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5 5 : (ubiquitin); 1. ( nucleosome)

1

D . , DA . , ( , ) . D . [1] . D D.

8-7 [2] , .( 8-7) , D . , D . D.[2] ,

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, Roger Kornberg 1974. Aaron Klug . . , . (3)2(4)2 2-2. D. , , . . , , D .[3] D , . , D. DA, . 1 , DA. , , . D D . D 200 bp 680 . D 100 . , D . , , , 104 . D. : 360 110 .

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D. D . D , . D .[3] ( ) 30 nm.( 8-8) , ' . [2]

8-8. [2] D "" D . . D . , D "" , D (linker DNA), . ( 2. 2. 3 4) D 146 . "" D.[2] , . . , 100. "" ' "" D.

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(FIGURE 10-20) D , D. ( ). D, . D, 200 ( 146 D D ). ( 60 ) D. [4]

( ) [5] ( ) : , 4 . , , . . , " ". , , 30 nm. 30 nm ( 2) ' , 1

5 .[4].

5

2. 1 . 11 nm ( 2) 1 , [6] D . , R D ; D;[7] , . , D . , R . ' ,

5 R D.[7]

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D . . , . , . , .[7] D . , , . D , D, . . . . D . , ( , ) "" . D , R . "" D - .[7] - : . , - . - , DNA .

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, ( HMG), . , HMG14/17 -

. HMGI/Y , , DNA, 1, - . HMG DNA. , DNA (q.v. SRY, enhanceosome ) -, , DNA . DNA DNA .[8] DNA . . DNA - (). , (DACs) . , K9 H3, (MTs) , .. K4 3 . K9 , .. 1, DNA (DNMTs), . DNA MBD. DNMTs HDACs MTs, . .[9]

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- DNA. H3 H4 (Figure 8.7) H3 Lys4 (K4). DNA , . . () . ' , -. , S . . , .[10] - .

. . D 250 ms ( 3) 10-15 ms. D

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9

. D.[11]

3. .

. SWI/SNF ISW ( 4) ( ).[11]

4.

5 - .

10

:

D

(SW).

.

(SWI/SNF).

D.[11] .

. , . : hsp70 (drosophila), , . .[11] : 1) D , 2) - . D .[11] .

. .

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11

. . ( 1) . . .[11]

1. .

:

D .

.

.[11] .

R pol.

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R ( 5) D 50 , 12 bp. D D . [11]

5. R D , D . D .( 6) R 30 bp D . 10 bp D. . ( D).[11]

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6.

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[1] [2]

Berg J., Tymoczko J., Stryer L., , , , 2005, . 984. Alberts, Bray, Johnson, Lewis, Raff, Roberts, Walter, , , .. , .291-292.

[3] [4]

Berg J., Tymoczko J., Stryer L., , , , 2005, . 986. Alberts, Bray, Johnson, Lewis, Raff, Roberts, Walter, , , .. , .293-294.

[5] [6]

Lodish H., Berk A., Matsudaira P.,..,Molecular Cell Biology, fifth ed.,. 425 ., , : : ,...2005, .10

[7]

Alberts, Bray, Johnson, Lewis, Raff, Roberts, Walter, , , .. , .311-312.

[8] [9].167.

Twyman R.M., Advanced Molecular Biology, bios scientific publishes, .37 Wolfgang Arthur Schulz, Molecular Biology of Human Cancers, Spinger 2005, Wolfgang Arthur Schulz, Molecular Biology of Human Cancers, Spinger 2005, .180. ., , 14/7/06, http://multimedia.biol.uoa.gr/media2005_06/karamanos_files/lobby0.htm

[10] [11]

5 2. (Ubiquitin)

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. (Ubiquitination) ( Ubiquitylation) - ( ) . 1975 8,5 kDa . , '80 Aaron Ciechanover, Avram Hershko Irwin Rose 2004. , . (polyubiquitination), , . ( , )

.[1] , , ( ) . . MPF . , .(Figure 1244) [2]

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[3] ( 1) 76

8500 DA. : 96% ( 3 76 ). : MQIFVKTLTGKTITLEVEPSDTIENVKAKIQDKEGIPPDQQRLIFAGKQLEDGRTLSDY NIQKESTLHLVLRLRGG [1]

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1.[4] , 1, ,( 2) 2, , 3.

2.[5] , 1 . . (C-

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1. , 2. , 3 2 - -, . , : - 48 . .[5] . , . , . 1, 2 3. , 2 3. 3 , 2-3 .[5] ; . . - . 20 , 2 . , , , , , . 3 - . , , , , ( S). [6] . ,

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(V) 3. p53 (Fig.2.11) D, . 3 90% . , , .[6]

[7] " - "

, , . :

-

de novo

. -

. mRNA . :

. . . , . (tab.2.1)

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. , . . , , . . --, . .[8]

[9] , ; 26S . , . 26S : 20S, , 19S. 20S ( 3) 14 700 kd .[6]

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3.[6] 14 . 7 . . . . . , . , .( 4) 20S . 19S, 700 kd 20 . 20S 26S.( 5,6)

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5.[10]

Figure

8.

Binding and to its short

of

the

polyubiquitinated substrate to the 26S proteasome degradation subsequent peptides.

Illustration was kindly provided by Dr. Ed Yeh, M. D. Anderson Cancer Center, Houston, TX. ( 26S .) 4.[11] 19S . 19S ( ) - 230 , , . . , . , A 19S 20S,

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. , 19S ;. . .[10]

6.[12]

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.

[10] . . , , . F- (F: ) , . . -. F- , - (: ). , - , 3. 3 - F-. (Figure 43-10)[10]

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[13] -. F- - 1- : , , .[10] . . . , . 28 .( 7)

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7.[14] , , . , . . , , . , , , . . .[14]

8.[14]

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( 8) . - hiS, . hiS hiF. 1 160 28% hiF; .coli. hiS, ( 9) , 14% .

9.[14] , .[14]

, . ( ) , . : , , Alzheimer arkinson ( ), ( , , Liddle ( , Na+ , ) . . ,

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28

, .[15]

.

Von Hippel-Lindau

Angelman, UBE3,

(E3) 6-AP. (E3) VHL VHL . Liddle Na+ (ENaC) . [16]

Figure2 . E1 E2 .(Figure2) , E2 E3, 2 3. E3 , . E3, . [17]