. 3-13. -, . , . - ( ) (). , .

131 , () (); - () (. . 3.2.10), (, , . . 3.2.10) . , , , , (. . 10.2.1). , . . , , , . , - : 104 , 105 , 4 109 .

3.2.7. , [15] , . 1977 . , , ( ), ( ). , , , ( ). , , , , . , , (. . 3-13). , , , . , , . . , , , , , , , , .

132

. 3-14. , . 5'- 3'- . .

3.2.8. [16] - - 60- . , - - . , , , , (. 3-14). . , , , . , , 43 = 64 (, ). , 20 , , ; . , . 3-15, : , , .

3.2.9. [17] - , . -

. 3-15. . () . , GUG GAG . , U ( 2-5).

133

. 3-16. . . , ( ) . . , . . , , , , , . . . ; . - , , . ( 80 ) , ( ). , , . . , , . , . L- , (. 3-16). L : , (); , 3' , (. 3-16, ).

3.2.10. [18] , (. 3-17). , , 50 , (), . , , . -

134

. 3-17. . - . . () . , . . , , 5'-. . , (. 3-18). , 3'- . : 1 20 . . 5.

3.2.11. [19]- . 1981 . , . Tetrahymena . , . , in vitro . , , . 3-19.

. 3-18. . 5'- 3'-, . , ; .

135

. 3-19. , Tetrahymena. G , . 3'- , . , 400 , . , : , , (. 3-20).

. 3-20. , Tetrahymena. , , . 3-19, , G . , . .

136

. 3-21. , . 3-19 . 3-20. , . 240 ; , . , , ( 4). , . 3-19, , . , , Tetrahymena, , . , 1,5 . . . , , , - , . , , . , , , . , , . ? , . Tetrahymena, , . 3-21. ( - . . 3-16) . , , . . 1, (. . 1..4.).

137 . , , GC- - . , , . , , . . . , . , , , . . . , .

3.3. [20] , (. . 3-1). ; , . , , . , : , . , , - , . - , . , , . , 20 , . , , -, , .

3.3.1. [21] , . {).

138

. 3-22. , . , . , . , . , (. 3-1). , , . . , ; . , . , , . , ( ). , , , (. 3-22). , . , ; , , (. 3-23). , , . , -

. 3-23. ( ), . .

139

. 3-24. . ( SS-) SH- , , (. 3-24). , , SS-. , SS- ( ) , , SH- (. . 8.6.11). , : , . , , , . (. ). - .

3.3.2. [22] , , , . . 100 . , . , , , . , , .

140

. 3-25.

- - . 115

.

- , , .

.

-. ,

-, , ,

;

- (. . 3-27).

1951 . , .

-

-.

- ,

-

- - (, ).

- (core) ( ) .

. 3-25 ;

-

180, . , . - , . - (. 3-26); , , - , (. . 6.2.1). - . -, , . , -

- , .

141

. 3-26. - - , . . ( 153 ); - . . -. -, . . . , ( . R - (. . 3-27).

. 3-27. - -. , - . . - ( ). . - ( ). ( R . 3-25 . 3-26); . ( Richard J. Feldmann.)

142 -, . - - . 3-27.

3.3.3. [23] . , , - , . , , (. . 14.2.6). , . , (. 3-28). - , , , , , . . 3-29, .

. 3-28. - , . (), , .

. 3-29. , , . , . .

143

. 3-30. 300 . (D. . Metzler, Biochemistry, New York; Academic Press, 1977; ). , - - . , , , , - , . . 3-30 , 300 . , , .

3.3.4. [24] , . . - -, . - , , , . , 50 350 ; -, , (. ). , , . , , , . (. . 3.1.1), . . , , - , 58 , . , (. 3-31, ), , (. 3-31, ). , , (. 3-31, , ). , , , (. 3-32). . 3-33 , , . , , .

144

. 3-31. . . , . , - . . , (. 3-1). . , , - . . , (-), (-), ; . . , . , , , . ( Richard J. Feldmann; - Jane Richardson.)

145

. 3-32. . . b562, , -. . NAD- -, - -. . -. - , , -, . , , , , . ( Jane Richardson.)

. 3-33. , . , . AMP, , . , - . , , , , ( ) - . ( Jane Richardson.)

146

. 3-34. . ( 149 245- ). , 195 . , , (. . 3-35). . . ( J. Greer, Proc. Natl, Acad. Sci. USA 77: 3393-3397, 1980.)

3.3.5. 20 , 20202020 = 160000 , 20n . , 10390 300 . , , , . . , , . . , . , .

3.3.6. [25] , , (, . 10.5.1). , - , . . , , .

