サイクリン依存性キナーゼ8とは？わかりやすく解説

CDK8
PDBに登録されている構造
PDB	オルソログ検索: RCSB PDBe PDBj
PDBのIDコード一覧
	3RGF, 4F6S, 4F6U, 4F6W, 4F70, 4F7J, 4F7L, 4F7N, 4F7S, 4G6L, 4CRL, 5HBH, 5CEI, 5FGK, 5HBE, 5BNJ, 5HBJ, 5HVY, 5HNB, 5I5Z
識別子
記号	CDK8, K35, cyclin-dependent kinase 8, cyclin dependent kinase 8, IDDHBA
外部ID	OMIM: 603184 MGI: 1196224 HomoloGene: 55565 GeneCards: CDK8
遺伝子の位置 (ヒト)
染色体	13番染色体 (ヒト)
終点	26,405,238 bp
遺伝子の位置 (マウス)
染色体	5番染色体 (マウス)
終点	146,239,684 bp
RNA発現パターン
	さらなる参照発現データ
遺伝子オントロジー
分子機能	• トランスフェラーゼ活性; • protein kinase activity; • ヌクレオチド結合; • キナーゼ活性; • 血漿タンパク結合; • RNA polymerase II CTD heptapeptide repeat kinase activity; • ATP binding; • protein serine/threonine kinase activity; • cyclin-dependent protein serine/threonine kinase activity;
細胞の構成要素	• メディエーター; • 核質; • 細胞核; • 核小体; • 高分子複合体;
生物学的プロセス	• regulation of transcription, DNA-templated; • リン酸化; • タンパク質リン酸化; • transcription initiation from RNA polymerase II promoter; • positive regulation of transcription by RNA polymerase II; • transcription, DNA-templated; • regulation of mitotic cell cycle; • regulation of cell cycle;
	出典:Amigo / QuickGO
オルソログ
	1024
	264064
	ENSG00000132964
	ENSMUSG00000029635
	P49336,A0A0D9SEP3
	Q8R3L8
	NM_001260; NM_001318368; NM_001346501
	NM_153599; NM_181570; NM_001359990; NM_001359991
	NP_001251; NP_001305297; NP_001333430
	NP_705827; NP_001346919; NP_001346920
	ウィキデータ
閲覧/編集ヒト	閲覧/編集マウス

サイクリン依存性キナーゼ8（英: cyclin dependent kinase 8、CDK8）は、ヒトではCDK8遺伝子によってコードされている酵素（プロテインキナーゼ）である^[5]^[6]。

機能

CDK8遺伝子にコードされているCDK8タンパク質は、サイクリン依存性キナーゼ（CDK）ファミリーの一員である。CDK8とサイクリンCはメディエーター複合体（英語版）へ結合し、いくつかの機構で転写を調節する。CDK8はいくつかの転写因子に対して結合やリン酸化を行うことで、転写因子の機能を活性化もしくは阻害的な影響を及ぼす場合がある^[7]^[8]。具体的には、CDK8はNotchの細胞内ドメイン^[9]、SREBP^[10]、STAT1のセリン727番^[11]をリン酸化することが知られている。また、CDK8はメディエーター複合体のテールモジュールのサブユニットのターンオーバーに影響を及ぼすことで、転写活性化を阻害する^[12]^[13]。さらに、CDK8はRNAポリメラーゼIIのメディエーターへの結合にも影響を及ぼす^[14]^[15]。

