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. .

TRiC. TRiC .

5- . . : ..., .. ... ..

2008

.............................................................4 ...................................................................................................................................5 ..............................................................................................................7

.......................................................................7

........................................................................................................9 ..............................................................................................10 ..............................................................................................13

TRIC ....................................................................................................................16

...............................................................................................................................16 TRiC ............................................................................................16 ................................................................................................................20 TRiC ................................................................................................22 .....................................................................................................................23 .....................................................................................................................25 ....................................................................................................26

...............................................................................................................................26

.....................................................................................................................................26

TRiC ....................................................................................................................26 GroEL GroES ...................................................27 TRiC .......................................................................................29 ....................................................30 ............................................31 - ..............................................................................31 BrCN- 4 ......................................32 , 4 ................................................33 TRiC ..............34 ..............................................................................34 .........................................................35 ............................................................35

2

..................................................................................................35 ......................................................................................36 () .................................................36 () ............................................36 - ..........................................................................37 .................................................................................39

TRIC ......................................................................................................................39 TRIC ...................44 .................................................46 TRIC ........................48

-

TRIC

..........................................54 GROEL TRIC ......................56

..........................................................................................................64

TRIC

..........................................................................................................................69

.........................................................................................................................72 ....................................................................................................................................75 .......................................................................................................76

3

- D--3- - 1,4--DL- - - TM - () 3- - 3- - ADP ATP GroEL , 14 x 60 (840 ) GroES , 7 x 10 (70 ) GuHCl- HEPES N-2--N'- NAD+ - NADH - PMSF TRiC , 16

4

TRiC (TCP-1 Ring Complex), CCT (Cytosolic Chaperonin containing TCP-1), . , , , TRiC . , TRiC 10% . , , , . TRiC . - GroEL , . , -. , GroEL TRiC, , . , , GroEL,

. , GroEL , TRiC, , TRiC . TRiC . -, , , -, .

5

TRiC . , TRiC , , , . , TRiC . (Nam et. al. 2006; Kitamura et. al, 2006). GroEL -3, (Naletova et. al. 2006). GroEL TRiC

.

6

. , , . , in vitro, (Anfinsen, 1973). () . , . . . , , . , « », , , , (Yon, 1997). in vivo, in vitro, , , . in vitro 7

«»

:

. , , , . 300 /. . ( ) , , , (Minton, 2000; Zimmerman and Minton, 1993). , in vitro , , , , (van den Berg et. al., 1999). 80- . . , , . , . , , . (Heat shock protein/ Hsp) (Schlesinger 1990; Yura and Nakahigashi, 1999; Nover and Scharf, 1997). , «» , . « « » («molecular » 8 chaperones») (Ellis, 1987; Hartl, 1996; Martin and Hartl, 1997).

, , , . , , , (Saibil, 2000). . , . , . . , «» ATP . , . ATP- , - Hsp 70 ( Dna K E.coli), Hsp90 . . , , . : 1 , (Ranson et al., 1998; Fenton and Horwich, 2003); 2 (Kim et al., 1994; Cowan and Lewis, 2002). -, 9

, , . , , . GroEL. GroEL E.coli . (Gething and Sambrook, 1992; Horwich and Willison, 1993). GroEL E.coli, , , .
35

,

GroEL, E.coli, S-, (Houry et. al., 1999). , GroEL 1000 4300, E.coli. , , . , , , GroEL. -, , , . GroEL , . GroEL , . , 10

, , . , . , GroEL GroES ATP. , , 20 , ( 1). , GroEL . (Fenton and Horwich, 1997). , (, ) . , 2% , (Ranson et al., 1997).

11

1. GroEL/GroES . GroEL . : , / . (1), . () , , , . ATP (2) GroES (3) GroEL. , GroES, , . ATP (5) GroEL ( ), . ATP GroES , GroES -. . (Walter, 2002)

12

. , Pyrodictium ( ) , (Phipps et al., 1991). , ATP- GroEL/Hsp60/Cpn60 , , , . , , . 9- Sulfolobus shibatae (TF55 55 ), in vitro, (Trent et al., 1991). TCP-1 ( tailless complex polypeptide 1) (Gupta, 1990). , , , (Ursic, Culbertson, 1991) - . , TCP-1 TRiC. TRiC , . (Frydman et al., 1992; Gao et al., 1992). . 13

( 2): - ; , ; , .

