Эпигенетика

Приложение 2. Модификации гистонов и литература

Сайт | Модель | Энзим | Функция | Ссылка

K5ac | Hs, Sc | Tip60, p300/CBP, Hat1 | Активация транскрипции | Yamamoto and Horikoshi, 1997; Kimura and Horikoshi, 1998; Verreault etal., 1998

K9bio | — | HCS-биотинидаза | Зависимая от ацетилирования и метилирования. Участвует в клеточной пролиферации, сайленсинге генов и в клеточном ответе на повреждение ДНК | Stanley et al., 2001; Kothapalli et al., 2005b; Chew et al., 2006

K7ac | Sc | Hat1 Esa1 | Активация транскрипции | Suka et al., 2001

K13bio | — | HCS-биотинидаза | Зависимая от ацетилирования и метилирования. Участвует в клеточной пролиферации, сайленсинге генов и клеточном ответе на повреждение ДНК | Stanley et al., 2001; Kothapalli et al., 2005b; Chew et al., 2006

K119ub1 | Dm. Hs | dRing, RING IB | Polycomb-сайленсинг, Реакция на УФ-повреждение | Wang et al., 2004; Kapetanaki et al., 2006

S121ph | Sc | Mec1 PIKK | Ответ на повреждение ДНК. | Теломерный сайленсинг | Wyatt et al., 2003; Harvey et al., 2005

T125ph | Sc | Mec1 PIKK | Ответ на повреждение ДНК. Теломерный сайленсинг | Wyatt et al., 2003

K126bio | Hs | HCS биотинидаза | Зависимая от ацетилирования и метилирования. Участвует в клеточной пролиферации, сайленсинге генов и клеточном ответе на повреждение ДНК | Stanley et al., 2001; Kothapalli et al., 2005b, Chew et al., 2006

K126su | Sc | Репрессия транскрипцииБлокирует ацетилирование и убиквитинирование гистонов | Nathan et al., 2006

K127bio | Hs | HCS-биотинидаза | Зависимая от ацетилирования и метилирования. Участвует в клеточной пролиферации, сайленсинге генов и клеточном ответе на повреждение ДНК | Stanley et al., 2001; Kothapalli et al., 2005b; Chew et al., 2006

S128ph | Sc | Mec1 PIKK | Ответ на повреждение ДНК. Теломерный сайленсинг | Downs et al., 2000; Redon et al., 2003; Wyatt et al., 2003; Downs et al., 2004

K130bio | Hs | HCS-биотинидаза | Зависимая от ацетилирования и метилирования. Участвует в клеточной пролиферации, сайленсинге генов и клеточном ответе на повреждение ДНК | Stanley et al., 2001; Kothapalli et al., 2005b; Chew et al., 2006

Дополнительные модификации Н2А:

S1ph, К4ас, K13me, K15ас, K21ас, K36ас, K74me, K75me, R77me, K95me, T120ph, карбонилирование Н2А (Pantazis and Bonner, 1981; Song et al., 2003; Alhara et al., 2004).

Сайт | Модель | Энзим | Функция | Ссылка

К5ас | Hs | — | Активация транскрипции | Puerta et al., 1995; Galasinski et al., 2002

S10ph | Sc | Ste20 | Апоптоз | Ahn et al., 2005

S14ph | Hs, Mm | Mst1/krs2 киназа | Апоптоз. Соматическая гипермутация и рекомбинация с переключением классов | Ajiro, 2000; Cheung et al., 2003; Odegard et al., 2005

K16su | Sc | — | Репрессия генов | Nathan et al., 2006

K17su | Sc | — | Репрессия генов | Nathan et al., 2006

S33ph | Dm | CTK | Активация транскрипции | Maile et al., 2004

K120ub | Hs | — | Прогрессия по клеточному циклу в сочетании с SAGA для транскрипционной активации посредством метилирования H3, мейоз | Robzyk et al., 2000; Sun and Allis, 2002; Kao et al., 2004

K123ub | Sc | Rad6(E2)Brel(E3)-ub1 | Теломерный сайленсинг путем снижения метилирования гистонов по H3K4 и H3K79 | Emre et al., 2005

Дополнительные модификации Н2В:

E2am, K5me, K6su, K7su, K11ac, K12ас, K15ас, K16ас, K20ас, K23me, K24ас, S32ph, K43me, K85ас, R99me, K108ас, K116ас, Kарбонилирование Н2В, биотинилирование Н2В (Rouleau et al., 2004).

