Epigenetic Regulation on Gene Expression Induced

Embed Size (px)

Citation preview

  • 7/24/2019 Epigenetic Regulation on Gene Expression Induced

    1/15

    See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/237004244

    Epigenetic regulation on gene expression inducedby physical exercise

    ARTICLE in JOURNAL OF MUSCULOSKELETAL & NEURONAL INTERACTIONS JUNE 2013

    Impact Factor: 1.74 Source: PubMed

    CITATIONS

    23

    READS

    1,254

    4 AUTHORS:

    Ioannis Ntanasis-Stathopoulos

    National and Kapodistrian University of Athens

    5PUBLICATIONS 25CITATIONS

    SEE PROFILE

    Ioannis - Georgios Tzanninis

    National and Kapodistrian University of Athens

    4PUBLICATIONS 23CITATIONS

    SEE PROFILE

    Anastassios Philippou

    National and Kapodistrian University of Athens

    47PUBLICATIONS 664CITATIONS

    SEE PROFILE

    Michael Koutsilieris

    National and Kapodistrian University of Athens

    272PUBLICATIONS 4,892CITATIONS

    SEE PROFILE

    Available from: Anastassios Philippou

    Retrieved on: 18 February 2016

    https://www.researchgate.net/profile/Anastassios_Philippou?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_4https://www.researchgate.net/profile/Michael_Koutsilieris?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_4https://www.researchgate.net/profile/Ioannis_Ntanasis-Stathopoulos?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_4https://www.researchgate.net/profile/Ioannis_Ntanasis-Stathopoulos?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_5https://www.researchgate.net/profile/Ioannis_Georgios_Tzanninis?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_5https://www.researchgate.net/?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_1https://www.researchgate.net/profile/Michael_Koutsilieris?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_7https://www.researchgate.net/institution/National_and_Kapodistrian_University_of_Athens?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_6https://www.researchgate.net/profile/Michael_Koutsilieris?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_5https://www.researchgate.net/profile/Michael_Koutsilieris?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_4https://www.researchgate.net/profile/Anastassios_Philippou?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_7https://www.researchgate.net/institution/National_and_Kapodistrian_University_of_Athens?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_6https://www.researchgate.net/profile/Anastassios_Philippou?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_5https://www.researchgate.net/profile/Anastassios_Philippou?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_4https://www.researchgate.net/profile/Ioannis_Georgios_Tzanninis?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_7https://www.researchgate.net/institution/National_and_Kapodistrian_University_of_Athens?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_6https://www.researchgate.net/profile/Ioannis_Georgios_Tzanninis?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_5https://www.researchgate.net/profile/Ioannis_Georgios_Tzanninis?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_4https://www.researchgate.net/profile/Ioannis_Ntanasis-Stathopoulos?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_7https://www.researchgate.net/institution/National_and_Kapodistrian_University_of_Athens?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_6https://www.researchgate.net/profile/Ioannis_Ntanasis-Stathopoulos?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_5https://www.researchgate.net/profile/Ioannis_Ntanasis-Stathopoulos?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_4https://www.researchgate.net/?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_1https://www.researchgate.net/publication/237004244_Epigenetic_regulation_on_gene_expression_induced_by_physical_exercise?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_3https://www.researchgate.net/publication/237004244_Epigenetic_regulation_on_gene_expression_induced_by_physical_exercise?enrichId=rgreq-8bfc217c-83fa-4ee2-a43f-bdfbfdce3463&enrichSource=Y292ZXJQYWdlOzIzNzAwNDI0NDtBUzo5NzI3MjE1NTA4Mjc2MkAxNDAwMjAyODk3MzQw&el=1_x_2
  • 7/24/2019 Epigenetic Regulation on Gene Expression Induced

    2/15

    133

    I(+)c%)(

    @ @-

    @ @ J @ @ @ @ @ J@@ J>. -

    J, >J @ @ -

    @ J I,

    I@ @ @ 1. ,

    @ DA @ @>R

    @ @ J

    @ >J (F 1). >@ @>>@ -

    J I @ >@@,

    >J@ J@, DA >J@,

    I@ @ J @ >@A (>A)2.

    H@ >@@ @-@ @ @

    J- @ @ @, J @ H3 H4 -

    @. H@ J (HA) @ J

    (HDAC) KJ> DA J@, HA J @ HDAC >@ > @>DA. , @ J J@ @ @ @ @3,4, HA HDAC > @ @ J@ , J, @ -@4,5. A@ -@ >J@, @, J (H$), KJ> >J @ @ J @@ @. KJ> J @, J > 4 ('AD4), J- >J- 1 (#*D1) !>@ C-@>-@ @>J (!HD$), >@ >J @6,7.

    DA >J@ @ @ JK J >J @ DA >J(D$). KJ> >J @, @ @- >@@, >J @ CG @. CG -@ J @ , CG ,@ >@ @ DA >J@ @ CG @, @ @ CG 8. -J , @ @@ @() @ @ > @ >J-CG @> ($BD) @, - HDAC, @ @> @ 6,9, @.