, , (Kim et al. 1994) Swissprot (36 106 ). , , , , . , . . , , , . (TRiC), , (Gutsche et al., 1999). GroEL ( GroES GroEL). - ( 2) Termoplasma acidophilum. GroEL , , ( 2). , GroES, 14

( 2) (Meyer et al. 2003). , , ( ) 50 ( Ditzel et al. 1998).

()

()

()

( )

,

2. . - Termoplasma acidophilum. - () - () (). , ; . a. , . TRiC ; . ( ); ATP- (Meyer et al. 2003).

15

TRiC TRiC . 8 (, , , , , , , ). TRiC ; , (Waldmann et al., 1995) . 8 TRiC 30%. TRiC. - (Kubota et al., 1997). . , TRiC. , TRiC , . : . (Hartl, 1996, Kubota et al., 1995). , , . TRiC TRiC, , GroEL. 16

, TRiC . , , , . , , , . , , GroEL. TRiC - , , , , . (Gao et al., 1992). , . , TRiC (Dunn et al., 2001; Gomez-Puertas et al., 2004) , , -, , , , G- (Feldman et. al., 1999; Srikakulam, Winkelmann, 1999; Gutsche, Essen, 1999; Willison, 1999) TRiC - , - ( 1). , , 10%

TRiC (Thulasiraman et. al., 1999). TRiC - . , 17

100 . , , , , TRiC . - . , - TRiC . , - TRiC - E, . , TRiC (Archibald et al. 2001). GroEL , , GroEL . - . TRiC GroEL/GroES (Leroux and Hartl 2000). , GroEL . - ( ) (Naletova et. al. 2006). , GroEL - TRiC, , , GroEL, , TRiC. , TRiC - ( 1). (Dunn et al. 18

1. TRiC (Spiess et. al., 2004)
()

-, - -, -, 4- G - G1-S- E1 1 - V () EBNA-3 Gag M-PMV 3 SET 3 HOS2 20 (Cdc20p) 15 (Cdh1p) (Cdc55) PP2A (Pex7p) 2 (PRP46) - '- Sec27p - (Ste4p)

42.1, 41.7 50.2, 49.8, 51.2 223.0 60.1 40.0 24.2 47.1 18.5 16.1 42.6 20.9 89.1 73.1 48.8 85.5 51.5 67.4 110.4 59.6 42.3 50.7 99.4 46.6

19

2001). , , , , -, CDC20, , . , , . , TRiC -, BC (VBC- ), Cdc20 Cdh1 . , , . TRiC , . , (Spiess et. al., 2004). TRiC . . , in-vitro, in-vivo, . , TRiC . GimC. ,

20

, TRiC. TRiC. 1-4 . , . , (. 3) (Valpuesta et. al., 2002).

3. TRiC. , . , , . N- , C- (Valpuesta et. al., 2002). , TRiC, . . (). TRiC , . 21

TRiC ( - ). . ( ) N- - , . N- C- . , GroEL, , (Gomez-Puertas et.al., 2004). , , . 4 TRiC. TRiC GroEL , 2% , TRiC . , . TRiC (Frydman et al, 1992) (Gao et al., 1992). TRiC , (Liou et al., 1998).

22

-

()

()

()

()

4. TRiC. () , (U-). ( ). () - , . () . () - . , , , , . () (Spiess et. al., 2004)

, TRiC . TRiC , . 1- 23

(Kunisawa and Shastri, 2003). , TRiC , , (Plath and Rapoport 2000). -, . TRiC . , , (Lee et al. 2003). TRiC , VHL , in vivo. , , , TRiC (Feldman et al. 2003). Caenorhabditis elegans , TRiC - (Nollen et al. 2004). , TRiC (Nam et. al. 2006; Kitamura et. al, 2006). Hsp70 , . (Behrends et. al. 2006). .

24

TRiC,

TRiC. : 1. 2. 3. 4. TRiC. TRiC : -3

. TRiC . -3-

25

, "Fluka"; NAD+, NADH, -, , HEPES, -3-, ATP, , , , , 4, , PMSF, , , MgCl2, "Sigma"; KH2PO4, G-100, "Merck". BrCN KCN Br2 . . TRiC - , . 75 ( 7 ) (: 250 , 100 NaCl, 2 , 1 , 50 HEPES-NaOH, pH 7,4). 150 , 1 PMSF, ( 30 ), . 30 16 000 g, 4- . TRiC 30%, 50%. 30 % . 30 30 16 000 g (40 ). 50% . 10 (: 5 , 0,5 , 10% , 1 , 1 26