臨床的意義

CDK8は大腸がんにおいてがん遺伝子としてはたらく。ヒトの大腸がんではCDK8遺伝子が増幅し、β-カテニンを介した転写が活性化されることで結腸における腫瘍形成が駆動されている場合がある^[16]。一方で、CDK8は全ての細胞種で発がん性を有するわけではなく、Notch経路やEGFR経路においてはがん抑制因子として作用している可能性がある。具体的には、CDK8はNotchの細胞内ドメインのターンオーバーを促進し^[9]、またC. elegansではEGFRシグナルによって駆動される細胞運命決定を阻害することが示されている^[13]。さらに、CDK8はがん抑制タンパク質p53によって媒介される転写活性化を促進することから、がん抑制に重要な役割を果たしている可能性が示唆されている^[17]。さまざまな組織におけるCDK8阻害の影響を明らかにするためにはさらなる研究が必要であり、現時点ではCDK8を標的としたがん治療薬のヒトでの試験は行われていない。一方で、コルチスタチンA（英語版）と呼ばれる天然物はCDK8とCDK19に対する強力な選択的阻害剤となることが示されている^[18]。急性骨髄性白血病（AML）の動物モデルでは、コルチスタチンAはCDK8とCDK19を阻害することで、CEBPA（英語版）やIRF8（英語版）といった細胞のアイデンティティに関わる遺伝子を含むスーパーエンハンサー（英語版）関連遺伝子の選択的かつ不均衡なアップレギュレーションを引き起こし、細胞増殖を抑制して抗がん作用を発揮することが示されている^[18]。

CDK8のキナーゼドメインのATP結合ポケットの変異と関連した、常染色体顕性型症候群が記載されている^[19]。この症候群の臨床像としては、脳梁の欠損、軽度から中等度の知的障害、筋緊張低下（英語版）、てんかん発作、聴覚もしくは視覚の障害、行動障害、さまざまな顔面奇形、先天性心疾患、直腸肛門奇形が挙げられる^[19]。

相互作用

CDK8は次に挙げる因子と相互作用することが示されている。

CCNC^[5]^[20]^[21]^[22]^[23]
CREBBP^[22]
CRSP3（英語版）^[20]^[21]
MED1（英語版）^[20]^[24]
MED12（英語版）^[20]^[24]
MED14（英語版）^[21]^[24]
MED16（英語版）^[20]^[24]
MED17（英語版）^[20]^[24]
MED21（英語版）^[20]^[22]^[24]^[25]
MED24（英語版）^[20]^[24]
MED26（英語版）^[26]
MED6（英語版）^[20]^[21]^[24]^[27]
Notch^[9]
POLR2A（英語版）^[22]
SMARCB1（英語版）^[22]
STAT1^[11]
SREBP^[10]

