ヌクレオソームは、4つのコアヒストンタンパク質 (H2A、H2B、H3、H4) とリンカーヒストンH1から構成される、クロマチンの主要な基礎構成成分です。ヒストンはもともと、DNAをパッケージングする静的な足場として機能すると考えられていましたが、最近では、多様な翻訳後修飾を受けクロマチン凝縮とDNAへの接近しやすさを調節する、動的なタンパク質であるということが明らかになってきました。例えば、リジン残基のアセチル化は、ヒストンの移動や転写活性化に関わっており、最近ではDNA修復にも関与することがわかっています。セリンおよびスレオニン残基のリン酸化は、有糸分裂中のクロマチン凝縮および最初期遺伝子の転写活性化を促進します。リジンおよびアルギニン残基のメチル化は、クロマチンの転写活性化領域および転写不活性領域の形成の主要な決定因子として機能し、発生段階のゲノムの適切なプログラミングに重要です。
ヒストン | 部位 | ヒストン修飾酵素 | 提唱されている機能 | 参考文献# |
---|---|---|---|---|
H2A | Lys4 (S. cerevisiae) | Esa1 | transcriptional activation | (1) |
Lys 5 (mammals) | Tip60, p300/CBP | transcriptional activation | (2,3) | |
Lys7 (S. cerevisiae) | Hat1 | unknown | (4) | |
Esa1 | transcriptional activation | (1) | ||
H2B | Lys5 | p300, ATF2 | transcriptional activation | (5,3) |
Lys11 (S. cerevisiae) | Gcn5 | transcriptional activation | (6) | |
Lys12 (mammals) | p300/CBP, ATF2 | transcriptional activation | (5,3) | |
Lys15 (mammals) | p300/CBP, ATF2 | transcriptional activation | (5,3) | |
Lys16 (S. cerevisiae) | Gcn5, Esa1 | transcriptional activation | (6) | |
Lys20 | p300 | transcriptional activation | (3) | |
H3 | Lys4 (S. cerevisiae) | Esa1 | transcriptional activation | (1) |
Hpa2 | unknown | (7) | ||
Lys9 | unknown | histone deposition | (8) | |
Gcn5, SRC-1 | transcriptional activation | (10,9) | ||
Lys14 | unknown | histone deposition | (8) | |
Gcn5, PCAF | transcriptional activation | (11,3) | ||
Esa1, Tip60 | transcriptional activation | (2,1) | ||
DNA repair | (11,12) | |||
SRC-1 | transcriptional activation | (10) | ||
Elp3 | transcriptional activation (elongation) | (13) | ||
Hpa2 | unknown | (7) | ||
hTFIIIC90 | RNA polymerase III transcription | (14) | ||
TAF1 | RNA polymerase II transcription | (15) | ||
Sas2 | euchromatin | (16) | ||
Sas3 | transcriptional activation (elongation) | (17) | ||
p300 | transcriptional activation | (3) | ||
Lys18 | Gcn5 | transcriptional activation, DNA repair | (9) | |
p300/CBP | DNA replication, transcriptional activation | (18,3) | ||
Lys23 | unknown | histone deposition | (8) | |
Gcn5 | transcriptional activation, DNA repair | (9) | ||
Sas3 | transcriptional activation (elongation) | (17) | ||
p300/CBP | transcriptional activation | (18,3) | ||
Lys27 | p300/CBP | transcriptional activation | (93, 94) | |
Lys36 | Gcn5 | transcriptional activation | (82) | |
Lys56 (S. cerevisiae) | Spt10 | transcriptional activation | (19) | |
DNA repair | (20) | |||
H4 | Lys5 | Hat1 | histone deposition | (21) |
Esa1, Tip60 | transcriptional activation | (2,1) | ||
DNA repair | (11,12) | |||
ATF2 | transcriptional activation | (5) | ||
Hpa2 | unknown | (7) | ||
p300 | transcriptional activation | (3) | ||
Lys8 | Gcn5, PCAF | transcriptional activation | (22,3) | |
Esa1, Tip60 | transcriptional activation | (2,1) | ||
DNA repair | (11,12) | |||
ATF2 | transcriptional activation | (5) | ||
Elp3 | transcriptional activation (elongation) | (13) | ||
p300 | transcriptional activation | (3) | ||
Lys12 | Hat1 | histone deposition | (21) | |
telomeric silencing | (23) | |||
Esa1, Tip60 | transcriptional activation | (2,1) | ||
DNA repair | (11,12) | |||
Hpa2 | unknown | (7) | ||
p300 | transcriptional activation | (3) | ||
Lys16 | Gcn15 | transcriptional activation | (22) | |
MOF (D. melanogaster) | transcriptional activation | (24) | ||
Esa1, Tip60 | transcriptional activation | (2,1) | ||
DNA repair | (11,12) | |||
ATF2 | transcriptional activation | (5) | ||
Sas2 | euchromatin | (6,2) | ||
Lys91 (S. cerevisiae) | Hat1/Hat2 | chromatin assembly | (25) |
ヒストン | 部位 | ヒストン修飾酵素 | 提唱されている機能 | 参考文献# |
---|---|---|---|---|
H1 | Lys26 | Ezh2 | transcriptional silencing | (48, 49) |
H2A | Arg3 | PRMT1/6, PRMT5/7 | transcriptional activation, transcriptional repression | (83) |
H3 | Arg2 | PRMT5, PRMT6 | transcriptional repression | (83) |
Arg8 | PRMT5, PRMT2/6 | transcriptional activation, transcriptional repression | (31, 88) | |
Arg17 | CARM1 | transcriptional activation | (18) | |