    A@ >> @ @ > J>A. $A @ @ > @@ A >@-

    ! $@ @ EI> 'J@@J, $ *@@, @ "@ J @ A, A, G

    Ab+ac

    J I >@ @ @ >J J@@ J>, >@, J @ @ @ > @ /@ >@ @ I. $J @ @ @ @ @ DA, @ >@@, DA >J-

    @, I@ @ >@A @ @> . A @ >J -, J >@ @ @ @@@ @ @J@@ @, >@ J@>,@ . H @@ J J I @J@@ J> @@, @ >@.

    K!)+: AJ@, DA $J@, G *, H@ $@@, >@A

    R!/%! A+%c&! HJ@@>

    @ @ @ @ .

    C(**e+)('d#'! a"(*: M#c"ae% K(+#%#e*#+, MD, P".D, De)a*&e' ( E/)e*-

    #&e'a% P"+#(%(!, Med#ca% Sc"((%, Na#('a% a'd Ka)(d#+*#a' U'#e*+#

    ( A"e'+, 75 M#c*a+ A+#a+, G(d#, A"e'+, 115 27, G*eece

    E-&a#%: &$(+#%@&ed.(a.!*

    Ed#ed b: S. Wa*de'

    Acce)ed 26 Feb*a* 2013

  • 7/24/2019 Epigenetic Regulation on Gene Expression Induced

    3/15

    !. -*@@@ .: E @ I

    134

    , @ 22 @ , J J @

    @ > @ @ @, @

    @10,11. $A @ J@12 @> >@13. @ @

    >A >J > @ ,

    @>> @ @. J, >A -

    @ @ >> @11,14,15

    @ >> @10, >-

    @ @ @ @ >J@> >@16,17.

    E @ J OP @ O@@

    P @ J I @ > @ @

    @@ @18,

    @J@@ >>, @ -

    >@, @@ @ @

    . F, @ @

    @ @ >@ >>@J @; ,

    >> @ J I @ >-

    @ @@ ;

    #J, I @ @ @ -

    @ @ @ @ J> @ J> .

    E*%#!(!%c +!#&a%)( )" '!ab)&%c *+)c!!

    %(c! b !!+c%!

    J I -

    @ I@ @ > ,

    >> @ @ @ @J @ > @

    @ @ >@ @ I. $J @ @

    I@ @ @ @

    J I @ >@@19-21.

    , I J@>J@ @

    @ @> @ J @.A@ J@>J@ >J @ @>@@ >@ (.., 'GC-1, FA$, ''A-V, 'D"4, - J) I@, @- @ @ >- @ @,

    $J@D1 >J@J- @ ($EF)2A, @ @ @ @ > >@ >@1. $@-@, @>@ >J@ @ @ @- @ J @ I; J I @ @>@ >J-@ @ @I@> @@- @>> (''A-U), @@ 1 ('GC-1), @@ A >@@ (FA$), J J@@> 4 ('D"4) $EF2A, >>J

    I, @ @>@R >J-@ @ @I@> @@- @ (''A-V), 3 @ I. *> @@ e/ #( >@1,8.

    @, I @ @ @ J-@@ > C2+ A$', @ > >, @ @ J @ @ @. -@ @ @ @ @ C2+/C>@@-- @

  • 7/24/2019 Epigenetic Regulation on Gene Expression Induced

    4/15

    !. -*@@@ .: E @ I

    135

    'GC-1S @-@, I@

    @ @ @ @ , , @ -

    @ 23. $@@, @-I@ @ 'GC-1S @J @ 1 (F-1) @

    I@ @ G#4 J @ $EF2 >24,25

    >J A$'" @ I@ @ G#4

    @ @ 'GC-1S J.

    , I@ @ G#4 @ -

    @ @ J (F 2). *J

    > , HDAC, @ @

    HDAC4,5,7 9, J I5 J -

    @> J4, @, J @-

    >@R @, J I. @ @

    @ -> @@> @26,27,

    @ @@J@ J C$"28,29, A$'"30, @ @-

    > @ J@ @ G#4 -

    @ $EF2 @>@ @ 38,39.

    @ @ $EF2 I @

    @ @ I37. $@@, HDAC

    I@ @ 'GC-1S, I

    J- >40,

    > @ @ I41.

    * I- >@@ @

    >@. *J, J 2 ,

    ''A-U 'GC-1S J>J > . J>J@ @ -

    >A I@ @ 'GC-1S >@@DA42. , I >J @ -

    @ @@@ @ J 2 @ >-

    @ @43,44 @ @->@

    >>, @ @J I@ @

    G#4 >, @ J@>J@ @ ''A-U

    'GC-1S.

    E @ @ @ @

    >J@ J ($HC)45. , J@

    >J@ @ @ H3 @

    I@ @ $HC, I $HC $HC

    F%#+! 2. H@ >@@ @ @ J 4 (G#4) I@ @ @ I. A$'": 5R A$'-@

  • 7/24/2019 Epigenetic Regulation on Gene Expression Induced

    5/15

    !. -*@@@ .: E @ I

    136

    >@ @ > @@ > J(> @), @ @>46,47. $@@, HDAC5 @ @

    > @ J @I @@ I >26.A@, @ @ J > >I> @ J (2 >I) >

    @J @ I@ @ J- $*4 (>@@J K @-- @)48, HA I@ @-@> @ @ (2)49, -

    @ @@ @ @ @>@50.$@@, @ @J >>

    @ J@@J >A.A J >@, >A

    >@ J @ @ @ @. 'J > , >A (>J@>A) @ @ >J@J @-

    @ @, >@ @ > J, @ > J@J @J,

    @ J > J@51. @ @ >J@>A @@ J @

    @ @, J@ @J -@ ($F), $J@D1 >J@, $EF2,> @ @ (*F) >J@- -

    @ @-A ($F-A)52. A @> >J@>A, @ QQRR >A (->A) >, > >J J@@ @ @,>>@, @J -

    > @. *@> @ >A J I @ I (>-21 >-221), @J @ I (>-20), @ J

    @ J @ I (>-146 >-222), @ >

    (>-133, >-210, >-328) J @ I-, @ (>-133) @ I- > @@ @ I53,54.

