植物中介因子复合体亚基尾部模块研究进展
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摘要
中介因子复合体是当今分子生物学领域中热门的研究方向。复合体被分为头部、中部、尾部和细胞周期素依赖性激酶4个模块部分。其中尾部模块能直接和转录激活因子结合并调控该基因的表达,是中介因子复合体发挥调控功能的重要模块。本文系统阐述了植物体内尾部模块中各个亚基(主要为MED16、MED25)的调控作用机制,并进行了展望,旨在为亚基新功能的开发提供参考。
Intermediate factor complex is a hot research direction in the field of molecular biology.The complex is divided into four modules: head,middle,tail and cyclin-dependent kinase.The tail module can directly bind to transcription activator and regulate the expression of the gene.It is an important module for the mediator complex to play a regulatory role.In this paper,the regulatory mechanism of each subunit(mainly MED16 and MED25) in the tail module of plant was systematically described,and the prospect was given,in order to provide reference for the development of new functions of the subunit.
Intermediate factor complex is a hot research direction in the field of molecular biology.The complex is divided into four modules: head,middle,tail and cyclin-dependent kinase.The tail module can directly bind to transcription activator and regulate the expression of the gene.It is an important module for the mediator complex to play a regulatory role.In this paper,the regulatory mechanism of each subunit(mainly MED16 and MED25) in the tail module of plant was systematically described,and the prospect was given,in order to provide reference for the development of new functions of the subunit.
引文
[1] Jeronimo C,Robert F.The mediator complex:At the nexus of RNA polymerase II transcription.[J].Trends in Cell Biology,2017,27(10):765-783.
[2] Schilbach S,Hantsche M,Tegunov D,et al.Structures of transcription pre-initiation complex with TFIIH and Mediator[J].Nature,2017,551:204-209.
[3] Nozawa K,Schneider T R,Cramer P.Core Mediator structure at 3.4 ? extends model of transcription initiation complex[J].Nature,2017:248-251.
[4] Rival P,Press M O,Bale J,et al.The conserved PFT1 tandem repeat is crucial for proper flowering in Arabidopsis thaliana[J].Genetics,2014,198(2):747-754.
[5] Alonso José M,Stepanova A N,Leisse T J,et al.Genome-wide insertional mutagenesis of Arabidopsis thaliana[J].Science,2003,301(5633):653-657.
[6] Samanta S,Thakur J K.Characterization of mediator complex and its associated proteins from rice[J].Methods in Molecular Biology,2017,1629:123.
[7] YangY,Li L,Qu,Li Jia.Plant Mediator complex and its critical functions in transcription regulation[J].Journal of Integrative Plant Biology,2016,58(2):13.
[8] Plaschka C,Larivière,L,Wenzeck L,et al.Architecture of the RNA polymerase II-Mediator core initiation complex[J].Nature,2015,518(7539):376-380.
[9] Canet J V,Dobón A,Tornero P.Non-recognition-of-BTH4,an Arabidopsis mediator subunit homolog,is necessary for development and response to salicylic acid[J].Plant Cell,2012,24(10):4220-4235.
[10] KnightH,Mugford N G S G,Ulker B,et al.Identification of SFR6,a key component in cold acclimation acting post-translationally on CBF function[J].Plant Journal for Cell & Molecular Biology,2010,58(1):97-108.
[11] Ansari S A,Morse R H.Mechanisms of Mediator complex action in transcriptional activation[J].Cellular & Molecular Life Sciences,2013,70(15):2743-2756.
[12] Ansari S A,He Q,Morse R H.Mediator complex association with constitutively transcribed genes in yeast[J].Proceedings of the National Academy of Sciences,2009,106(39):16734-16739.
[13] Gould P D,Locke J C W,Larue C,et al.The molecular basis of temperature compensation in the Arabidopsis circadian clock[J].Plant Cell,2006,18(5):1177-1187.
[14] Knight H,Thomson A J,Mcwatters H G.Sensitive to freezing6 integrates cellular and environmental inputs to the plant circadian clock[J].Plant Physiology,2008,148(1):293-303.
[15] Knight H,Knight M R.The sfr6 mutation in Arabidopsis suppresses low-temperature induction of genes dependent on the CRT/DRE sequence motif.[J].Plant Cell,1999,11(5):875-86.