出典

^ ^a ^b ^c GRCh38: Ensembl release 89: ENSG00000132964 - Ensembl, May 2017
^ ^a ^b ^c GRCm38: Ensembl release 89: ENSMUSG00000029635 - Ensembl, May 2017
^ Human PubMed Reference:
^ Mouse PubMed Reference:
^ ^a ^b Tassan JP, Jaquenoud M, Léopold P, Schultz SJ, Nigg EA (Sep 1995). “Identification of human cyclin-dependent kinase 8, a putative protein kinase partner for cyclin C”. Proceedings of the National Academy of Sciences of the United States of America 92 (19): 8871–5. Bibcode: 1995PNAS...92.8871T. doi:10.1073/pnas.92.19.8871. PMC 41069. PMID 7568034.
^ “Entrez Gene: CDK8 cyclin-dependent kinase 8”. 2025年11月9日閲覧。
^ Nemet J, Jelicic B, Rubelj I, Sopta M (Feb 2014). “The two faces of Cdk8, a positive/negative regulator of transcription”. Biochimie 97: 22–7. doi:10.1016/j.biochi.2013.10.004. PMID 24139904.
^ Poss ZC, Ebmeier CC, Taatjes DJ (2013). “The mediator complex and transcription regulation”. Critical Reviews in Biochemistry and Molecular Biology 48 (6): 575–608. doi:10.3109/10409238.2013.840259. PMC 3852498. PMID 24088064.
^ ^a ^b ^c Fryer CJ, White JB, Jones KA (Nov 2004). “Mastermind recruits CycC:CDK8 to phosphorylate the Notch ICD and coordinate activation with turnover”. Molecular Cell 16 (4): 509–20. doi:10.1016/j.molcel.2004.10.014. PMID 15546612.
^ ^a ^b Zhao X, Feng D, Wang Q, Abdulla A, Xie XJ, Zhou J, Sun Y, Yang ES, Liu LP, Vaitheesvaran B, Bridges L, Kurland IJ, Strich R, Ni JQ, Wang C, Ericsson J, Pessin JE, Ji JY, Yang F (Jul 2012). “Regulation of lipogenesis by cyclin-dependent kinase 8-mediated control of SREBP-1”. The Journal of Clinical Investigation 122 (7): 2417–27. doi:10.1172/JCI61462. PMC 3386818. PMID 22684109.
^ ^a ^b Bancerek J, Poss ZC, Steinparzer I, Sedlyarov V, Pfaffenwimmer T, Mikulic I, Dölken L, Strobl B, Müller M, Taatjes DJ, Kovarik P (Feb 2013). “CDK8 kinase phosphorylates transcription factor STAT1 to selectively regulate the interferon response” (英語). Immunity 38 (2): 250–62. doi:10.1016/j.immuni.2012.10.017. PMC 3580287. PMID 23352233.
^ Gonzalez D, Hamidi N, Del Sol R, Benschop JJ, Nancy T, Li C, Francis L, Tzouros M, Krijgsveld J, Holstege FC, Conlan RS (Feb 2014). “Suppression of Mediator is regulated by Cdk8-dependent Grr1 turnover of the Med3 coactivator”. Proceedings of the National Academy of Sciences of the United States of America 111 (7): 2500–5. Bibcode: 2014PNAS..111.2500G. doi:10.1073/pnas.1307525111. PMC 3932902. PMID 24550274.
^ ^a ^b Grants JM, Ying LT, Yoda A, You CC, Okano H, Sawa H, Taubert S (Feb 2016). “The Mediator Kinase Module Restrains Epidermal Growth Factor Receptor Signaling and Represses Vulval Cell Fate Specification in Caenorhabditis elegans”. Genetics 202 (2): 583–99. doi:10.1534/genetics.115.180265. PMC 4788237. PMID 26715664.
^ Taatjes DJ, Näär AM, Andel F, Nogales E, Tjian R (Feb 2002). “Structure, function, and activator-induced conformations of the CRSP coactivator”. Science 295 (5557): 1058–62. Bibcode: 2002Sci...295.1058T. doi:10.1126/science.1065249. PMID 11834832.
^ Tsai KL, Sato S, Tomomori-Sato C, Conaway RC, Conaway JW, Asturias FJ (May 2013). “A conserved Mediator-CDK8 kinase module association regulates Mediator-RNA polymerase II interaction”. Nature Structural & Molecular Biology 20 (5): 611–9. doi:10.1038/nsmb.2549. PMC 3648612. PMID 23563140.
^ Firestein R, Bass AJ, Kim SY, Dunn IF, Silver SJ, Guney I, Freed E, Ligon AH, Vena N, Ogino S, Chheda MG, Tamayo P, Finn S, Shrestha Y, Boehm JS, Jain S, Bojarski E, Mermel C, Barretina J, Chan JA, Baselga J, Tabernero J, Root DE, Fuchs CS, Loda M, Shivdasani RA, Meyerson M, Hahn WC (Sep 2008). “CDK8 is a colorectal cancer oncogene that regulates beta-catenin activity”. Nature 455 (7212): 547–51. Bibcode: 2008Natur.455..547F. doi:10.1038/nature07179. PMC 2587138. PMID 18794900.