    A@ I @ @ >J -@ I@ @ @ J @ >A >, 22% @ -

    @ 16% @ > >-@>, J @I @@J@55. , >A I@ I-@ @ >@@ @I@ KJ>, -

    @ >@ @ >A @ >@ .A@, @ >A @ J (-@) I 1, 'A3 *9,

    >J >@@ @ > @ J @ I55. @, >A >J J @ @ @ > K - I @ @ >@

    > @ @, @ @ -. , @ @> I@ @ J @ >A

    @ J@ @ >56. F->@, I @ @ I-@ @ @ J @ >A >, J @ > @

    >. *J, I- -@ >A I@ >@ I@ @ -J @ >J@>A60.

    E!+c%!-%(c! !*%#!(!%c +!#&a%)( )"

    %("&a''a)+ *+)c!!

    >-@J @61-63. A@@-@ (A*C) >@@ J@@-J >>@J J64,65. - @-166 @>@ @ @ -J @ @ @ >>J. @ @ A*C J >>. *J, >J@ @ CG @ I@ 1 J @ A*C @ I@69,70. @ @>@ I - >J@ @ A*C, J @ > >@@-J 71 , , @ @ >>-@J J@ >->@J @>, @ @ @ @ @> @ - , @@ @.

    @, @ I -J I@ @ >A @@ ('B$C)73,74. A @ @-, *@> .73 JK @- >A @ I@ J J -@ I, J @-@ >A I@ . >@ @> -> J, -> >>@J @75. @ @ @ @@76,77. @ H@ J, @ @ @ @ >>@78. >.

    @ 'B$C, J @ J @ 'B$C- >@A @ I-@ J J I, J 74. *@> @ @ J J >@ @ @ @ @- ->>@J

  • 7/24/2019 Epigenetic Regulation on Gene Expression Induced

    6/15

    !. -*@@@ .: E @ I

    137

    J@@J92,93. A J@>J@ > >J @ @ -@> J @ @> >J@ @ 94. 'J J J @ @ @ @> DA >J@ , , @@, @ @> I, J@>J @> 95.

    A@ J >> @ @ @ >>@ > J A*C @71.A @>@, J I I@ @ A*C @ >>, ,

    , @ @ >>@J J@ >>@J @> @ @>@ @.

    @ @J J@>J@ @ J>J@ @ @> >@ @>.AJ, >@ J @ > @>, >J@ @J@ CG @ @>@ @ @ @ 96,97. A - DA >J@ >J @ >@--@ , @>@ @ >>@ @98. CACA2D3 > >@ @ , @ @@ @ 98. .98 @ CACA2D3 >J@ >@ @ J J @> @ @ @> J >J@ @ @- >@. #3$B#1 @ >@@ >J@ @ @ J @- I@ >@99. .99 @ #3$B#1 >J@ I >@-I I@@ , @, @J @-@ >, @ @>@ @ @-

    >@, @ < @ . @ @ > - A'C A**F1A 100. @ >@ >@ J>J@ @ >@ -@ @-> , @ I-@. F>@, J I @, @@ @ @>@ J->J@ @ >@ @ > @100,101. A@ J I @ I@ @ >@ -@ @ 53, @- >J J @ -, @ >> >A18. @@, I >J @@ @ @- >@ @ @ @ @ >@ @ @ >>@J @.

    E!+c%!-+!#&a! !*%#!(!%c '!c$a(%' %(

    a#%(# *+)c!

    A @ J @ ->@ @@ @>@ @. > @ @@ @> @ - . @ .102 .103 @ >J@ , @> J, @ @. @ @ >@A @ @, @ >@ @ >J @@@104. F>@, J @ @ @ @> @ >-105. @> @ @ @@>@@> @ J @ @106. @ @> @ @ I @-@ A >J @> @> 107, - >@@ @>R. , @ > >@108-110 >109, J I @@> J @, @ @ @ K @>, @ >>. F-, @ @> J I >J@ @ DA, @ >@-@ @@ @ >A2. A @ >@@ @, @@> I, @ @ @.

    A@ >J @ >@ @ 111.* @ J @ >J @ @ @ @ @ J@-

  • 7/24/2019 Epigenetic Regulation on Gene Expression Induced

    7/15

    !. -*@@@ .: E @ I

    138

    > >@@113. @ *1 J @J >> >A >-134, @ J J >>@J @>@ >114-116. , 106, J I @*1 J, @ J @ I @

    @ > @ > @J *1 J >. @ J I , @-J @ >>, *1 J , , >@ >@, 'GC-1, 53, F-WB ( @ @ J @>, @ , @J, . , @@ I @ @> -@J@@ @>@ , @ @ J@106.