Сайт | Модель | Энзим | Функция | Ссылка

R2me | Hs Mm | CARM1-me2a | Экспрессия генов | Chen et al., 1999; Schurter et al., 2001

T3ph | Hs At | Haspin | Центромерная функция митотического веретена | Polioudaki et al., 2004; Dai et al., 2005

K4me | Sc | Setl-me3 | Сайленсинг rDNA, теломерный сайленсинг. Активация транскрипции | Briggs et al., 2001; Roguev et al., 2001; Nagy et al., 2002; Bryk et al., 2002; Bernstein et al., 2002; Santos-Rosa et al., 2002

K4me | Tt | ? | Активация транскрипции | Strahl et al. 1999

K4me | Hs | SET7/Set9-me1 | Активация транскрипции | H. Wang et al., 2001a; Nishioka et al., 2002a; Wilson et al., 2002; Zegerman et al., 2002

K4me | Ds Hs | MLLMLL2-me3 MLL3 | Активация Trithorax | Milne et al., 2002; Nakamura et al., 2002

K4me | Ds Hs | Ash1-me2 | Активация Trithorax | Beisel et al., 2002; Sanchez-Eisner et al., 2006

K4me | Hs | SMYD3-me3 | Активация транскрипции | Hamamoto et al., 2004

K9ac | Mm Sc | Meisetz-me3SAGA | Прогрессия мейотической профазы. Активация транскрипции | Hayashi et al., 2005 Grant et al., 1999

K9ac | Hs | SRC1 | Коактиватор ядерного рецептора | Spencer et al. 1997; Schubeler et al., 2000: Vaquero et al., 2004

K9ac | Dm | Активация транскрипции | Nowak et al., 2000

K9me | Mm, Hs | G9a-me1, me2 | Активация транскрипции. Импринтинг | Tachibana et al., 2001, 2002; Ogawa et al., 2002; Xin et al., 2003

K9me | Dm | Su(var)3-9-me2 | Доминантный модификатор PEV | Czermin et al., 2001; Schotta et al., 2002; Ebert et al., 2004

K9me | Mm | Suv39h1-me3 Suv39h2-me3 | Перипентрический гетерохроматин | O’Carroll et al., 2000; Rea et al., 2000; Lachner et al., 2001; Peters et al., 2001

K9me | Hs | SUV39H1-me3 | Rb-опосредованный сайленсинг | Nielsen et al., 2001; Vandel et al., 2001

K9me | Sp | Clr4-me1, me2 | Сайленсинг центромеры и типа спаривания | Bannister et al., 2001; Nakayama et al., 2001

K9me | Nc | Dim5-me3 | Метилирование ДНК | Tamaru and Selker, 2001

K9me | At | KRYPTONITE-me2 | Метилирование ДНК | Jackson et al., 2002, 2004

K9me | Hs | EuHMTase 1-me1, me2 | Репрессия транскрипции | Ogawa et al., 2002; Tachibana et al., 2005

K9me | Hs, Mm | ESET-me2, me3 | Репрессия транскрипции | Schultz et al., 2002; Yang et al., 2002; Dodge et al., 2004; Wang et al., 2004

K9me | Dm, Hs | Ash1-me2 | Активация Trithorax | Beisel et al., 2002

K9me | Hs | R1Z1-me2 | Супрессия опухоли и ответ на женские половые гормоны | Kim et al., 2003; Carling et al., 2004