PMSF, 250 KCl, 50 HEPES-NaOH, 7,2). 16 . 2 1 4 , 3,5 240 000 g (40 ). , . . ( 50 ) 2,5x10 - . 250-500 KCl . SDS-. , TRiC, . . , TRiC, (0,1 NaCl, 5 MgCl2, 5% , 0,5 , 1 DTT, 20 HEPES-KOH, 7,2). 12 TRiC 0, 68 /. GroEL GroES (Corrales and Fersht, 1996). E. coli W3110 F39, GroEL. 8 30 (100 l, 8,1 (4 ), 0,1 , 10 , 0,2 / PMSF) 22 ( 45 ). 30 5000 g. (NH4)2SO4. 30-80 % 27

30 12000 g, 10 (50 -l, 7,2 (4 ), 0,1 , 2 ) . D. (3 8 ), , 15-25 . 100 . , , 300 NaCl (0-0,5 ) . GroES 0,18 0,23 NaCl. , GroES ( -) . GroES (50 -l, 7,2 (4 ), 0,1 , 2 , 0,2 NaCl), , 60 10 25 . 30 15000 g. 80 10 . GroES 80 . GroES 10 - , pH 7,5 Centriprep 2-7 /. GroES -70 . GroES 12% . 15 , ( GroES 10,5 ) (Marusich et. al., 1998). GroEL 0,31 0,37 NaCl. - 12% . ( 3 ) 28

GroEL D- . - . 2 10 - , 7,5, Centriprep 7,2 /. 10 , 7,5 - 70 . GroEL14 GroES7 , al., 2002). GroEL14 840000 ( 60000 ), GroES7 70000 ( 10000 ). TRiC 0 TRiC : .
280 0,1%

0,18 0,14, (Walters et.

,1%, 280

= 0,4.

. 0
,1%, 280

TRiC .

TRiC , GroEL.

29

5. , GroEL , , , (Asryants et. al., 1985). , ( 5).

6. , , .

30

(Laemmli, 1970) "Mini-Protean 3" (BioRad). 0,5 , ( -I

0,4%- , 6,8 1,5 -I , 0,4%- , 8,8) (0,025 , 0,192 , 6,8) . ( ) , . : 0 125 -I 6,8 4%- 20%- , 0,002% , 10% 1-10 . 0,04 % R-250 2 : 1 : 1 : 6. 10% . 4- 3 . , . , . - - . 31 -. 3 .

. (5,5 - 12%), . . 200 - 150 - ( 7,0). 5 . (45 ), (66 132 ) (443 886 ) 5 /. 40 . (50 , 15 -, 0,02 % G 250, 7,0 40 ), 20 ( , ). , , , , . 50 - ( 7,0 4є). 100 V, 300 V. (0,04% R250 10% 20% ) 10% (Schagger, 1994). BrCN- 4 4 0,1 11 , . 5 CNBr 1 . 40 CNBr 800 8 1 - 11 32

( : 1:1). 6 . 20 , 20 0,1 - 8,3. 1 1 ( 0,1 - , 8.3). 3 , . 50 +40 . 0,1 - (5 , 2 , 8,3) 280 . . 238 / ( 1:1). 14,6 /. , 4 (50 -HCl, 8 , 7,5) 24 . , 280 . . 104 1 ( 1:1). 33

TRiC 50 100 TRiC 10 - 7,5. TRiC 1 : 2. 30 . . 5 10 - 7,5, 50 4- . NAD+ 3-, 340 NAD 6,22 103 -1-1. ( 8,9) 50 , 100 KH2PO4, 5 , 50 , 1 3- 5 3-. NADH , 340 ( NAD 6,22 103 1

NAD+.

3-

, 1 NAD+ 5

-1). ( 7,0) 100 KH2PO4 50

, 1,6 5 NADH. , 0,2 NADH 5 20 3-.

34

25 , NADH 340 3-. ( 1 ) 50 , 2 , 100 KH2PO4, pH 8,9, 1 3-, 1 NAD+ 0,8-2 . , 1 NAD+ . 25 , NADH 340 . ( 1 ) 100 KH2PO4, pH 7,0 1,6 , 0,2 NADH 0,5-1 . , 1 NADH . ( 2-3 /) 4 GuHCl . 100- . : 50 KH2PO4, 7,5, 1 , 5 , 1,5 NAD+ ( NAD+ 1,5 ), 2 ATP, 2 Mg2+. - TRiC. . 1,5 2,5 c 20 .

