(), 90- .
: 5/; /; 7/; 8/D; 9/; 10/F; 11/G; 12/; 26/1; 44/J; 45/K; 48/L; 51, 60 61, - 5/, 9/ 45/ - . , , , 1991. , , ; ; . , , (7/), (48/ L) , (/). , - , -, .
. . () 5 : 6/, 7/, 9/(1), 10/F 48/L. 6/ 7/, llulomonas fimi - 6/, 10/F 48/L, - 9/(1) 48/L.
. (GH) : GH-A ( 1, 2, 5, 10, 17, 26, 30, 35, 39, 42, 51, 53), GH-B ( 7 16), GH-C ( 11 12), GH-D ( 27 36) GH-E ( 33 34).
, (- (β/α)8-) " ", β-, (α,α)6- .
- , -, . II Trichoderma reesei - , α- β- -. {(β/α)8}-, 1, 2, 3, 5/, 6/, 10/F, 17, 26/I, 30, 35, 39, 42, 51 53.
lC - ( 8/D) (α,α)6- 365 . D (α,α)6- - α-, α-, . - , α-, . , , 450 . D N- 90 , β-. . " " , . 9/, , 4 , . (α,α)6- , . 9/2.
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(α,α)6- 8/D, 9/ 48/L, .
I β- , , , β-, α- . I 434 405060 Å.
11/G 12/ " ". 7/ β- . ( 16). 7, 11, 12 16 12-14- β-.
"" 6-β- 45/, , () .
() , .
. - II, III, IY, YI, YII, IX. 11/G, 12/H 45/. II, III, IY 45/ β- " " (.6).
6
( Tomme et al., 1995)
e | * | ** | C | -*** | |||
I | Agaricus bisporus | Cel1 | C | -, β- | M86356 | ||
Fusarium oxysporum | Xyn | N | 10/F | L29380 | |||
Neocallimastix patriciarum | XylB | C | 10/F | S71569 | |||
Humicola insolens | EGY | C | 45/K | [Schou et al., 1993] | |||
Neurospora crassa | CBHI CBHI CBHI | C | 7/C | X7778 | |||
Penicillium janthinellum | X59054 | ||||||
Phanerochaete chrysosporium | M22220 | ||||||
Porphyra purpurea | PBP | X4 | U08843 | ||||
Trichoderma reesei | CBHI CBHII EGI EGII EGY Man | C N C N C C | 7/C 6/B 7/C 5/A5 45/K 5/A1 | P00725 M16190 M15665 M19373 Z33381 L25310 | |||
IIa | Butirivibrio fibrisolvens | End1 | C | - β-, β- | 5/A4 | X17538 | |
Cellulomonas fimi | CenA CenB CenD CbhA CbhB Cex | N C C C C C | 6/B 9/E2 5/A1 6/B 48/L 10/F | M15823 M64644 L02544 L25809 L29042 L11080 | |||
Clostridium cellulovorans | EngD | C | 5/A4 | M37434 | |||
Dictiostelium discoideum | SGSP | C,I | 98/106 | ( - ) | M33861 | ||
Microbispora bispora | CelA | C | 6/B | P26414 | |||
Pseudomonas fluorescens | CelA CelB CelC CelE XynA | C N N C N | 9/E1 45/K 5/A1 5/A2 10/F | X12570 X52615 X61299 X86798 X15429 | |||
Streptomyces lividans | CelA CelB ChiC | N C N | 5/A2 12/H 18 () | M82807 U04629 D12647 | |||
Thermomonspora fusca | E2 E3 E5 | C N N | 6/C 6/C 5/A2 | M73321 U18978 L01577 | |||
IIb | Cellulomonas fimi | XynD | X2C | 11/G | X76729 | ||
Streptomyces lividans | Axe | C | - | [Tomme et al., 1995,] | |||
Thermomonospora fusca | XynA | C | b- | 11/G | U01242 | ||
IIIa | Bacillus lautus | CelA | C | - β-, 2+ | 44/J | M76588 | |
Bacillus subtilis | Cel | C | 5/A2 | X67044 | |||
Caldicellulosiruptor (Caldocellum) saccharolyticum | CelA CelB CelC ManA | x2I I x2I x2I | 9/E2//48/L 10/F//5/A1 9/E2//5/A4 5/A4//44/J | L32742 X13602 [] L01257 | |||
Clostridium cellulovorans | CbpA | N | M73817 | ||||
Clostridium stercorarium | CelZ | C | 9/E2 | X55299 | |||
Clostridium thermocellum | Cbh3 CipA CelI | C I C | 9/E1 9/E2 | X80993 L08665 L04735 | |||
Erwinia carotovora | CelY | C | 9/E2 | X76000 | |||
IIIb | Cellulomonas fimi | CenB | I | 9/E2 | M64644 X55299 L04735 | ||
Clostridium stercorarium | CelZ | ||||||
Clostridium thermocellum | CelI | ||||||
IY | Cellulomonas fimi | CenC | x2N | " " β- (.. 5 6 ) | 9/E1 | X57858 | |
Clostridium cellulolyticum | CelE | N | M87018 | ||||
Streptomyces reticuli | Cel1 | L04735 | |||||
Thermomonospora fusca | E1 | L20094 | |||||
Myxococcus xantus | CelA | 6/B | X76726 | ||||
Y | Erwinia chrysanthemi | EgZ | C | , b- b-, | 5/A2 | Y00540 | |
YI | Bacillus polymyxa | XynD | C | X57094 | |||
Clostridium stercorarium | XynA | X2C | 87/92 | 11/G | D13325 | ||
Clostridium thermocellum | XynZ | I | 10/F | M22624 | |||
- | G-α | X2C | 16 ( ) | D16623 | |||
Microbispora bispora | BglA | C | L06134 | ||||
YII | Clostridium thermocellum | CelE | I | 5/A4 | M22759 | ||
YIII | Dictiostelium discoideum | CelA | N | 9/E2 | M33861 | ||
IX | Clostridium thermocellum | XynX | X2C | 174/189 | 10/F | M67438 | |
Thermoanaerobacterium sp. | XynA | 174/187 | M97882 | ||||
Thermotoga maritima | XynA | 170/180 | Z46264 | ||||
X | Cellvibrio mixtus | XynA | C | b- 2- 3- , a- . | 11/G | Z48925 | |
Pseudomonas fluorescens | CelA CelB CelC CelE XynA XynE | I | 9/A1 45/K 5/A1 5/A2 10/F 11/G | X12570 X52615 X61299 X86798 X15428 Z48927 | |||
C |
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* , N- (I) ; 4 2 ; ** ; *** SWISS-PROT, EMBL/GenBank, PIR
, , , . , tyr, trp, , . . -, N- , , , . , .