    @, - >@ 117, @@118, J -

    119

    @ @ >>@.C@ I >J I@ @ @->>@J J@@@ ,@, @ >>@J 72. #J,@ J @ J, @ @ >, @, >> - J I I@ @ >J@ -@J @, $J@, $J@D, $J5 $F4, @ > @J @ >- @18.

    E!+c%!-%(c! !*%#!(!%c a&!+a%)( %(

    c!(+a& (!+/) !'

    @ J - @J >@ @ @ I @ J @@. @ @ I >J > @ @ @ - -@@ @ (BDF), @@ J I- @> @ @ @@>120. , @ BDF @ @ @J @ I @ -J121-123, @ @ @ IJ, J @ @ >>@J124-127.$@@, >@ >@> J, J @ >@, A$'", @ @ - JJ I 128.

    A>@ BDF @>@, @>@ @ @ @ @ J, >>@J @6. $J-CG- @($C'2) @ @ - @ DA>J@129 , @ >@, @ @ BDF @>@ , @@ BDF @130. @ @K@ @$C'2 @> BDF @>@ , @->@R >J@ BDF @, >@@

    131. J @ -@ @ J@ @(CEB) .

    J, G@>K-' .136 @ JI >> @ @>@ -@ BDF I@ . *J,

    @ J I >J@ @CG BDF @>@ $C'2 @@ BDF. F>@, @ I- J@ @ @ H3 BDF @>@

    , @ J@ @ @ @ H3.H@, J@ @ @ H3 @ @@ HDAC5 @ @ BDF , - H3 >@ >@ > -

    @ @@ I. @, G@>K-' .136 @ I -

    @@J@ @ CEB C$".

    (@@J) C$" @@-J@ @ CEB, CEB- @

    (CB'). >@ @ @ J@ --@>@ J , , BDF

    @. *J, CB' @ @ @J >@- @ @ @>@ @ @ >J, @ J @ I@ @ @ J J, @>

    >@ BDF @. F, @ @ @ >@- @ HA J @ CB' @ @->

    >>@J @ @-> >>@J , > @->>137-139.

    @ I- @@ @ > @

    @@ @ J@>, >J @>, AK>, @, > @, @@, E$ @, @ J-

    J @ (ADHD) J < @ BDF121,140-143. $@ J, A .144 @ JI J>@> @ ADHD. -@J 136, @ >@

    @> @ @ BDF , >

  • 7/24/2019 Epigenetic Regulation on Gene Expression Induced

    8/15

    !. -*@@@ .: E @ I

    139

    ), J @ @ BDF, >-@ @ @@ @> @ I >> @-> >>@J @> E-#' J @. , I-- @>@J >>, J - >@@136, @- @->@ @ '-C$" BDF, @ @145.

    A@ , J I @ J - BDF , @. D @> J >@ >@ >@ @ @-@ @ BDF >> J I > @146. , @ @ @ @ J @ @@ @ @ BDF/ @ I @ @>J @ @ @, AK>R .

    A @> - BDF, I @ J @ @> J HA/HDAC @.A @ >, I HDAC J HA J @>, - HA/HDAC @148. JJ@ @ @ @ @ -J4,27,149-155. A@, @ @ @@ J@ @ @@, @ @@, @ @ J@ >156. @ @ CB'-HAJ @ @ @-@. , HA/HDAC @ @

    HDAC J KJ> @. @I,

    I, @ J@ @HA/HDAC , >@ -

    J @@@ >>@J @157, @ @ -

    @ @J @ 158,159.

    A @ @J >@ >>@J,

    @

    160

    , (F 3). C@ .

    155

    @ @- I >> I@ J @ > @>,

    >@ @ @ @ J@@ . @-

    @ @@J@ @ @

    H3 -F@ @, J I. @@J@ @ H3 @ @ -F@

    @ @@ @ @J @

    @, I I@@ @@@ @ (G) , , -

    @ - @ @ @@@ @>@

    @161,162. H, J I -

    >@@, @ >-> @ , @@

    @ @ @ .

    E*%#!(!%c !""!c )" !!+c%! )( ca+%)/ac&a+

    !'

    'J I I @ > @ @-

    J>163. >@ >> @>@

    J @ @ @@ >@ @I -

    F%#+! 3. A @@ >@ @ @ I @ >@, @ @ @ . GF-1: -

  • 7/24/2019 Epigenetic Regulation on Gene Expression Induced

    9/15

    !. -*@@@ .: E @ I

    140

    J, @ @>@@ @ I >I, >@@164.

    @ @, @ @

    @ @@ >@@ J I-, @, @J.

    @ >@@ J JI J @ @@ I@ @ @->>@J J@J@ @ J I 71.

    E @ @ @ -@ @ FWB @ DA, @@ @->>@J J@

    166. A @> , @ @-@, >@J J@@@ @ -

    @ , >>@J @ J I@ @ ->>@J 168.