35

"SIM Aminco DW2000" (). 320 . (50 HEPES, 50 NaCl, 7,4) 450 . 0,8 ( 1 ) . TRiC. TRiC 0,27 . , 450 , TRiC. TRiC 1:3. "ORIGIN 7.0". ()

4 (, , ) 0,47 1 /. 1-2 /. Tmax ( ) , Arina, . .. . () «Zetasizer nanoZS» (Malvern Instruments, Malvern, ). 633 (He-Ne, 4 ) , 36

173°. « » (size distribution by number) « » ( size distribution by intensity). 250. 5 30 . - , Coomassie Brilliant Blue (G-250), : 1 1 , 100 40 % 0 ,1 NH4HCO3 30 37 , . 4 100 . (Promega) 0.05 NH4HCO3 15 -1. 12 37 , 8 0.5 % 10 % . MALDI--. -
1

: 1 0.3 2,5- (Aldrich,10 . - MALDI- - Ultraflex II BRUKER (), (Nd). - ; 37 20 % 0.5 % ),

0 ,007 %. - «». Mascot (www.matrixscience.com). « » NCBI , .

38

TRiC TRiC TRiC . (Gao et. al., 1992; Norcum 1996; Ferreyra, Frydrnan, 2000). , , , (, ). TRiC . : , . ( ). . (. « ») , . 7 ( 1), . 2-5 . . ( 6-8). ,

39

1

2

3

4

5

6

7

8

9

10

11

66 45 36 29 24 14

7. , TRiC (14, 20, 24, 29, 36, 45, 60 ); 1 ; 2 , 30% ; 3 , 30% ; 4 , 50% ; 5 , 50% ; 6 ; 7 ; 8 , 1 ; 9 , ; 10 , ; 11

40

9

13

15

17 21

23 25

29

31

33

35

37

39

43

45 49 53

60 45 36 29 24 20 14

8. , -. (14, 20, 24, 29, 36, 45, 60 ) .

41

, -. 8 . , KCl. 360 470 KCl (. 8, 21 49). , , , () . , . , , , , 470 KCl, 250-500 KCl. 9 1 2 ( 1 2, ). 9 , . 50 , . , , , 50 60 . , (Frydman et al., 1992; Lewis et al., 1992).

42

60 45

9. . 1 , 2 .

43

TRiC .

( www.expasy.org) TRiC. , 0
,1%, 280

0,1%, 280

. ,

TRiC

0,4. (240 300 ), 10. 10 , 260 280 . D280/D2 ATP),
60

0,703. 260 .

, ADP ( ,

. , ( 1,5 ) . . , , GroEL (. 5).

44

0,18 0,16 0,14 0,12 0,1 0,08 0,06 0,04 0,02 0 240 250 260 270 280 290 300

D

,

10. TRiC 240 300 . TRiC 0,68 / ( ) 10 7,5

45

. , , , 2,1 , . , , TRiC . - , TRiC, - . , TRiC , , 8 . 50-60 ( 11). - 1 2, , 50 60 . , . ( 2). 2 , 7 , TRiC. «Tcomplex protein 1 subunit epsilon» . , , ( 59,6 ), . , , TRiC. , 46

2. , . 50-60 . 50-60 T-complex protein 1 subunit beta T-complex protein 1 subunit zeta T-complex protein 1 subunit eta T-complex protein 1 subunit delta T-complex protein 1 subunit theta T-complex protein 1 subunit theta T-complex protein 1 subunit alpha T-complex protein 1 subunit gamma 57,3 57,8 58,1 59,4 59,4 59,4 60,2 60,5 ,

47

50-60 (. 11). TRiC, Hsp60, . 2, Hsp60 . TRiC , TRiC, . , , , . , , 950 , ( 7 ). , 886 , 443 , 66 45 . 12 , , (886 ). , -, , , , , 886 , .

48

gamma, alpha, theta theta, delta, eta, zeta beta

11. , SDS, TRiC. .

49

886 443

66 45 12. -. (886, 443, 66, 45 ) TRiC.

50

, , , TRiC. (), . 13 « » TRiC. 17 . swissprot TRiC 12 16 . , , , , , -, , . , , , 13. , , TRiC , TRiC . , - . , . 14 « » TRiC. , : TRiC , 160 . 10 .

51

30

25

20

, %

15

10

5

0 1 10 100 1000 10000

,

13. TRiC. TRiC 250 10 , 7,5. TRiC 0,27 . , « ».

52

9 8

, %

7 6 5 4 3 2 1 0 -1

1

10

100

1000

10000

,

14. TRiC. TRiC 250 10 , 7,5. TRiC 0,27 . , « ».

53

( 13), , , . 5000 , , . TRiC TRiC 4. « » , . (. « » 6). 238 1 ( 1:1). 104 1 . 45% , , . , , BrCN (30 5 ). , , , , . , , , .