^ Donner AJ, Szostek S, Hoover JM, Espinosa JM (Jul 2007). “CDK8 is a stimulus-specific positive coregulator of p53 target genes”. Molecular Cell 27 (1): 121–33. doi:10.1016/j.molcel.2007.05.026. PMC 2936241. PMID 17612495.
^ ^a ^b Pelish HE, Liau BB, Nitulescu II, Tangpeerachaikul A, Poss ZC, Da Silva DH, Caruso BT, Arefolov A, Fadeyi O, Christie AL, Du K, Banka D, Schneider EV, Jestel A, Zou G, Si C, Ebmeier CC, Bronson RT, Krivtsov AV, Myers AG, Kohl NE, Kung AL, Armstrong SA, Lemieux ME, Taatjes DJ, Shair MD (Oct 2015). “Mediator kinase inhibition further activates super-enhancer-associated genes in AML”. Nature 526 (7572): 273–6. Bibcode: 2015Natur.526..273P. doi:10.1038/nature14904. PMC 4641525. PMID 26416749.
^ ^a ^b Calpena, Eduardo; Hervieu, Alexia; Kaserer, Teresa; Swagemakers, Sigrid M. A.; Goos, Jacqueline A. C.; Popoola, Olajumoke; Ortiz-Ruiz, Maria Jesus; Barbaro-Dieber, Tina et al. (2019-04-04). “De Novo Missense Substitutions in the Gene Encoding CDK8, a Regulator of the Mediator Complex, Cause a Syndromic Developmental Disorder”. American Journal of Human Genetics 104 (4): 709–720. doi:10.1016/j.ajhg.2019.02.006. ISSN 1537-6605. PMC 6451695. PMID 30905399.
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ Kang YK, Guermah M, Yuan CX, Roeder RG (Mar 2002). “The TRAP/Mediator coactivator complex interacts directly with estrogen receptors alpha and beta through the TRAP220 subunit and directly enhances estrogen receptor function in vitro”. Proceedings of the National Academy of Sciences of the United States of America 99 (5): 2642–7. Bibcode: 2002PNAS...99.2642K. doi:10.1073/pnas.261715899. PMC 122401. PMID 11867769.
^ ^a ^b ^c ^d Wang G, Cantin GT, Stevens JL, Berk AJ (Jul 2001). “Characterization of mediator complexes from HeLa cell nuclear extract”. Molecular and Cellular Biology 21 (14): 4604–13. doi:10.1128/MCB.21.14.4604-4613.2001. PMC 87123. PMID 11416138.
^ ^a ^b ^c ^d ^e Cho H, Orphanides G, Sun X, Yang XJ, Ogryzko V, Lees E, Nakatani Y, Reinberg D (Sep 1998). “A human RNA polymerase II complex containing factors that modify chromatin structure”. Molecular and Cellular Biology 18 (9): 5355–63. doi:10.1128/MCB.18.9.5355. PMC 109120. PMID 9710619.
^ Zhang Y, Iratni R, Erdjument-Bromage H, Tempst P, Reinberg D (May 1997). “Histone deacetylases and SAP18, a novel polypeptide, are components of a human Sin3 complex”. Cell 89 (3): 357–64. doi:10.1016/s0092-8674(00)80216-0. PMID 9150135.
^ ^a ^b ^c ^d ^e ^f ^g ^h Ito M, Yuan CX, Malik S, Gu W, Fondell JD, Yamamura S, Fu ZY, Zhang X, Qin J, Roeder RG (Mar 1999). “Identity between TRAP and SMCC complexes indicates novel pathways for the function of nuclear receptors and diverse mammalian activators”. Molecular Cell 3 (3): 361–70. doi:10.1016/s1097-2765(00)80463-3. PMID 10198638.
^ Suñé C, Hayashi T, Liu Y, Lane WS, Young RA, Garcia-Blanco MA (Oct 1997). “CA150, a nuclear protein associated with the RNA polymerase II holoenzyme, is involved in Tat-activated human immunodeficiency virus type 1 transcription”. Molecular and Cellular Biology 17 (10): 6029–39. doi:10.1128/MCB.17.10.6029. PMC 232452. PMID 9315662.
^ Sato S, Tomomori-Sato C, Parmely TJ, Florens L, Zybailov B, Swanson SK, Banks CA, Jin J, Cai Y, Washburn MP, Conaway JW, Conaway RC (Jun 2004). “A set of consensus mammalian mediator subunits identified by multidimensional protein identification technology”. Molecular Cell 14 (5): 685–91. doi:10.1016/j.molcel.2004.05.006. PMID 15175163.
^ Yang F, DeBeaumont R, Zhou S, Näär AM (Feb 2004). “The activator-recruited cofactor/Mediator coactivator subunit ARC92 is a functionally important target of the VP16 transcriptional activator”. Proceedings of the National Academy of Sciences of the United States of America 101 (8): 2339–44. Bibcode: 2004PNAS..101.2339Y. doi:10.1073/pnas.0308676100. PMC 356952. PMID 14983011.