[16] Kim TW,Kwon Y J,Kim J M,et al.MED16 and MED23 of Mediator are coactivators of lipopolysaccharide and heat-shock-induced transcriptional activators[J].Proceedings of the National Academy of Sciences,2004,101(33):12153-12158.
[17] Kidd B N,Aitken E A,Schenk P M,et al.Plant mediator: mediating thejasmonate response[J].Plant Signal Behav,2010,5(5):718-720.
[18] Wang C,Yao J,Du X,et al.The Arabidopsis mediator complex subunit16 is a key component of basal resistance against the mecrotrophic fungal pathogen Sclerotinia sclerotiorum[J].Plant Physiology,2015,169(1):856-72.
[19] Ding Z J,Yan J Y,Li G X,et al.WRKY41 controls Arabidopsis seed dormancy via direct regulation of ABI3 transcript levels not downstream of ABA[J].The Plant Journal,2014,79(5):810-823.
[20] Zhang X,Wang C,Zhang Y,et al.The Arabidopsis mediator complex subunit16 positively regulates salicylate-mediated is acquired resistance and jasmonate/ethylene-induced defense pathways[J].Plant Cell,2012,24(10):4294-4309.
[21] Sorek N,Szemenyei H,Sorek H,et al.Identification of MEDIATOR16 as the Arabidopsis COBRA suppressor MONGOOSE1[J].Proceedings of the National Academy of Sciences of the United States of America,2015,112(52):16048-16053.
[22] B?ckstr?m S,Elfving N,Nilsson R,et al.Purification of a plant mediator from Arabidopsis thaliana identifies PFT1 as the Med25 subunit[J].Molecular Cell,2007,26(5):717-729.
[23] Wathugala D L,Richards S A,Knight K M R.OsSFR6 is a functional rice orthologue of SENSITIVE TO FREEZING-6 and can act as a regulator of COR gene expression,osmotic stress and freezing tolerance in Arabidopsis[J].New Phytologist,2011,191(4):984-995.
[24] Cerdán P D,Chory J.Regulation of flowering time by light quality[J].Nature,2003,423(6942):881.
[25] Kidd B N,Edgar C I,Kumar K K,et al.The mediator complex subunit PFT1 is a key regulator of jasmonate-dependent defense in Arabidopsis[J].The Plant Cell,2009,21(8):2237-2252.
[26] Sun J Q,Jiang H L,Li C Y.Systemin/jasmonate-mediated systemic defense signaling in tomato[J].Molecular Plant,2011,4(4):607-615.
[27] Cevik V,Kidd B N,Zhang P,et al.MEDIATOR25 acts as an integrative hub for the regulation of jasmonate-responsive gene expression in Arabidopsis[J].PLANT PHYSIOLOGY,2012,160(1):541-555.
[28] Kim YJ,Bj?Rklund S,Li Y,et al.A multiprotein mediator of transcriptional activation and its interaction with the C-terminal repeat domain of RNA polymerase II[J].Cell,1994,77(4):599-608.
[29] IňIgo S,Alvarez M J,Bárbara Strasser,et al.PFT1,the MED25 subunit of the plant mediator complex,promotes flowering through CONSTANS dependent and independent mechanisms in Arabidopsis[J].Plant Journal,2012,69(4):601-612.
[30] Liu J,Zhang T,Jia J,et al.The wheat mediator subunit TaMED25 interacts with the transcription factor TaEIL1 to negatively regulate disease resistance against powdery mildew[J].Plant Physiology,2016,170(3):1799.
[31] Chini A,Fonseca S,Fernández G,et al.The JAZ family of repressors is the missing link in jasmonate signalling[J].Nature,2007,448(7154):666-671.
[32] Berger S,Bell E,Mullet J E.Two methyl jasmonate-insensitive mutants show altered expression of AtVsp in response to methyl jasmonate and wounding[J].Plant Physiology,1996,111(2):525-531.
[33] Robbins C,Kamath Y.The Arabidopsis mediator subunit MED25 differentially regulates jasmonate and abscisic acid signaling through interacting with the MYC2 and ABI5 transcription factors[J].Plant Cell,2012,24(7):2898-2916.