147

. 3-35. () (). , . . ; (. . 3-47). ( ) , , . () , , , , - , . , 40% (. 3-34). , : (. 3-35). . , , -, , , . , . . , , . - , , , . , . , , , .

1483.3.7. [26] . : 1 . , 40000-50000 ( 300-400 ); . , - , , (. . 10.5.4). , , , . , ,

. 3-36. , , , . , - .

. 3-37. -3-- . ( ). - , -- . , , . 2-21. , . ( Alan. J. Wonacott.)

149

. 3-38. . , , , . . - ; ( ) , - (. . 3-33). - - , lac- er-. , - , . . , 40 , . , () - 839 , . , , , - ( ), , ( ), . , , (. 3-36). . 3-37 . , , (. . 3-28). . , , , .

3.3.8. [27] (. . 4.6.6). . , , - . , , , . , , , (. 3-38). . , . , , . , , , .

150

. 3-39. . , . . , . , , . , , , , . , , Drosophila, , , , . , . , .

3.3.9. [28], , . , , , , , , , , . : 1) ; 2) , ; 3) , .

3.3.10. [29] , - , . , . , (. 3-39 . 3-40). , . . , , (. 3-41). . ,

151

. 3-40. pa, (). , . 3-33 . 3-38, . ( Jane Richardson.)

. 3-41. . . 3-3, , , . , (. . 3-3). , , ; . , , , , , (. 3-42, ). . (. . 6.2.8). (. 3-42, ). (. 3-43). - , - ; . , , , . , . , (. 3-43). , , .

152

. 3-42. , . . . , , - "coiled coil". . . , .

. 3-43. . , , . 60 . , . , (TBSV) 33 . () () , , 60 . . 180 ( 386 ) , 4500 . , ( ). ( Steve Harisson; - John Finch.)

153

. 3-44. (). , , . . ( Robley Williams.)

3.3.11. [30] , , , . . (). , (. 3-44 . 3-45). , . : - , . , , - . 55 (. . 5.1.8). , . , . , , : , .., . , . , , , . , . . 3-46 . , . , . , (. 3-47).

. 3-45. . 6000 , 2130 ( 158 ).

154

. 3-46. , : . , ; . . . . , .

. 3-47. . , , , . , , . 3-46, .

3.3.12. [31] , , . , , , , , . . , , , , . , , , .

. 3-48. - , , . . , , , .

155, . , (. 348). , , , , , , , .

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

3.4. [32] , (. . 3.1.1). ( ) , . , . , . . , , , . (. 3-49) . , . , .

156

. 3-49. - , , . , (. . 3-40). ( Tom Steitz.)

3.4.1. [19, 33] , . . -, , . , . ( ) , . , , , . | ( 2-1)

157

. 3-50. - . , , (. . 3-35). , , 195; , . 3-53. , . -, . , , , . , . (, 2- , . 3-50). , . , , , , . . , , . , . : ; , ; , (. . 2.4.3). ,

158

. 3-51. , , , , , . , . , , , . . , . , (. 3-51). . 3-52, , .

3.4.2. - [34] . , - (. 3-52, ). - , . . -, , . , . , , , . , . , . , . . . , , (. 3-53). .

. 3-52. . . - . . . . ( Richard J. Feldmann.)

159

. 3-53. . . ( ) : , 195 (. . 3-50). , . (. 3-54). , ( Vmax) , . , , 1000 , 106 . - , , (. 3-54). , .

3.4.3. , , . . , , , , (. . 3-7). + 108 , + 108 . , . , .

. 3-54. V , Vmax. , , . , , . , .

1603.4.4. , [35]. : , . , . 2 ADP (. . 2.4.2). , , , , . , , , . .

3.4.5. [36] . , , , . , . , 1-2 (.. ). , 107 , , , . . , , : , . , . , . . (10-6 ), . ? . , , . . . , . . - -

161

. 3-55. . - - . 2-40. , , . , - , , -, , , . , .

3.4.6. , [37] , . -, , , . , 10% , 10 , (. 3-55). , , , . , , , , 100 , (. 3-56). , , , ; . , , . , , , . . , .

. 3-56. , - . . . 30 , 10 . . , . 1 , 2 , . . 10 , 10 , , 10 , . , 100 , .

1623.4.7. [38] , . , , , , . , , , . , , , . , , . ; (. 3-57). , , . .

3.4.8. [39] , (. . 2.5). , , - , , . - . . - . ( ). , (-

. 3-57. . . , X , Y .

163

. 3-58. , , . , . 3-57. , . , (. . 3-59). ), , . , , , , . ( ) (. . 3-57). , , , .

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172

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