外部リンク

CDK8 protein, human - MeSH・アメリカ国立医学図書館・生命科学用語シソーラス（英語）

[refGRCh38Ensembl-1] GRCh38: Ensembl release 89: ENSG00000132964 - Ensembl, May 2017

[refGRCm38Ensembl-2] GRCm38: Ensembl release 89: ENSMUSG00000029635 - Ensembl, May 2017

[3] Human PubMed Reference:

[4] Mouse PubMed Reference:

[pmid7568034-5] Tassan JP, Jaquenoud M, Léopold P, Schultz SJ, Nigg EA (Sep 1995). “Identification of human cyclin-dependent kinase 8, a putative protein kinase partner for cyclin C”. Proceedings of the National Academy of Sciences of the United States of America 92 (19): 8871–5. Bibcode: 1995PNAS...92.8871T. doi:10.1073/pnas.92.19.8871. PMC 41069. PMID 7568034.

[entrez-6] “Entrez Gene: CDK8 cyclin-dependent kinase 8”. 2025年11月9日閲覧。

[:0-7] Nemet J, Jelicic B, Rubelj I, Sopta M (Feb 2014). “The two faces of Cdk8, a positive/negative regulator of transcription”. Biochimie 97: 22–7. doi:10.1016/j.biochi.2013.10.004. PMID 24139904.

[:1-8] Poss ZC, Ebmeier CC, Taatjes DJ (2013). “The mediator complex and transcription regulation”. Critical Reviews in Biochemistry and Molecular Biology 48 (6): 575–608. doi:10.3109/10409238.2013.840259. PMC 3852498. PMID 24088064.

[Fryer_CJ_2004-9] Fryer CJ, White JB, Jones KA (Nov 2004). “Mastermind recruits CycC:CDK8 to phosphorylate the Notch ICD and coordinate activation with turnover”. Molecular Cell 16 (4): 509–20. doi:10.1016/j.molcel.2004.10.014. PMID 15546612.

[ReferenceA-10] Zhao X, Feng D, Wang Q, Abdulla A, Xie XJ, Zhou J, Sun Y, Yang ES, Liu LP, Vaitheesvaran B, Bridges L, Kurland IJ, Strich R, Ni JQ, Wang C, Ericsson J, Pessin JE, Ji JY, Yang F (Jul 2012). “Regulation of lipogenesis by cyclin-dependent kinase 8-mediated control of SREBP-1”. The Journal of Clinical Investigation 122 (7): 2417–27. doi:10.1172/JCI61462. PMC 3386818. PMID 22684109.

[Bancerek_J_2012-11] Bancerek J, Poss ZC, Steinparzer I, Sedlyarov V, Pfaffenwimmer T, Mikulic I, Dölken L, Strobl B, Müller M, Taatjes DJ, Kovarik P (Feb 2013). “CDK8 kinase phosphorylates transcription factor STAT1 to selectively regulate the interferon response” (英語). Immunity 38 (2): 250–62. doi:10.1016/j.immuni.2012.10.017. PMC 3580287. PMID 23352233.