[34] Adel Z,Paula K A,Parisa Y,et al.Two GCC boxes and AP2/ERF-domain transcription factor ORA59 in jasmonate/ethylene-mediated activation of the PDF1.2 promoter in Arabidopsis[J].Plant Molecular Biology,2011,75(4-5):321-331.
[35] Yang L,Li B,Zheng X Y,et al.Salicylic acid biosynthesis is enhanced and contributes to increasedbiotrophic pathogen resistance in Arabidopsis hybrids[J].Nature Communications,2015,6:7309.
[36] Rayagonzález J,Ortizcastro R,Ruízherrera L F,et al.Phytochrome and flowering TIME1/MEDIATOR25 Regulates lateral root formation via auxin signaling in Arabidopsis[J].Plant Physiology,2014,165(2):880.
[37] Ito J,Fukaki H,Onoda M,et al.Auxin-dependent compositional change in mediator in ARF7-and ARF19-mediated transcription.[J].Proceedings of the National Academy of Sciences of the United States of America,2016,113(23):6562.
[38] Yang Y,Ou B,Zhang J,et al.The Arabidopsis Mediator subunit MED16 regulates iron homeostasis by associating with EIN3/EIL1 through subunit MED25[J].Plant Journal,2014,77(6):838-851.
[39] Nogi Y,Fukasawa T.A novel mutation that affects utilization of galactose in Saccharomyces cerevisiae II.A partial aneuploid for chromosome II isolated as a revertant of the mutant[J].Current Genetics,1981,3(2):91.
[40] Malik S,Roeder R G.The metazoan mediator co-activator complex as an integrative hub for transcriptional regulation[J].Nature Reviews Genetics,2010,11(11):761.
[41] Klose C,Buche C,Fernandez A P,et al.The mediator complex subunit PFT1 interferes with COP1 and HY5 in the regulation of Arabidopsis light signaling[J].Plant Physiology,2012,160(1):289-307.
[42] Kumar V,Waseem M,Dwivedi N,et al.KIX domain of AtMed15a,a mediator subunit of Arabidopsis,is required for its interaction with different proteins[J].Plant Signaling & Behavior,2018,13(2):e1428514.
[43] Thakur,Kumar J,Agarwal,et al.Sequence and expression analyses of KIX domain proteins suggest their importance in seed development and determination of seed size in rice and genome stability in Arabidopsis[J].Molecular Genetics & Genomics,2013,288(7-8):329-346.
[44] Stout J,Romero-Severson E,Ruegger M O,et al.Semidominant mutations in reduced epidermal fluorescence 4 reduce phenylpropanoid content in Arabidopsis[J].Genetics,2008,178(4):2237-2251.
[45] Bonawitz N D,Soltau W L,Blatchley M R,et al.REF4 and RFR1,subunits of the transcriptional coregulatory complex mediator,are required for phenylpropanoid homeostasis in Arabidopsis[J].Journal of Biological Chemistry,2012,287(8):5434-45.
[46] Pieterse C M,Leonreyes A,Van d E S,et al.Networking by small-molecule hormones in plant immunity[J].Nature Chemical Biology,2009,5(5):308-316.
[47] Jane G.Contrasting mechanisms of defense against biotrophic and necrotrophic pathogens[J].Annual Review of Phytopathology,2005,43(1):205-227.
[48] Sundaravelpandian K,Chandrika N N P,Schmidt W.PFT1,a transcriptional mediator complex subunit,controls root hair differentiation through reactive oxygen species(ROS)distribution in Arabidopsis[J].New Phytologist,2012,197(1):151-161.
[49] Wang C,Du X,Mou Z.The mediator complex subunits MED14,MED15,and MED16 are involved in defense signaling crosstalk in Arabidopsis[J].Frontiers in Plant Science,2016,7(ra20):1947.
[50] Hemsley P A,Hurst C H,Kaliyadasa E,et al.The Arabidopsis mediator complex subunits MED16,MED14,and MED2 regulate mediator and RNA polymerase II recruitment to CBF-responsive cold-regulated genes[J].Plant Cell,2014,26(1):465-84.
[51] Shaikhali J,Davoine,Céline,Bj?Rklund S,et al.Redox regulation of the MED28 and MED32 mediator subunits is important for development and senescence[J].Protoplasma,2016,253(3):957-963.