    H@, @ @ @ >> @ @ @ ,@ >J -

    @> @ @@ @71,169. D@@ HA/HDAC @, @ @ @, @ @ >@- I@ @ >I >@@ ($$'), @ @@ @ @ 170, @

    @@ @ @> >J@J -171, @ @>J@J172. JI @ @ @ -

    , J @ HA HDAC J @ @> @-

    @, J >>164. @, >A @ @ @ @ >J-

    @> >@ @, J, @ J @@

    >>(). EI @-@@ - @ >J@ >, J@J @-@ N O R P17. D I-

    - J@J, @> @ , @ - . @ @, >@ @ J16.

    @ @ I >@ ->@ >A, >A, , @@ >@, -@ I @ @ I-@ @ >A-1, -133 -133, @@@@ >J-

    A (@A), @ @@ @ 42 (CDC42), @@ @ A (E#FA) @, @ @-H@ J@> 2 (2)174. -

    @, @ I @>A-29, -29 -29, I@@ @ (C#A C#A)175, I@ @ >A-27 -27, -

    @ @-@ KJ> 1 (ACE)16, @ >A-143, I@ @ @-@ KJ> 2 (ACE2)16.A @ @>@ @ @@ , @>, -@ ,J, J@@ J@J174. $@-

    @, J@J @-@ , >, @@ @ I - I@ @ >A-126 , , - I@ @ >A ('3"2 *-1). , @ I @>@ @- @ EGF J @- @ J $A'" '3"/A @ @@ >@, J @> @ @@J - I@ @ >A173,174. @ @> >J , > @ >A >A->>, @ @ >A, >A--@, @ @ @ >J @ @ -17,177. @> > >@ @ @ >A > @ >@ 59.

    C)(c&%)( a( *+)*!c

    @ @ @ @ -@ J J I @ J@@

    J> @@. @ >@@ @ @ @ @ >@> J @>@ . J @ >@ >@ -J @ >>@J @. $@@, @> - >@@ @J @ @ JI @ @ @ @>J @ @, @J @ . F @ >> @ @ @ J J I > J I @ /@@>>J J - -. F>@, @ @ > J> J @ @>, @ J> @ J J I, @, @ @@ > @ @. - I- @ >@ @ /@ @@ @ @-, >@ @ @ , J @ J @ @ @ @-> @ >J @ @.A @@ @ > @

  • 7/24/2019 Epigenetic Regulation on Gene Expression Induced

    10/15

    !. -*@@@ .: E @ I

    141

    J I @ I, >@ @ @, @ @ J@-@ J>. @I, I >

    @ @ @>@ >@ @@ @ @@ >@ @ .

    R!"!+!(c!

    1. B , !, E B, . A I >@@>@ >J@ > . C$ 2012;15(3):405-11.

    2. A E@ D, C G, B $, D . E. 2007;447(7143):396-440.

    3. # B, CJ $, @ !#. @ @ @> - @. C 2007;128(4):707-19.

    4. $G *#, H $. H@ >@@ I- @. ! A 'J@ 2011;110(1):258-63.

    5. $"J A, C#, @ E. C@@ @ >@> J HA HDAC. C G D 2001;11(5): 497-504.

    6. F !, F@ *, F G. E @ @ I@ @ @. ' 2007;61(5 ' 2):58-63.

    7. !C, A CD. H@ >J@ @J@: @ @. C C B@ 2001;13(3):263-73.

    8. D@ A, '< H, B, . D >J@ @- - CG @ > @ > , >J@ > @. G 2009;41(12):1350-3.

    9. ' . @ @ >J@ I@. E@ 2008;1(1).

    10. B< D, !, * C, . > @ >@A@ @ @. 2008;455(7209):64-71.

    11. B>@ D, B@ $', RC@ $, . $@-A: @ @ >> @> @. >>@ 2008;9(8):839-45.

    12. D> A$, " $, * $, . >-148 - > D$3 @ @ @. A 2008;14(5):872-7.

    13. AK-* $, A@ G, J A, .>A-132 @ @> >@

    @ @@ @ @ $@

    G 20(4):731-51.14. #J $A. >@A >> @.

    >>@ 2008;29(7):343-51.15. A> A!, $ !, @> B. >A -

    @ @ J@-@ , J -@ @ @-@ KJ> 2-@(1-7). HJ@ 2011;58(2):182-9.

    17. E@ G$, CD, K C, @ D. 'J-@@ >@ @ @ I-

    : >@ >>. H2012;98(1):5-10.

    18. *-G@> F, G-G>K !#, 'K- C,

    . 'J I >@@: E,

    O@ P @ @ I@. !* C@ 2012;26(12):3469-3472.

    19. B@ , C@@ BD, . E @

    > I @ >>@J I-@ > . A 'J@ $2009;34(4):745-753.

    20. H $', "@@@@ B, . $ @- J I@ @J @>@ I . A> ! 'J@ C@> 'J@ 2010;299(5):1254-1262.

    21. D>>@ $!, F *, A , . H> > I@ @@ I @@ @@. $ * *@ EI 2008;40(4):691-8.

    22. !@ *B, EA, @K@ @->@ @ @- >J@. ! C 2007;117(9):2459-67.

    27. $G *#, F E, G> A', H $. EI-

    - @ >@@ > . ! 'J@ 2009;587(' 24):5951-8.