54

1

2

3

60 45 36 29 24 20 14

TRiC

15. 4 TRiC TRiC (0,04 ) 10 KH2PO4, 7,5, , 10 . : 1. TRiC 2. TRiC, 3. TRiC

55

30 TRiC. , - . SDS-. , . , , TRiC . 15. 1, 2 3 TRiC, , , , TRiC. , 3 , TRiC . . GroEL TRiC , TRiC, GroEL. GroEL GroES E. coli, « ». , GroEL GroES. 16 ( 1 2) GroES GroEL, . , .

56

1

2

3

60 50 36

25

15

16. GroEL GroES 1 GroES 2 GroEL 3 ( , )

57

GroEL (. 17). , Mg-ATP 4% ( 2,5-3,5 / ), GroEL/GroES ( 0,1 / ), « », GroEL-GroES Mg-ATP, 16-20% ( 13-20 /). TRiC, , TRiC, GroEL, . 4 30 120 . . , 15-20 4 . 100 . 18 TRiC- Mg-ATP. , 18, , Mg-ATP . , -, , -, , - , , « ». (2 : 3 TRiC

), , , , . 58

10

8

3

, /

6

4

1
2

2
0 0 20 40 60 80 100 120

,

17. GroEL- . (0,13 ) 4 , 100 1 (10 - , 7,5, 1 , 1,5 NAD+, 5 , 2-) 2 ATP 2 Mg2+ (1); 1 GroEL, 1 GroES (2); 2 ATP, 2 Mg2+, 1 GroEL, 1 GroES (3). (. « »).

59

4.0 3.5

+TRiC 1 +TRIC-MgATP

1 3

, .

3.0 2.5 2.0 1.5 1.0 0.5 0.0 0 20 40 60 80 100 120

2

140

160

,

18. TRiC- . (70 ) 4 , 100 0,6 10 - , 7,5, 1 , 1,5 NAD+, 5 , 2) 2 ATP 2 Mg2+ (1); 0,4 TRiC(2); 2 ATP, 2 Mg2+, 0,4 TRiC(3). (. « »).

60

, 18 , TRiC- . , , TRiC , GuHCl, . , GroEL ( (. 19). -

) (Naletova et. al. 2006). , , , , , TRiC , . TRIC Mg-ATP (. 19). , 20, . , . , , . , Mg-ATP. , TRiC 3 . , TRiC- 60 . , TRiC .

61

12 11 10 9

2

, /

8 7 6 5 4 3 2 1 0 0 20 40 60 80 100 120 140 160 180 200

1
,

19. GroEL- , . (18 ) 4 , 100 0,14 (10 - , 150 NaCl, 7,5, 1 , 1,5 , 5 , 2) (1); 0,14 GroEL, 0,14 GroES, 2 ATP, 2 Mg2+ (2). (. « »).

62

3.0 2.8 2.6 2.4 2.2

B E

2

, .

2.0 1.8 1.6 1.4 1.2 1.0 0.8 0.6 0.4 0.2 0.0 0 20 40 60 80 100 120 140 160

1
,

20. TRiC- , . (71,4 ) 4 , 100 0,6 (1) (10 , 150 NaCl, 7,5, 1 , 1,5 , 5 , 2-) (2) 0,4 TRiC 2 ATP, 2 Mg2+. (. « »).

63

, . -, TRiC- -3- , , -, TRiC- , . , . GroEL . 320 . 21 ( 1) 45 . 45 , . GroEL. ( 2). , , GroEL . TRiC . TRiC, , , TRiC. () , ( , ). 64

0.05

0,122

1

0.04

D320

0.03

0.02

0.01

0,122 + 0,122 GroEL
0.00 0 10 20 30 40 50

2
60

, .

21. GroEL . (0,122 ) (50 , 1 , 7,5) 45 . , . ( 1), 0,122 GroEL ( 2).

65

, , . . 35 70 0 1 22 . . 22 ( 2) 520 . . 22 ( 1) 400 . , 400 , TRiC 520 , 450 . TRiC , 1 50 NaCl, 50 HEPES, 7,5 45 . . 23. 23 , 450 ( 1). ( 2) , TRiC . TRiC ( 3) .

66

700 600 500 400 300 200 100 0 35 40 45 50 55 60 65 70

,

2

1

T, C

0

22.