S10ph | Sc | Snf1 | Активация транскрипции | Lo et al., 2001

S10ph | Dm | |il-1 | Транскрипционная ап-регуляция мужской Х-хромосомы | Jin et al., 1999; Y. Wang et al., 2001

S10ph | Hs | Rsk2 | Активация транскрипции немедленных ранних генов (в сочетании с ацетилированием H3K14) | Sassone-Corsi et al., 1999; Thomson et al. 1999;

S10ph | Hs | Msk1 | Активация транскрипции немедленных ранних генов (в сочетании с ацетилированием H3K14) | Cheung et al., 2000;

S10ph | Hs | Msk2 | Активация транскрипции немедленных ранних генов (в сочетании с ацетилированием H3K14) | Clayton et al. 2000

S10ph | Hs | IKKa | Транскрипционная ап-регуляция | Anest et al., 2003; Yamamoto et al, 2003

S10ph | Sc, Ce | Ipl1/AuroraB | Конденсация митотических хромосом | Hendzel et al., 1997; Wei et al.. 1999: Hsu et al., 2000

S10ph | An | NIMA | Конденсация митотических хромосом | De Souza et al., 2000

S10ph | Hs, Ce | Fyn kinase | UVB-индуцированный МАР-киназный путь | He et al., 2005

T11ph | Hs | Dlk/ZIP | Митоз-специфичное фосфорилирование | Preussetal., 2003

K14ac | Sc, Tt, Mm | Gcn5 | Активация транскрипции | Brownell et al., 1996; Kuo et al., 1996

K14ac | Hs, Dm TAF„230 TAF„250 | Активация транскрипции | Mizzen et al., 1996

K14ac | Hs | p300 | Активация транскрипции | Schiltz et al., 1999

K14ac | Hs | PCAF | Активация транскрипции | Schiltz et al., 1999

K14ac | Mm | SRC1 | Коактиватор ядерного рецептора | Spencer et al., 1997

R17me | Hs, Mm | CARM1 | Активация транскрипции (в сочетании с ацетилированием H3K18/23) | Chen et al., 1999; Schurter et al., 2001; Bauer et al., 2002; Daujat et al., 2002

K18ac | Sc | SAGA Ada Активация транскрипции | Grant etal., 1999

K18ac | Hs | p300 | Активация транскрипции | Schiltz etal., 1999

K18ac | Hs | СВР | Активация транскрипции (в сочетании с метилированием H3R17) | Daujat et al., 2002

K23ac | Sc | SAGA | Активация транскрипции | Grant etal., 1999

K23ac | Hs | СВР | Активация транскрипции (в сочетании с метилированием H3R17) | Daujat et al., 2002

R26me | Hs | CARM1 | Сайт метилирования in vitro | Chen et al., 1999; Schurter et al., 2001

K27me | Hs, Dm | E(z)/EZH2-me3 | Polycomb-репрессия Инактивация Х-хромосомы ранних В-клеток | Cao et al., 2002; Czermin et al., 2002; Kuzmichev et al., 2002; Muller et al., 2002; Su et al., 2003

S28ph | Hs | Aurora-В | Конденсация митотических хромосом | Goto et al., 1999, 2002

S28ph | Hs | MSK1 | UVB-индуцированное фосфорилирование | Zhong et al., 2001

K36me | Sc | Set2-me2 | Репрессия генов | Strahl et al., 2002; Kizer et al., 2005

K36me | Nc | Set2-me2 | Активация транскрипции | Adhvaryu et al., 2005

K36me | Sp | Set2-me2 | Элонгация транскрипции | Morris et al., 2005

K79me | Sc. Hs | Dot! /DOT! L-me2 | Теломерный сайленсинг, пахитенный checkpoint | Feng et al., 2002; Lacoste et al., 2002; Ng et al., 2002; van Leeuwen et al., 2002

Дополнительные модификации H3:

K14me, K23me, K27ас, T32ph, K37me, K56me, K64me, K115ас, K118ас, K118me, K122ас, R128me (Hyland et al., 2005).