    28. B@- K@ @J 4. $@ C B@ 2008;28(10):3437-45.

    29. $"J A, C#, # !, @ E. *--

    I@ @ @ J > @. 2000;408(6808):106-11.

    30. $G *#, H $. EI @ @ 4 I@: >@ >>.

    C EI '>@ 'J@ 2006;33(4):395-9.31. C *, BK@KJ *, # *, @ E. A I-

    @ >@@ @ @ J-

    @@J@. '@ A * * A 2005;102(23):8120-8125.

    32. "> $*, FK !, $AJ !, . '@ $EF2 J

  • 7/24/2019 Epigenetic Regulation on Gene Expression Induced

    11/15

    !. -*@@@ .: E @ I

    142

    > @>. $@ C B@ 2008;28(11):3600-9.33. # !, $"J A, C#, @ E. @ @

    J@ J @@ @ $EF2 -

    @ @ @ J. $@C 2000;6(2):233-44.

    34. F , D F, HK $!, . EKJ> -

    J @ HDAC @ >@ @>I @ HDAC3 *$/-C@. $@ C 2002;9(1):45-57.

    35. CKJ $', $AJ !, F> G, @ E. -

    @ @ @I@> @@- @>> @@ 1 ('GC-1 ) >@@- @ J $EF2 HDAC5. '@ A * * A 2003;100(4):1711-6.

    36. $G *#, * D, @ A#, H $. EI- $EF2- GEF DA- J > . FA*EB ! 2006;20(2):348-9.

    37. ", $G *#, @@ C, . E @ I @ > $EF2 @

    I. A> ! 'J@ E@@ $ 2008;294(2):E408-15.

    38. *> !A, "@ A, CJ A", @ #' #(. A> ! 'J-@ E@@ $ 2007;292(2):E413-20.

    40. E B, C@ B', G-@ '$, . EI -

    J- @ @ @I@> @@-

    @ @@-1 >A @ @ @ > - . ! 'J@ 2010;

    588(' 10):1779-90.41. !, " ", ' H. 'GC-1->

    @ . ' A 2010;

    460(1):153-62.42. B , $E, !, . @-CG >J@

    @ 'GC-1 @>@ @ D$3B @@>@@ J. C $ 2009;10(3):189-198.

    43. # C, G@@ #. E: >@ < @> @ J 2 . D 2009;58(12):2718-25.

    44. B , !. DA >J@ >@ -@. A> ! C 2011;93(4):897*-900.

    45. B ". E @@ @ J.A 'J@ (I) 2010;199(4):477-87.

    46. '@ CE, H F, C, . D - >@@ @ @ >J@ J @

    @ @ > @. A> ! 'J@ C 'J@2009;297(1):C6-16.

    47. @@ "A, #J $!, B@@ F. @ @ @ >J@ J .

    F@ @ OD >@@ @ -@ >J@ J @

    @ @ > @-P. A> ! 'J@ C 'J@ 2009;297(1):C1-3.

    48. 'I> @IJ ! 'J-

    @ E@@ $ 2008;294(6):E1152-9.49. ' , C> , * *, !. $

    $F @ J -

    @> @ @ I2. @ 2002;21(17):2729-40.

    50. # !B, ! A, *", . @J @@ @

    @@ @ @: @ @I/C/A$# @. C E, @.! 'J@ 2010;588(' 21):4075-87.

    53. > $, $@-

    @ I J . E ! ' C-@ 2012.

    54. B A#, H A, B, . DJ> -@ @ >@A I I-

    @ I . ! 'J@2011;589(' 16):3983-94.

    55. " ', @ B, G@ , . A -

    @ > @ > @ I @ @J. ! A 'J@ 2011;

    110(1):46-59.56. D>>@ $!, $CJ !!, FJ C*, . A -

    J > >@A I-@ @ > @ I >@ . A> ! 'J@ E-@@ $. 2008; 295(6): E1333-1340.

    57. A@ , @ , $K> ", . >@A >-696 'GC-1LM >@ @ @ J J. A> ! 'J@ E@@

    $ 2010;298(4):E799-806.58. * A, A A, AA -

    @ @ @ @ I C57B/6! > >. '#@* 2009;

    4(5):5610.59. F , $ FC, @ F, . EI

    >@ @ J-@ @ @@ @: @

    @ >@A-16, -21, -126. HJ@ 2012;59(2):513-20.

    60. *, * C, C, . $ >-

    @A J I >. ! 'J@ 2010;588(' 20):4029-4037.

    61. A>@ !#, @ . @ J- J >>@ >@ J J

  • 7/24/2019 Epigenetic Regulation on Gene Expression Induced

    12/15

    !. -*@@@ .: E @ I

    143

    >- @ * . A @ > 2002;162(11):1286-92.

    62. A C, * A, D@ A, . A@@ @> @ J J >>@. $ @

    I 2009;41(9):1706-13.

    63. @> E, > D. >>@J @, J- J, J J@ @. *- @ @ > & @ 2008;

    18(5):543-56.64. $>@@ !, *, AJ@ -1- -- J. ! >>@ 2006;177(7):4252-6.

    73. @>-AK< *, F !, *, . E@ >@A @> I-@ -@ I@ . ! A 'J@ 2010;

    109(1):252-61.74. @>-AK< *, F !, # *, . E @

    I @ >@A I@ J@ > - @@ >@@ . C @

    2012;5(1):32-8.75. * @@ @@ J@ @ -J . B@@@@J 2007;8(2):173-87.