. I II , . , , , .
IIb . , - , . , . , II .
131 172 III, . IIIa , , 5/1,2,4; 9/1,2; 10/F; 44/J; 48/L. - , Caldicellulosiruptor (Caldocellum) saccharolyticum, , . N- . IIIb , 9/2, IIa .
IY , . N- , , 9/1, 6/ 10/F. , .
Z - E. hrysantemi Y. - , 5/2 63 . , , , , .
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YI β- 10/F 11/G, 1,3-β- G-α -. -, .
YII YIII . : ( 5/4 ) , - 240 , ( 9/2), III IY (152 ) N-. IX - , 10/F. 170-189 . 51-55 , - , 5/1,2; 9/1, 10/F, 11/G 45/, . , XI XII 1,4-β-: , , . , , , . Trp Tyr, , . , , , . , , , , . , .
, , . , , - , , , , .. , . , , ( , , , ..). , .
( ) (), . , , . , .
, , , , , . , , , (1 - , 2 - ..). .
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, , , . , , , , , , , - . , .
, . / , . (.30).
, , , . , I II Trichoderma reesei . , .
. 30. : - , ; - , ; - , .
I, 7, 50 10 - -7 +3. , -7, . , , 180, , . -7 +2, +1 +2 -7 -1, E212, .
II ( ΙΙ). , Ι. 4 (-2, -1, +1, +2) . , +4 . -2, +1, +2 +4 Trp135 (-2), Trp269 (+1), Trp367 (+2) Trp272 (+4). N- 1.
I T. reesei 45% I. , , , .
, , , , - . - , ( ).
, β-1.4- , .
, β-, . , , -1 . , , . , -1 , . β- , (.31).
. 31. , . - , (, - ); - , -1 ( ); R - ,
"" "". , "". , . , .
, β -, . -1 , , , . , , (. 32).
.32. ,
, II Trichoderma reesei , -1 +1 , , 175 221. -, Asp221, Tyr169, -1, .
. , , , ( 60- ).
. , . . . . , (.33).
. , . .
, Trichoderma reesei , , 2-8% . Aspergillus niger .
/
Gn Gn/x G2 G
/
.33.
Gn - ; Gn/x- c , ; G2 - ; G
, - .
, . , , II I EGII/CBHI 5/95. 40/60, - .
: , , .
, I ( Ι) Trichoderma reesei , 44 , 18 (.34). Sporotrichum pulverulentum ellulomonas fimi.
( Ι) Trichoderma reesei β- , , SS- . ( Ι) - . 38 , β- , SS-. , . .
, , . , " - ".
. 34. ( I Trichoderma reesei): () - , . , ; () - ; () - ; CCD - ; LR - () ; BD -
"" "". , , .
, , , . , . : .
: 1- , ; - ; - ; , ; . , .
60- . , . , . . , , , , . , , , . . " - "
" - " , . , , , .
, , "" , - - , "" - , , , . , , .
, , 15 , .
, , , . I Trichoderma reesei , II , .
( , , ..) . , , - , , .
(2000), , Ι. , . , , , . , , . : , . , , , : (*** ) → (** *)→ (* **) → (Ė ***) → (** *) .. .
. .
, , . , I -7- -5 . , -5 - -2 . . , , . 1, +1 +2, .. - , . . . , - , .
, , , . - , , .
, Trichoderma, Penicillium, Fusarium, Humicola, Phanerochaete, Schizophillum . , . T. reesei (longibrachiatum) H. insolens (I II, 6/ 7/ , ) : I (7/); II (5/A5); III (12/), IY (61; T. reesei), Y (45/K) YI (6/; H. insolens). , 10/F 11/G 5/1. cbh1, cbh2, egl1, egl2, egl3, egl4, egl5, xyn1, xyn2, man1, T. reesei, bgl1 bxl1 (β-- ), axe1 (), glr1 (β-), arf1 (α-L-), agl1, agl2, agl3 (α-), .
I, . . I, . , -, 180 , . I , : , - .
II 20 Å. . .
I Trichoderma reesei . I , 4-. - I (T. reesei) (H. insolens).
II - - . . - (-). N- I.
. IY T. reesei, 61, - , . Y T. reesei H. insolens, , 6β- ( 45/), I -, 36 , 23 . ( 4 +2) .
H. insolens YI, II 6/. . II , .