Arg26 | CARM1 | transcriptional activation | (83) | |
Arg42 | CARM1 | transcriptional activation | (89) | |
Lys4 | Set1 (S. cerevisiae) | permissive euchromatin (di-Me) | (26) | |
Set 7/9 (vertebrates) | transcriptional activation (tri-Me) | (27) | ||
MLL, ALL-1 | transcriptional activation | (28, 29) | ||
Ash1 (D. melanogaster) | transcriptional activation | (30) | ||
Lys9 | Suv39h, Clr4 | transcriptional silencing (tri-Me) | (32, 33) | |
G9a | transcriptional repression genomic imprinting | (34) | ||
SETDB1 | transcriptional repression (tri-Me) | (35) | ||
Dim-5 (N. crassa), Kryptonite (A. thaliana) | DNA methylation (tri-Me) | (36, 37) | ||
Ash1 (D. melanogaster) | transcriptional activation | (30) | ||
Lys27 | Ezh2 | transcriptional silencing | (38) | |
X inactivation (tri-Me) | ||||
G9a | transcriptional silencing | (34) | ||
Lys36 | Set2 | transcriptional activation (elongation) | (39) | |
Lys79 | Dot1 | euchromatin | (40) | |
transcriptional activation (elongation) | (41) | |||
checkpoint response | (42) | |||
H4 | Arg3 | PRMT1/6 | transcriptional activation | (43) |
PRMT5/7 | transcriptional repression | (31) | ||
Lys20 | PR-Set7 | transcriptional silencing (mono-Me) | (44) | |
Suv4-20h | heterochromatin (tri-Me) | (45) | ||
Ash1 (D. melanogaster) | transcriptional activation | (30) | ||
Set9 (S. pombe) | checkpoint response | (46) | ||
Lys59 | unknown | transcriptional silencing | (47) |
ヒストン | 部位 | ヒストン修飾酵素 | 提唱されている機能 | 参考文献# |
---|---|---|---|---|
H1 | Ser27 | unknown | transcriptional activation, chromatin decondensation | (48,49) |
H2A | Ser1 | unknown | mitosis, chromatin assembly | (50) |
MSK1 | transcriptional repression | (51) | ||
Ser122 (S. cerevisiae) | unknown | DNA repair | (53) | |
Ser129 (S. cerevisiae) | Mec1, Tel1 | DNA repair | (54,55) | |
Ser139 (mammalian H2A.X) | ATR, ATM, DNA-PK | DNA repair | (56-58) | |
Thr119 (D. melanogaster) | NHK1 | mitosis | (52) | |
Thr120 (mammals) | Bub1, VprBP | mitosis, transcriptional repression | (90,91) | |
Thr142 (mammalian H2A.X) | WSTF | apoptosis, DNA repair | (92) | |
H2B | Ser10 (S. cerevisiae) | Ste20 | apoptosis | (59) |
Ser14 (vertebrates) | Mst1 | apoptosis | (60) | |
unknown | DNA repair | (61) | ||
Ser33 (D. melanogaster) | TAF1 | transcriptional activation | (62) | |
Ser36 | AMPK | transcriptional activation | (84) | |
H3 | Ser10 | Aurora-B kinase | mitosis, meiosis | (64,65) |
MSK1, MSK2 | immediate-early gene activation | (66) | ||
IKK-α | transcriptional activation | (67) | ||
Snf1 | transcriptional activation | (68) | ||
Ser28 (mammals) | Aurora-B kinase | mitosis | (70) | |
MSK1, MSK2 | immediate-early activation | (66,71) | ||
Thr3 | Haspin/Gsg2 | mitosis | (63) | |
Thr6 | PKCβI | (85) | ||
Thr11 (mammals) | Dlk/Zip | mitosis | (69) | |
Tyr41 | JAK2 | transcriptional activation | (86) | |
Tyr45 | PKCδ | apoptosis | (87) | |
H4 | unknown | mitosis, chromatin assembly | (50) | |
Ser1 | CK2 | DNA repair | (71) |
ヒストン | 部位 | ヒストン修飾酵素 | 提唱されている機能 | 参考文献# |
---|---|---|---|---|
H2A | Lys119 (mammals) | Ring2 | spermatogenesis | (73) |
H2B | Lys120 (mammals) | UbcH6 | meiosis | (74) |
Lys123 (S. cerevisiae) | Rad6 | transcriptional activation | (75) | |
euchromatin |
ヒストン | 部位 | ヒストン修飾酵素 | 提唱されている機能 | 参考文献# |
---|---|---|---|---|
H2A | Lys126 (S. cerevisiae) | Ubc9 | transcriptional repression | (76) |
H2B | Lys6 or Lys7 (S. cerevisiae) | Ubc9 | transcriptional repression | (76) |
H4 | N-terminal tail (S. cerevisiae) | Ubc9 | transcriptional repression | (77) |
ヒストン | 部位 | ヒストン修飾酵素 | 提唱されている機能 | 参考文献# |
---|---|---|---|---|
H2A | Lys9 | biotinidase | unknown | (78) |
Lys13 | biotinidase | unknown | (78) | |
H3 | Lys4 | biotinidase | gene expression | (79) |
Lys9 | biotinidase | gene expression | (79) | |
Lys18 | biotinidase | gene expression | (79) | |
H4 | Lys12 | biotinidase | DNA damage response | (80,81) |
こちらの表を監修いただいたマサチューセッツ大学医学部Craig Peterson教授 (マサチューセッツ州ウースター) に深く感謝いたします
作成日: 2007年5月
改訂日:2018年3月