    77. R* !!, $J '!. CJ@@ -

    >>@J @. >>J 2008;28(4):477-87.

    78. B@ A, #@J !A, #> !, H@ *E. @ *@ @ J >>J.$@ >>@@J. 2004;41(6-7):715-25.

    79. H@ $, C. @ @ @J I. EI >>@@J 2010;16:163-83.

    80. $> @ I. EI

    >>@@J 2011;17:6-63.85. # !, C !, E '!, . >-181

    >@@ @ J @. C 2007;129(1):147-61.

    86. *>@ !, GJ ". C@ >>J

    J J: > J @? - >-. G@@@J 2010;56(5):449-58.

    87. @ "D, B@ $', C "!, B>@ D. F-@A >-146, @ @ @ @ >>@. '@ @ @ A>J @ *-

    @ * @ A> 2006;103(33):12481-6.

    88. R #A, *J F!, $C@J CE. $@A:

    - @ @->@@J 2011;11(3):163-75.

    89. @ , FJJ-"K H, E , . H> - >@A : @ @ >@A-31

    >@A-21 F'3 I@. E@ @@ >>@@J 2009;39(6):1608-18.

    90. ' , *, $, . $@A-21 >-@A-148 @ @ DA J@>J@

    CD4+ J J J DA >J 1. ! >>@ 2010;184(12):6773-81.

    91. D , A> *A. $@A, > @ - >>@@, >>@, -@>> . @ : @ @@@J > 2011;157(4):163-79.

  • 7/24/2019 Epigenetic Regulation on Gene Expression Induced

    13/15

    !. -*@@@ .: E @ I

    144

    92. $>@ $ A, BJ , C !, H@ , . F@@,@, J J, @ @ : @ . @ C F/A>-

    @ C 2007: 517.93. "> !B, B *, B@ CE, "@ H 3. 'J

    J, J , - >@-

    J: @ J @ > @>. A @>@@J 1996;6(5):452-7.

    94. H@> $!, *K A. C @ @DA J@>J@ > . B@>J

    @@J = B@> @@ 2005;83(3):296-321.

    95. FF, C , C@ !, . 'J J @ @> DA >J@ -

    @@. E: @ @ @ DA$J@ *@J 2011;6(3):293-9.

    96. E $, C@ 'G, BJ *B, H> !G. A J-

    >J@ @ @ > . C .2001;61(8):3225-9.

    97. !@ 'A, BJ *B. > @ @ . G 2002;3(6):415-28.

    98. , J@ J @ @ J>J@ @>- . C E>@ B@> A@ C 2005;46:2744.

    102.@ #, *> A, $ *, C G. G@> 5->J@IJJ . !@ @@@ >J 1987;262(21):9948-51.

    103. !, "@@ , # , . @ @ -

    CG >J@ J @ >< @> . $>@ @> 2002;123(11):1487-503.

    104. C #, @ A. C@> @ . E-: @ @ @ DA $J@ *@J2009;4(1):23-6.

    105.B . .

    2000;408(6808):53-6.106. "> ', 'K $, #K !F, . @J@@-

    @ J I @

    @. A 2011;10(4):475-86.107. *@ *, B@ $A. C@> @ @-

    @ A >>> @>. *> & @> @@J 2010;21(2):186-93.

    108. C, H@ $, > , . E @ J- I @ >J@ @>- @,

    J, @@. !@ @ A>- C@ @ C@@J 2008;52(6):470-82.

    109. C, F , > , . 'J I

    @ @>. 2000;408(6809):255-62.116. G@ !, , $@ , . A @ J -

    >>@J J *1 >-134. - 2010;466(7310):1105-9.

    117. C@J EH, 'J G*. CJ@>@ >@ . E ! $ 2001;344(12):907-16.

    118. C@ ', @ !, E. >>@J J@ C- @ @@ J > @@-

    J J >@@ > . C@ 2003;108(16):1930-2.

    119. * !, "@>@J J-@BDF > J @ I @ J -J @@. E ! @ 2004;20(10):2580-90.

    122. C, C C, !> , . E @ I @

    $DA @ @@ @ @J J >@ J. H@>- 2007;17(12):1201-8.

    123. # , *@@ G. B@@ >> @ J

    J @ . C $@ @-@ 2009;30(4):493-503.

    124. $@ ", $ H, # B. @ BDF

    @ @ IJ. @ 2007;10(9):1089-93.

    125. B@ C, C G, C@ $, . EI > >@ >>@J @ - @@ @

  • 7/24/2019 Epigenetic Regulation on Gene Expression Induced

    14/15

    !. -*@@@ .: E @ I

    145

    @ @ @>. @2005;133(3):853-61.

    126. B@ #$, '@@ $$. 'J @K@ - @@- J @@. @ 1999;2(4):346-51.

    127. G E, B G, B A$, "J A$. EI

    @>- : - @ BDF- >>. H@>2009;19(10):973-80.

    128. G@>K-' F, J> *, . B- -@@ @ @ >@@ @ > @ I @ @@. E ! @ 2008;

    28(11):2278-87.129. C@ H, @ H. J J @ $C'2 @-

    @J@. '@ A * * A 2009;106(12):4577-8.