[52] Zhang X,Yao J,Zhang Y,et al.The Arabidopsis mediator complex subunits MED14/SWP and MED16/SFR6/IEN1 differentially regulate defense gene expression in plant immune responses[J].Plant Journal,2013,75(3):484-497.
[53] Elfving N,Céline D,Benlloch R,et al.The Arabidopsis thaliana Med25 mediator subunit integrates environmental cues to control plant development[J].Proceedings of the National Academy of Sciences of the United States of America,2011,108(20):8245-8250.
[54] Autran D,Jonak C,Belcram K,et al.Cell numbers and leaf development in Arabidopsis:A functional analysis of the STRUWWELPETER gene[J].Embo Journal,2014,21(22):6036-6049.
[2] Schilbach S,Hantsche M,Tegunov D,et al.Structures of transcription pre-initiation complex with TFIIH and Mediator[J].Nature,2017,551:204-209.
[3] Nozawa K,Schneider T R,Cramer P.Core Mediator structure at 3.4 ? extends model of transcription initiation complex[J].Nature,2017:248-251.
[4] Rival P,Press M O,Bale J,et al.The conserved PFT1 tandem repeat is crucial for proper flowering in Arabidopsis thaliana[J].Genetics,2014,198(2):747-754.
[5] Alonso José M,Stepanova A N,Leisse T J,et al.Genome-wide insertional mutagenesis of Arabidopsis thaliana[J].Science,2003,301(5633):653-657.
[6] Samanta S,Thakur J K.Characterization of mediator complex and its associated proteins from rice[J].Methods in Molecular Biology,2017,1629:123.
[7] YangY,Li L,Qu,Li Jia.Plant Mediator complex and its critical functions in transcription regulation[J].Journal of Integrative Plant Biology,2016,58(2):13.
[8] Plaschka C,Larivière,L,Wenzeck L,et al.Architecture of the RNA polymerase II-Mediator core initiation complex[J].Nature,2015,518(7539):376-380.
[9] Canet J V,Dobón A,Tornero P.Non-recognition-of-BTH4,an Arabidopsis mediator subunit homolog,is necessary for development and response to salicylic acid[J].Plant Cell,2012,24(10):4220-4235.
[10] KnightH,Mugford N G S G,Ulker B,et al.Identification of SFR6,a key component in cold acclimation acting post-translationally on CBF function[J].Plant Journal for Cell & Molecular Biology,2010,58(1):97-108.
[11] Ansari S A,Morse R H.Mechanisms of Mediator complex action in transcriptional activation[J].Cellular & Molecular Life Sciences,2013,70(15):2743-2756.
[12] Ansari S A,He Q,Morse R H.Mediator complex association with constitutively transcribed genes in yeast[J].Proceedings of the National Academy of Sciences,2009,106(39):16734-16739.
[13] Gould P D,Locke J C W,Larue C,et al.The molecular basis of temperature compensation in the Arabidopsis circadian clock[J].Plant Cell,2006,18(5):1177-1187.
[14] Knight H,Thomson A J,Mcwatters H G.Sensitive to freezing6 integrates cellular and environmental inputs to the plant circadian clock[J].Plant Physiology,2008,148(1):293-303.
[15] Knight H,Knight M R.The sfr6 mutation in Arabidopsis suppresses low-temperature induction of genes dependent on the CRT/DRE sequence motif.[J].Plant Cell,1999,11(5):875-86.
[16] Kim TW,Kwon Y J,Kim J M,et al.MED16 and MED23 of Mediator are coactivators of lipopolysaccharide and heat-shock-induced transcriptional activators[J].Proceedings of the National Academy of Sciences,2004,101(33):12153-12158.
[17] Kidd B N,Aitken E A,Schenk P M,et al.Plant mediator: mediating thejasmonate response[J].Plant Signal Behav,2010,5(5):718-720.
[18] Wang C,Yao J,Du X,et al.The Arabidopsis mediator complex subunit16 is a key component of basal resistance against the mecrotrophic fungal pathogen Sclerotinia sclerotiorum[J].Plant Physiology,2015,169(1):856-72.