. 0,8 (1) TRiC 0,27 (2) 1 22 (50 HEPES, 50 NaCl, 7,5).

67

0,8 + 0,27 TRiC
0.18 0.16 0.14 0.12

3 2
0,8

D320

0.10 0.08 0.06 0.04 0.02 0.00 0 10 20 30 40 50

0,27 TRiC

1
60

,

23. TRiC . (0,8 ) (50 HEPES, 50 NaCl, pH 7,5) 45 . , . (0,8 ) (1) TRiC (0,27 ) (2) (1,2) (3).

68

, , GroEL, TRiC . TRiC () . , , . , . ( ) ( Ptitsyn, 1995; Privalov, 1979). , , (), (Sturtevant, 1977). , TRiC TRiC. TRiC , . . 24. , 55 ( 1). TRiC 69

300

TRiC 1,8 3 GAPD 27

TRiC 1,8 + GAPD 1,7

250

/(*)

200

2 1

150

100

50

0 30

40

50

60

70

80

90

100

,

24. . 10 - , pH 7,5, 1 /: 27 ( 1), 1,8 TRiC ( 2), 1,8 TRiC 1,7 ( 3).

70

59,4 (. 24, 2). , (, TRiC), . TRiC TRiC. ( 3) ( 2). TRiC - GroEL, 30 , 1,47 (Naletova et. al. 2006). , TRiC .

71

, TRiC ( TRiC TRiC ). , TRiC . , , , , . -, , , TRiC. -, , 886 17 , TRiC. , , TRiC

. , 7 8 , TRiC. «T-complex protein 1 subunit epsilon» , . TRiC -3- TRiC GroE. 72 GroE -3-

. , , , TRiC, . , , TRiC -3-, , , , GuHCl. ("misfolded forms") (Polyakova et al, 2005; Naletova et al, 2006) . , , , TRiC , GroE. , , , . TRiC ("misfolded forms") . TRiC . , TRiC, , . , , 73

GroE (, ), TRiC, , GroE (, ) (Naletova et al, 2006), TRiC- . , , .

74

1.

TRiC .

, , 886 17 , , 7 , TRiC. 2. 3. , TRiC GuHCl , -3- . GroEL TRiC 4.

. , TRiC GroEL .

75

1. 2.

Anfinsen, C.B. (1973) Principles that govern the folding of protein Archibald J.M., Blouin C., Doolittle W.F. (2001) Gene duplication

chains. Science. 181:223-230 and the evolution of group II chaperonins: implications for structure and function. J. Struct. Biol. 135, 157169 3. Behrends C., Langer C.A., Boteva R., BЖttcher U.M., Stemp M.J., Schaffar G., Rao B.V., Giese A., Kretzschmar H., Siegers K., Hartl F.U. (2006) Chaperonin TRiC promotes the assembly of polyQ expansion proteins into nontoxic oligomers. Mol. cell, 23(6):887-97 4. 5. 6. 7. 8. Corrales, F.J., Fersht, A.R. (1996) Kinetic significance of GroEL14. Dunn A.Y., Melville M.W. and Frydman J. (2001) Cellular substrates Ellis J. (1987) Proteins as molecular chaperones. Nature 328:378-379 Ewbank J.J., Creighton T.E., Hayer-Hartl M.K., Ulrich Hartl F. (1995) Feldman D.E., Thulasiraman V., Ferreyra R.G., Frydman J. (1999) (GroES7)2 complexes in molecular chaperone activity. Fold. Des. 1:265-273 of the eukaryotic chaperonin TRiC/CCT. J. Struct. Biol.135, 176184

What is the molten globule? Nat Struct Biol. 2:10-11 Formation of the VHL-elongin BC tumor suppressor complex is mediated by the chaperonin TRiC. Mol Cell, 4:1051-1061 9. Feldman D.E., Spiess C., Howard D.E., Frydman J. (2003) Tumorigenic mutations in VHL disrupt folding in vivo by interfering with chaperonin binding. Mol. Cell 12, 12131224 10. 11. Fenton W.A. and Horwich A.L. (2003) Chaperonin-mediated protein Ferreyra R.G., Frydman J. (2000) Purification of the cytosolic folding: fate of substrate polypeptide. Q. Rev. Biophys. 36, 229256 chaperonin TRiC from bovine testis. Methods Mol Biol. 140:153-60 76

12.

Frydman J., Nimmesgern E., Erdjument-Bromage H., Wall J. S.,

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