Сайт | Модель | Энзим | Функция | Ссылка

S1ph | Hs, Sc | Casein kinase II | Реакция на повреждение ДНК | Ruiz-Carrillo et al., 1975; Cheung et al., 2005; van Attikum and Gasser, 2005

R3me | Hs, Sc | PRMT1 | Активация транскрипции | H. Wang et al., 2001 a,b

K5ac | It, Dm, Hs | Hatl | Откладка гистонов | Sobeletal., 1995; Parthunetal., 1996; Taplicket al., 1998; Turner, 2000; Kruhlak et al., 2001

K5ac | Sc | Esa1/NluA4 | Прогрессия по клеточному циклу | Smith etal., 1998; Allard etal., 1999; Clarke et al., 1999; Bird et al., 2002; Miranda et al., 2006

K5ac | Hs, Mm | ATF2 | Сиквенс-специфичный TF | Kawasaki et al., 2000a

K5ac | Hs p300 | Активация транскрипции | Turner and Fellows, 1989; Schiltz et al., 1999

K8ac | Hs, Mm | Y-ATF2 | Исключен из Xi. Сиквенс-специфичный транскрипционный фактор | Jeppesenetal., 1993: Choyetal., 2001; Kruhlak et al., 2001; Kawasaki et al., 2000b

K5ac | Hs | PCAF/p300 | Активация транскрипции | Turner and Fellows, 1989; Schiltz et al., 1999

К12ас | Sc, Hs | Hatl | Исключен из Xi. Откладка гистонов | Turner and Fellows, 1989; Jeppesenetal., 1993; Kleffetal., 1995; Sobeletal., 1995; Parthunetal., 1996; Chang et al., 1997; Kruhlak et al., 2001

К12ас | Sc | NuA4 | Прогрессия в митозе и в мейозе | Choy et al., 2001

K12bio | Hs | HCS Биотинидаза | Уменьшение реакции на двунитевые разрывы ДНК. Влияние на пролиферацию клеток | Stanley et al., 2001; Kothapalli et al., 2005a,b

К16ас | Mm | ? | Исключен из Xi. Зависимое от клеточного цикла ацетилирование | Ieppesen et al., 1993; Taplick et al., 1998

К16ас | Dm | MOF | Транскрипционная ап-регуляция мужской Х-хромосомы | Akhtar and Becker, 2000; Hsu et al., 2000

К16ас | Hs, Mm | ATF2 | Сиквенс-специфичный транскрипционный фактор | Turner and Fellows, 1989; Kawasaki et al., 2000a; Turner, 2000; Kruhlak et al., 2001; Vacquero et al., 2004

K20me | Mm, Dm | Suv4-20hl-me2, me3 Suv4-20h2-me2, me3 | Сайленсинг генов | Schotta et al., 2004

K20me | Hs, Dm | Pr-SET7/Set8-me1 | Транскрипционный сайленсинг, митотическая конденсация | Fang et al., 2002; Nishioka et al., 2002b; Rice et al., 2002

K20me | Dm | Ash1-me2 | Trithorax-активация в сочетании с метилированием H3K4 и H3K9 | Beisei et al., 2002

K59me | Sc | Формирование «молчащего» хроматина | Zhang et al., 2003

K59me | Su Hs | SUMO-1 SUMO-3 | Репрессия транскрипции | Shiio and Eisenman, 2003

Дополнительные модификации Н4:

K12me, S47ph, K31ubl, К59me, К77ас, К79ас, К79me, K91ub1, R92me (Hyland et al., 2005).