    130. $@ ", H@ D, H, . DA >J-@- @> >@ J-BDF @. * 2003;302(5646):890-3.

    131. C G, C , # , . D@ @ BDF@ @ >- @@J-@ @ $C'2. * 2003;302(5646):885-9.

    132. * #, *> . C 1994;77(1):83-93.

    133. J> *, , G@>K-' F. J

    - @@ @ @>@@ @ @ @ I @J-J. @ 2003;122(3):647-57.

    134. *A, G@>K-' F, C@ !, C@> C. 'J- J >A @ - -

    @@ @ @ @ .B 1996;726(1-2):49-56.

    135. G@>K-' F, , @J , . @J I- BDF-> >> @>@@J. ! @J@ 2002;88(5):2187-95.

    136. G@>K-' F, , F !, . EI >-

    - @@ @ J J - >> @ @. E !@ 2011;33(3):383-90.

    137.B !. @ @> @ >>@J @>@ >. A A * 1990;608:457-77; @ 477-84.

    138. $" D, * #. @> @ -

    >>@J. ! EI 'J@ # $> C@ 1993;19(2):397-404.

    139. $@ G, G ', !, R" !. ' -

    @ > @> @. 1982;297(5868):681-3.

    140. C F, C@ A, @ F, D@ F. D@ AK>R : @@@ >@

    >@@ . E ! '>@ 2010;626(1):64-71.

    141. D> CH, * #, D*, D> *. @-J I @ I@J

    @ >. B 2008;1199:148-58.142. # FD, @ #, * !D. E @ @

    BDF @ @@@ @ >>@J. ! @ 2008;28(42):10576-86.

    143. @> @ >@ >@ @ -

    @ @. @ 2006;9(4):519-25.

    144.A , "@K $. 'J I

    ADHD J>@>: @ @> -@J. @@I 2012;21(2):195-209.

    145. $, A , D@ A, .

    @ I @ @@ E$ -@: @, @J@@ >@ -. @@ D 2011;45(3):1153-62.

    146. # ', # , . EI

    @@ >>@J >> >@ @> @. F@ B@ 2010;3:34.

    147. # ', #@: @- @ C'D. A@I @I * 2008;10(4):799-811.

    152.* BD, A CD. @ @ @

    >@@. 2000;403(6765):41-5.153. C@ !", H@ #!. H@ J@: @ @-

    ? B@> C B@ 2009;87(1):139-50.

    154. # $, B@@ G. @@ DA . B@ -@ 2005;73(2):289-96.

    155. C@ A, H #E, C>@ , . EI >-

    @ @ @ @ J@@ @ > @ >> I@ J. '#@* 2009;4(1):4330.

    156. * , ' ". HA HDAC @-@: @ @ J@ @>@. CD D 2006;13(4):539-50.

    157. * B, C !C, " ", B@ A#. J@ HA @: -J @ . B@> B@J A2010;1799(10-12):840-53.

  • 7/24/2019 Epigenetic Regulation on Gene Expression Induced

    15/15

    !. -*@@@ .: E @ I

    146

    158. "> AF, H *, C@ !, . A, @@ @. 1999;400(6743):418-9.

    159.$@ $'. @@ : J @@ > . B 2000;886(1-2):47-53.

    160.F A, * F, , . @J @

    >>@J @ @> >@-. . 2007; 447(7141): 178-182.

    161. !$, C>@ . E >> - >>@J @>@. 'J@@-@@@J.2007;32(*.1):*21-25.

    162. D@ *", C>@ , H #E, . @JI > @ J@>-J-@@ I >J . @-@@@J 2007;86(1):26-37.

    163. G@ *, # . EI @ J->. C@ ' 2012;2012:210852.

    164. B@ , * F, >> '. $@ >> @I- @ @. @

    , >@@ . HK2012;37(5):508-15.

    165.B '!, " $. @->@J . E ! $ 1997;336(15):1066-71.

    166. @ ", HK , @> ", . I - @ J 2 J #-8 I@: @ @ J@ @. B@> B@-J C@>> 2004;315(1):240-45.

    167.$@ F, G@K@ G, # #, . I' #(@> >@ @ >>@J @ @@. ! B@ C>.2004;279(17):18091-7.

    168. @I >@@ >I - J @ @ > @@. ! C 1994;94(6):2493-503.

    171.' G$, C@ E, A@ G, . $@ - @ @: @ -@. ! B@> * 2005;12(6):839-53.

    172. $@@>J #, '@@ $!, H $, . $- @ J >@> J @ J- 3 >. ! C 2008;118(11):3588-97.

    173. F-* $$, C@ , G> G, . 'J I >@A: @ . A B C@ 2012;98(5):459-66.

    174. F , *@ ', E$. E @-

    J@J J I :>@A >@ >. BK ! $ B@ 2011;44(9):836-47.

    175. *@ ', F , H>@@ , . $@A29 @ >@> @ @>- @>@ J @ I .'J@ G@> 2011;43(11):665-73.

    176. DA * D !, F , *@ ', . *>> - $@A-126 I@ @. $ * *@ EI 2012; 44(8): 1453-62.

    177. B@ BC,