[:2-12] Gonzalez D, Hamidi N, Del Sol R, Benschop JJ, Nancy T, Li C, Francis L, Tzouros M, Krijgsveld J, Holstege FC, Conlan RS (Feb 2014). “Suppression of Mediator is regulated by Cdk8-dependent Grr1 turnover of the Med3 coactivator”. Proceedings of the National Academy of Sciences of the United States of America 111 (7): 2500–5. Bibcode: 2014PNAS..111.2500G. doi:10.1073/pnas.1307525111. PMC 3932902. PMID 24550274.

[ReferenceB-13] Grants JM, Ying LT, Yoda A, You CC, Okano H, Sawa H, Taubert S (Feb 2016). “The Mediator Kinase Module Restrains Epidermal Growth Factor Receptor Signaling and Represses Vulval Cell Fate Specification in Caenorhabditis elegans”. Genetics 202 (2): 583–99. doi:10.1534/genetics.115.180265. PMC 4788237. PMID 26715664.

[:3-14] Taatjes DJ, Näär AM, Andel F, Nogales E, Tjian R (Feb 2002). “Structure, function, and activator-induced conformations of the CRSP coactivator”. Science 295 (5557): 1058–62. Bibcode: 2002Sci...295.1058T. doi:10.1126/science.1065249. PMID 11834832.

[:4-15] Tsai KL, Sato S, Tomomori-Sato C, Conaway RC, Conaway JW, Asturias FJ (May 2013). “A conserved Mediator-CDK8 kinase module association regulates Mediator-RNA polymerase II interaction”. Nature Structural & Molecular Biology 20 (5): 611–9. doi:10.1038/nsmb.2549. PMC 3648612. PMID 23563140.

[:6-16] Firestein R, Bass AJ, Kim SY, Dunn IF, Silver SJ, Guney I, Freed E, Ligon AH, Vena N, Ogino S, Chheda MG, Tamayo P, Finn S, Shrestha Y, Boehm JS, Jain S, Bojarski E, Mermel C, Barretina J, Chan JA, Baselga J, Tabernero J, Root DE, Fuchs CS, Loda M, Shivdasani RA, Meyerson M, Hahn WC (Sep 2008). “CDK8 is a colorectal cancer oncogene that regulates beta-catenin activity”. Nature 455 (7212): 547–51. Bibcode: 2008Natur.455..547F. doi:10.1038/nature07179. PMC 2587138. PMID 18794900.

[:7-17] Donner AJ, Szostek S, Hoover JM, Espinosa JM (Jul 2007). “CDK8 is a stimulus-specific positive coregulator of p53 target genes”. Molecular Cell 27 (1): 121–33. doi:10.1016/j.molcel.2007.05.026. PMC 2936241. PMID 17612495.

[:5-18] Pelish HE, Liau BB, Nitulescu II, Tangpeerachaikul A, Poss ZC, Da Silva DH, Caruso BT, Arefolov A, Fadeyi O, Christie AL, Du K, Banka D, Schneider EV, Jestel A, Zou G, Si C, Ebmeier CC, Bronson RT, Krivtsov AV, Myers AG, Kohl NE, Kung AL, Armstrong SA, Lemieux ME, Taatjes DJ, Shair MD (Oct 2015). “Mediator kinase inhibition further activates super-enhancer-associated genes in AML”. Nature 526 (7572): 273–6. Bibcode: 2015Natur.526..273P. doi:10.1038/nature14904. PMC 4641525. PMID 26416749.