Сайт | Модель | Энзим | Функция | Ссылка

E2ar1 | Rn | PARP-1 | Участие в нейротрофной активности | Ogata et al., 1980; Visochek et al., 2005

T10ph | Hs | — | Митоз-специфичная транскрипционная активация Н1b | Chadee et al., 1995; Garcia et al., 2004; Sarg et al., 2006

E14ar1 | Rn | PARP-1 | Участие в нейротрофной активности Ogata et al., 1980; Visochek et al., 2005

S17ph | Hs | — | Интерфаза-специфичная транскрипционная активация Н1b Garcia et al., 2004; Chadee et al., 1995; Sarg et al., 2006

K26me | Hs | EZH2-me2 | Опосредует связывание НР1 Kuzmichev et al., 2004; Daujat et al., 2005

S27ph | Hs | — | Блокирует связывание НР1 | Garcia et al., 2004; Daujat et al., 2005

T137ph | Hs | — | Митоз-специфичная транскрипционная активация H1b | Chadee et al., 1995; Garcia et al., 2004; Sarg et al., 2006

T154ph | Hs | — | Митоз-специфичная транскрипционная активация H1b | Chadee et al., 1995; Garcia et al., 2004; Sarg et al., 2006

S172ph | Hs | — | Интерфаза-специфичная транскрипционная активация H1b | Chadee et al., 1995; Garcia et al., 2004; Sarg et al., 2006

S188ph | Hs | — | Интерфаза-специфичная транскрипционная активация H1b | Chadee et al., 1995; Garcia et al., 2004; Sarg et al., 2006

K213arl | Rn | PARP-1 | Участие в нейротрофной активности | Ogata et al., 1980; Visochek et al, 2005

Дополнительные модификации Н1:

К26ас, E114arn, убиквитинирование Н1, карбонилирование HI (Pham and Sauer, 2000: Rouleau et al., 2004).

Сайт | Модель | Энзим | Функция | Ссылка

S139ph | Hs, Sc, Dm. XI | ATM DNA-PK | Репарация ДНК. Связана с М-фазой. Известна также как ?Н2АХ | Rogakou et al., 1998,1999; Burma et al., 2001 ; Stiff et al., 2004; Ichijima et al., 2005; Mukheijee et al., 2006

Дополнительные модификации H2AX: T136ph.

Сайт | Модель | Энзим | Функция | Ссылка

K17me | Hs | — | — | Chu et al., 2006

K115ub1 | Hs | — | — | Ogawa et al., 2005; Chu et al., 2006

K122me | Hs | — | — | Chu et al., 2006

T128ph | Hs | — | — | Chu et al., 2006

K238me | Hs | — | — | Chu et al., 2006

Сайт | Модель | Энзим | Функция | Ссылка

K4me | Dm | me1, me2, me3 | Активация транскрипции | McKittrick et al., 2004

K9me | Dm | me1, me2 | Репрессия транскрипции | McKittnck et al., 2004

K9ac | Dm, Hs | — | Активация транскрипции | McKittrick et al., 2004; Hake and Allis, 2006

K14me | Dm | me1, me2 | McKittrick et al., 2004

K14ac | Dm, Hs | — | Активация транскрипции | McKittrick et al., 2004; Hake and Allis, 2006

K18aс | Hs | — | Активация транскрипции | Hake and Allis, 2006

K23ac | Hs | — | Активация транскрипции | Hake and Allis, 2006

K27me | Dm | me1, me2, me3 | Репрессия транскрипции | McKittrick et al., 2004

K36me | Dm, Hs | me1, me2, me3 | Активация транскрипции | McKittrick et al., 2004; Hake and Allis, 2006

K37me | Dm | me1, me2 | — | McKittrick et al., 2004

K79me | Dm, Hs | me1, me2 | Активация транскрипции | McKittrick et al., 2004; Hake and Allis, 2006

S31ph | Млекопитающие | — | Митоз-специфичное фосфорилирование | Hake et al., 2005

Сайт | Модель | Энзим | Функция | Ссылка

S7ph | Hs | — | Митоз | Zeitlin et al., 2001

Дополнительные модификации CENP-A: S17ph.