[19] Ding Z J,Yan J Y,Li G X,et al.WRKY41 controls Arabidopsis seed dormancy via direct regulation of ABI3 transcript levels not downstream of ABA[J].The Plant Journal,2014,79(5):810-823.
[20] Zhang X,Wang C,Zhang Y,et al.The Arabidopsis mediator complex subunit16 positively regulates salicylate-mediated is acquired resistance and jasmonate/ethylene-induced defense pathways[J].Plant Cell,2012,24(10):4294-4309.
[21] Sorek N,Szemenyei H,Sorek H,et al.Identification of MEDIATOR16 as the Arabidopsis COBRA suppressor MONGOOSE1[J].Proceedings of the National Academy of Sciences of the United States of America,2015,112(52):16048-16053.
[22] B?ckstr?m S,Elfving N,Nilsson R,et al.Purification of a plant mediator from Arabidopsis thaliana identifies PFT1 as the Med25 subunit[J].Molecular Cell,2007,26(5):717-729.
[23] Wathugala D L,Richards S A,Knight K M R.OsSFR6 is a functional rice orthologue of SENSITIVE TO FREEZING-6 and can act as a regulator of COR gene expression,osmotic stress and freezing tolerance in Arabidopsis[J].New Phytologist,2011,191(4):984-995.
[24] Cerdán P D,Chory J.Regulation of flowering time by light quality[J].Nature,2003,423(6942):881.
[25] Kidd B N,Edgar C I,Kumar K K,et al.The mediator complex subunit PFT1 is a key regulator of jasmonate-dependent defense in Arabidopsis[J].The Plant Cell,2009,21(8):2237-2252.
[26] Sun J Q,Jiang H L,Li C Y.Systemin/jasmonate-mediated systemic defense signaling in tomato[J].Molecular Plant,2011,4(4):607-615.
[27] Cevik V,Kidd B N,Zhang P,et al.MEDIATOR25 acts as an integrative hub for the regulation of jasmonate-responsive gene expression in Arabidopsis[J].PLANT PHYSIOLOGY,2012,160(1):541-555.
[28] Kim YJ,Bj?Rklund S,Li Y,et al.A multiprotein mediator of transcriptional activation and its interaction with the C-terminal repeat domain of RNA polymerase II[J].Cell,1994,77(4):599-608.
[29] IňIgo S,Alvarez M J,Bárbara Strasser,et al.PFT1,the MED25 subunit of the plant mediator complex,promotes flowering through CONSTANS dependent and independent mechanisms in Arabidopsis[J].Plant Journal,2012,69(4):601-612.
[30] Liu J,Zhang T,Jia J,et al.The wheat mediator subunit TaMED25 interacts with the transcription factor TaEIL1 to negatively regulate disease resistance against powdery mildew[J].Plant Physiology,2016,170(3):1799.
[31] Chini A,Fonseca S,Fernández G,et al.The JAZ family of repressors is the missing link in jasmonate signalling[J].Nature,2007,448(7154):666-671.
[32] Berger S,Bell E,Mullet J E.Two methyl jasmonate-insensitive mutants show altered expression of AtVsp in response to methyl jasmonate and wounding[J].Plant Physiology,1996,111(2):525-531.
[33] Robbins C,Kamath Y.The Arabidopsis mediator subunit MED25 differentially regulates jasmonate and abscisic acid signaling through interacting with the MYC2 and ABI5 transcription factors[J].Plant Cell,2012,24(7):2898-2916.
[34] Adel Z,Paula K A,Parisa Y,et al.Two GCC boxes and AP2/ERF-domain transcription factor ORA59 in jasmonate/ethylene-mediated activation of the PDF1.2 promoter in Arabidopsis[J].Plant Molecular Biology,2011,75(4-5):321-331.
[35] Yang L,Li B,Zheng X Y,et al.Salicylic acid biosynthesis is enhanced and contributes to increasedbiotrophic pathogen resistance in Arabidopsis hybrids[J].Nature Communications,2015,6:7309.
[36] Rayagonzález J,Ortizcastro R,Ruízherrera L F,et al.Phytochrome and flowering TIME1/MEDIATOR25 Regulates lateral root formation via auxin signaling in Arabidopsis[J].Plant Physiology,2014,165(2):880.