[:8-19] Calpena, Eduardo; Hervieu, Alexia; Kaserer, Teresa; Swagemakers, Sigrid M. A.; Goos, Jacqueline A. C.; Popoola, Olajumoke; Ortiz-Ruiz, Maria Jesus; Barbaro-Dieber, Tina et al. (2019-04-04). “De Novo Missense Substitutions in the Gene Encoding CDK8, a Regulator of the Mediator Complex, Cause a Syndromic Developmental Disorder”. American Journal of Human Genetics 104 (4): 709–720. doi:10.1016/j.ajhg.2019.02.006. ISSN 1537-6605. PMC 6451695. PMID 30905399.

[pmid11867769-20] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ Kang YK, Guermah M, Yuan CX, Roeder RG (Mar 2002). “The TRAP/Mediator coactivator complex interacts directly with estrogen receptors alpha and beta through the TRAP220 subunit and directly enhances estrogen receptor function in vitro”. Proceedings of the National Academy of Sciences of the United States of America 99 (5): 2642–7. Bibcode: 2002PNAS...99.2642K. doi:10.1073/pnas.261715899. PMC 122401. PMID 11867769.

[pmid11416138-21] Wang G, Cantin GT, Stevens JL, Berk AJ (Jul 2001). “Characterization of mediator complexes from HeLa cell nuclear extract”. Molecular and Cellular Biology 21 (14): 4604–13. doi:10.1128/MCB.21.14.4604-4613.2001. PMC 87123. PMID 11416138.

[pmid9710619-22] Cho H, Orphanides G, Sun X, Yang XJ, Ogryzko V, Lees E, Nakatani Y, Reinberg D (Sep 1998). “A human RNA polymerase II complex containing factors that modify chromatin structure”. Molecular and Cellular Biology 18 (9): 5355–63. doi:10.1128/MCB.18.9.5355. PMC 109120. PMID 9710619.

[pmid9150135-23] Zhang Y, Iratni R, Erdjument-Bromage H, Tempst P, Reinberg D (May 1997). “Histone deacetylases and SAP18, a novel polypeptide, are components of a human Sin3 complex”. Cell 89 (3): 357–64. doi:10.1016/s0092-8674(00)80216-0. PMID 9150135.

[pmid10198638-24] ^ ^a ^b ^c ^d ^e ^f ^g ^h Ito M, Yuan CX, Malik S, Gu W, Fondell JD, Yamamura S, Fu ZY, Zhang X, Qin J, Roeder RG (Mar 1999). “Identity between TRAP and SMCC complexes indicates novel pathways for the function of nuclear receptors and diverse mammalian activators”. Molecular Cell 3 (3): 361–70. doi:10.1016/s1097-2765(00)80463-3. PMID 10198638.

[pmid9315662-25] Suñé C, Hayashi T, Liu Y, Lane WS, Young RA, Garcia-Blanco MA (Oct 1997). “CA150, a nuclear protein associated with the RNA polymerase II holoenzyme, is involved in Tat-activated human immunodeficiency virus type 1 transcription”. Molecular and Cellular Biology 17 (10): 6029–39. doi:10.1128/MCB.17.10.6029. PMC 232452. PMID 9315662.

[pmid15175163-26] Sato S, Tomomori-Sato C, Parmely TJ, Florens L, Zybailov B, Swanson SK, Banks CA, Jin J, Cai Y, Washburn MP, Conaway JW, Conaway RC (Jun 2004). “A set of consensus mammalian mediator subunits identified by multidimensional protein identification technology”. Molecular Cell 14 (5): 685–91. doi:10.1016/j.molcel.2004.05.006. PMID 15175163.

[pmid14983011-27] Yang F, DeBeaumont R, Zhou S, Näär AM (Feb 2004). “The activator-recruited cofactor/Mediator coactivator subunit ARC92 is a functionally important target of the VP16 transcriptional activator”. Proceedings of the National Academy of Sciences of the United States of America 101 (8): 2339–44. Bibcode: 2004PNAS..101.2339Y. doi:10.1073/pnas.0308676100. PMC 356952. PMID 14983011.

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サイクリン依存性キナーゼ8とは？わかりやすく解説