(An) Aspergillus nidulans, (At) Arabidopsis thaliana, (Ce) Caenorhabditis elegans, (Dm) Drosophila melanogaster, (Hs) Homo sapiens, (Mm) Mus musculus, (Nc) Neurospora crassa, (Rn) Rattus norvegicus, (Sc) Saccharomyces cerevisiae, (Sp) Schizosaccharomyces pombe, (Tt) Tetrahymena thermophila, (Xl) Xenopus laevis.

Модификации гистонов — по номенклатуре, предложенной Тернером (Turner, 2005).

В таблицах перечисляются все известные модификации гистонов и известные энзимы с первичными ссылками (до мая, 2006 г.). Дополнительные модификации с пока еще неизвестными функциями перечислены под таблицами. Эти модификации взяты из разных источников: обзорных статей, информации от Abeam Cambridge, UK и Upstate Charlottesville, USA, а также из неопубликованных данных лаборатории Рейнберга (Reinberg).

Эта таблица была составлена на основе оригинального варианта, взятого из (Lachner et al., 2003) и значительно расширенного Roopsha Sengupta и Mario Richter (Лаборатория Jenuwein, IMP Vienna). Д-р Patrick Trojer (Лаборатория Reinberg, HHMI, New Jersey) проверил содержимое таблиц.

Эта таблица была проверена и дополнена информацией, предоставленной д-ром Steven Gray (Dept, of Clinical Medicine, Institute of Molecular Medicine, St. James Hospital, Dublin).

Adhvaryu K.K., Morris S.A., Strahl B.D., and Selker E.U., 2005. Methylation of histone H3 lysine 36 is required for normal development in Neurospora crassa. Eukaryot. Cell 4: 1455-1464.

Ahn S.H., Cheung W.L., Hsu J.Y., Diaz R.L., Smith M.M., and Allis C.D., 2005. Sterile, 20 kinase phosphorylates histone H2B at serine 10 during hydrogen peroxide-induced apoptosis in S. cerevisiae. Cell 120: 25-36.

Aihara H., Nakagawa T., Yasui K., Ohta T., Hirose S., Dhomae N., Takio K., Kaneko M., Takeshima Y., Muramatsu M., and Ito T., 2004. Nucleosomal histone kinase-1 phosphorylates H2A Thr 119 during mitosis in the early Drosophila embryo. Genes Dev. 18: 877-888.

Ajiro K., 2000. Histone H2B phosphorylation in mammalian apop-totic cells. An association with DNA fragmentation. J. Biol. Chem. 275: 439-443.

Akhtar A., and Becker P.B., 2000. Activation of transcription through histone H4 acetylation by MOF, an acetyltransferase essential for dosage compensation in Drosophila. Mol. Cell 5: 367-375.

Allard S., Utley R.T., Savard J., Clarke A., Grant P., Brandi C.J., Pillus L., Workman J.L., and Cote J., 1999. NuA4, an essential transcription adaptor/histone H4 acetyltransferase complex containing Esalp and the ATM-related cofactor Tralp. EMBO J. 18: 5108-5119.

Anest V., Hanson J.L., Cogswell P.C, Steinbrecher K.A., Strahl B.D., and Baldwin A.S., 2003. A nucleosomal function for IkB kinase-a in NF-KB-dependent gene expression. Nature 423: 659-663.

Bannister A.J., Zegerman P., Partridge J.E., Miska E.A., Thomas J.O., Allshire R.C., and Kouzarides T., 2001. Selective recognition of methylated lysine 9 on histone H3 by the HP1 chromo domain. Nature 410: 120-124.

Bauer U.M., Daujat S., Nielsen S.J., Nightingale K., and Kouzarides T., 2002. Methylation at arginine 17 of histone H3 is linked to gene activation. EMBO Rep. 3: 39-44.

Beisel C, Imhof A., Greene J., Kremmer E., and Sauer F., 2002. Histone methylation by the Drosophila epigenetic transcriptional regulator Ash 1. Nature 419: 857-862.

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