[37] Ito J,Fukaki H,Onoda M,et al.Auxin-dependent compositional change in mediator in ARF7-and ARF19-mediated transcription.[J].Proceedings of the National Academy of Sciences of the United States of America,2016,113(23):6562.
[38] Yang Y,Ou B,Zhang J,et al.The Arabidopsis Mediator subunit MED16 regulates iron homeostasis by associating with EIN3/EIL1 through subunit MED25[J].Plant Journal,2014,77(6):838-851.
[39] Nogi Y,Fukasawa T.A novel mutation that affects utilization of galactose in Saccharomyces cerevisiae II.A partial aneuploid for chromosome II isolated as a revertant of the mutant[J].Current Genetics,1981,3(2):91.
[40] Malik S,Roeder R G.The metazoan mediator co-activator complex as an integrative hub for transcriptional regulation[J].Nature Reviews Genetics,2010,11(11):761.
[41] Klose C,Buche C,Fernandez A P,et al.The mediator complex subunit PFT1 interferes with COP1 and HY5 in the regulation of Arabidopsis light signaling[J].Plant Physiology,2012,160(1):289-307.
[42] Kumar V,Waseem M,Dwivedi N,et al.KIX domain of AtMed15a,a mediator subunit of Arabidopsis,is required for its interaction with different proteins[J].Plant Signaling & Behavior,2018,13(2):e1428514.
[43] Thakur,Kumar J,Agarwal,et al.Sequence and expression analyses of KIX domain proteins suggest their importance in seed development and determination of seed size in rice and genome stability in Arabidopsis[J].Molecular Genetics & Genomics,2013,288(7-8):329-346.
[44] Stout J,Romero-Severson E,Ruegger M O,et al.Semidominant mutations in reduced epidermal fluorescence 4 reduce phenylpropanoid content in Arabidopsis[J].Genetics,2008,178(4):2237-2251.
[45] Bonawitz N D,Soltau W L,Blatchley M R,et al.REF4 and RFR1,subunits of the transcriptional coregulatory complex mediator,are required for phenylpropanoid homeostasis in Arabidopsis[J].Journal of Biological Chemistry,2012,287(8):5434-45.
[46] Pieterse C M,Leonreyes A,Van d E S,et al.Networking by small-molecule hormones in plant immunity[J].Nature Chemical Biology,2009,5(5):308-316.
[47] Jane G.Contrasting mechanisms of defense against biotrophic and necrotrophic pathogens[J].Annual Review of Phytopathology,2005,43(1):205-227.
[48] Sundaravelpandian K,Chandrika N N P,Schmidt W.PFT1,a transcriptional mediator complex subunit,controls root hair differentiation through reactive oxygen species(ROS)distribution in Arabidopsis[J].New Phytologist,2012,197(1):151-161.
[49] Wang C,Du X,Mou Z.The mediator complex subunits MED14,MED15,and MED16 are involved in defense signaling crosstalk in Arabidopsis[J].Frontiers in Plant Science,2016,7(ra20):1947.
[50] Hemsley P A,Hurst C H,Kaliyadasa E,et al.The Arabidopsis mediator complex subunits MED16,MED14,and MED2 regulate mediator and RNA polymerase II recruitment to CBF-responsive cold-regulated genes[J].Plant Cell,2014,26(1):465-84.
[51] Shaikhali J,Davoine,Céline,Bj?Rklund S,et al.Redox regulation of the MED28 and MED32 mediator subunits is important for development and senescence[J].Protoplasma,2016,253(3):957-963.
[52] Zhang X,Yao J,Zhang Y,et al.The Arabidopsis mediator complex subunits MED14/SWP and MED16/SFR6/IEN1 differentially regulate defense gene expression in plant immune responses[J].Plant Journal,2013,75(3):484-497.
[53] Elfving N,Céline D,Benlloch R,et al.The Arabidopsis thaliana Med25 mediator subunit integrates environmental cues to control plant development[J].Proceedings of the National Academy of Sciences of the United States of America,2011,108(20):8245-8250.
[54] Autran D,Jonak C,Belcram K,et al.Cell numbers and leaf development in Arabidopsis:A functional analysis of the STRUWWELPETER gene[J].Embo Journal,2014,21(22):6036-6049.