植物转录调控因子NF-Y研究进展
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摘要
NF-Y是在植物、动物和酵母细胞均存在的转录调节因子,由NF-YA、NF-YB以及NY-YC组成异源三聚体,在细胞核内结合在下游基因启动子上调节基因转录活性,或通过与核内其它蛋白的互作调节细胞生命活动.NF-YA(核因子Y,亚基A)是具有多种生物功能的植物转录调控因子,其与NF-YB、NF-YC组成异源三聚体,结合下游基因的CCAAT顺式元件,从而激活或抑制一些基因的表达. NF-YA、NF-YB和NF-YC参与植物体内许多生理过程,包括胚胎发育、种子发育及萌发、花的发育、根和根瘤发育以及各种逆境反应等重要的生理过程.本文就植物中转录调控因子NF-YA、NF-YB和NF-YC的基本结构特点、生物学功能,以及它们在植物响应生物和非生物胁迫方面的作用进行了综述.
NF-Y(nuclear factor Y) is an important transcription factor in plant, animal and yeast cells. It is generally composed of NF-YA, NF-YB and NF-YC, which is a heterotrimer that binds to the promoter of downstream gene in the nucleus to regulate the transcriptional activity, or regulating the cellular life activities by interaction with other proteins in the nucleus. NF-YA(nuclear factor Y, subunit A) is a transcription factor with multiple biological functions, which forms a heterotrimer together with NF-YB and NF-YC, thereby activating or inhibiting the expression of related genes via binding with the CCAAT motif of the downstream genes. NF-YA, NF-YB and NF-YC are involved in many physiological processes in plants, including embryo development, seed development and germination, flower development, root and nodule development and various stress responses. This article reviewed the basic structural features and the biological functions of the transcription factors NF-YA, NF-YB and NF-YC, as well as their roles in plant responses to biotic and abiotic stress.
NF-Y(nuclear factor Y) is an important transcription factor in plant, animal and yeast cells. It is generally composed of NF-YA, NF-YB and NF-YC, which is a heterotrimer that binds to the promoter of downstream gene in the nucleus to regulate the transcriptional activity, or regulating the cellular life activities by interaction with other proteins in the nucleus. NF-YA(nuclear factor Y, subunit A) is a transcription factor with multiple biological functions, which forms a heterotrimer together with NF-YB and NF-YC, thereby activating or inhibiting the expression of related genes via binding with the CCAAT motif of the downstream genes. NF-YA, NF-YB and NF-YC are involved in many physiological processes in plants, including embryo development, seed development and germination, flower development, root and nodule development and various stress responses. This article reviewed the basic structural features and the biological functions of the transcription factors NF-YA, NF-YB and NF-YC, as well as their roles in plant responses to biotic and abiotic stress.
引文
[1] DOLFINI D, GATTA R, MANTOVANI R. NF-Y and the transcriptional activation of CCAAT promoters.[J]. Crit Rev Biochem Mol Biol,2012,47(1):29-49.
[2] LALOUM T, DE MITA, STEPHANE, GAMAS P, et al.Erratum to:‘CCAAT-box binding transcription factors in plants:Y so many?’[J]. Trends in Plant Science,2013,18(10):594-595.
[3] PETRONI K, KUMIMOTO R W, GNESUTTA N, et al.The Promiscuous Life of Plant NUCLEAR FACTOR Y Transcription Factors[J]. Plant Cell,2012,24(12):4777-4792.
[4] LIU J X, HOWELL S H. bZIP28 and NF-Y Transcription Factors Are Activated by ER Stress and Assemble into a Transcriptional Complex to Regulate Stress Response Genes in Arabidopsis[J]. The Plant Cell, 2010,22(3):782-796.
[5] HACKENBERG D, WU Y, VOIGT A, et al. Studies on Differential Nuclear Translocation Mechanism and Assembly of the Three Subunits of the Arabidopsis thaliana Transcription Factor NF-Y[J]. Molecular Plant,2012,5(4):876-888.
[6] MANTOVANI R. The molecular biology of the CCAATbinding factor NF-Y[J]. Gene,1999,239(1):15.
[7] XING Y, ZHANG S, OLESEN J T, et al. Subunit interaction in the CCAAT-binding heteromeric complex is mediated by a very short alpha-helix in HAP2.[J]. Proceedings of the National Academy of Sciences,1994,91(8):3009-3013.
[8] MAITY S N, DE C B. Biochemical analysis of the B subunit of the heteromeric CCAAT-binding factor. A DNA-binding domain and a subunit interaction domain are specified by two separate segments.[J]. Journal of Biological Chemistry,1992,267(12):8286-92.
[9] SIEFERS N, DANG K K, KUMIMOTO R W, et al.Tissue-specific expression patterns of arabidopsis NFY transcription factors suggest potential for extensive combinatorial complexity.[J]. Plant Physiology, 2009,149(2):625-641.
[10] SORIN C, DECLERCK M, CHRIST A, et al. A miR169isoform regulates specific NF-Y targets and root architecture in Arabidopsis[J]. New Phytologist, 2014,202(4):1197-1211.
[11] SIRIWARDANA C L, KUMIMOTO R W, JONES D S,et al. Gene Family Analysis of the Arabidopsis NF-Y Transcription Factors Reveals Opposing Abscisic Acid Responses During Seed Germination[J]. Plant Molecular Biology Reporter, 2014, 32(5):971.
[12] MU J, TAN H, HONG S, et al. Arabidopsis Transcription Factor Genes NF-YA1, 5, 6, and 9 Play Redundant Roles in Male Gametogenesis, Embryogenesis, and Seed Development[J]. Molecular Plant,2013,6(1):188-201.
[13] FORNARI M, CALVENZANI V, MASIERO S, et al.The Arabidopsis NF-YA3 and NF-YA8 Genes Are Functionally Redundant and Are Required in Early Embryogenesis[J]. Plos One,2013,8(11):e82043.
[14] LALOUM T, BAUDIN M, FRANCES L, et al. Two CCAAT box-binding transcription factors redundantly regulate early steps of the legume-rhizobia endosymbiosis[J]. Plant Journal for Cell&Molecular Biology,2015,79(5):757-768.
[15] LAPORTE P, LEPAGE A, FOURNIER, JO LLE, et al. The CCAAT box-binding transcription factor NF-YA1 controls rhizobial infection[J]. Journal of Experimental Botany,2014,65(2):481-494.
[16] SIRIWARDANA C, GNESUTTA N, KUMIMOTO R, et al. NUCLEAR FACTOR Y, subunit A(NF-Y)proteins positively regulate flowering and act through FLOWERING LOCUS T[J]. Plos Genetics,2016,12(12):e1006496.
[17] ALAM M M, TANAKA T, NAKAMURA H, et al.Overexpression of a rice heme activator protein gene(OsHAP2E)confers resistance to pathogens, salinity and drought, and increases photosynthesis and tiller number[J]. Plant Biotechnology Journal,2015,13(1):85-96.
[18] HANEMIAN M, BARLET X, SORIN C, et al. Arabidopsis CLAVATA1 and CLAVATA2 receptors contribute to Ralstonia solanacearum pathogenicity through a miR169-dependent pathway[J]. New Phytol,2016,211(2):502-515.
[19] SINGH K, TALLA A, QIU W. Small RNA profiling of virus-infected grapevines:evidences for virus infection-associated and variety-specific miRNAs[J].Functional&Integrative Genomics,2012,12(4):659-669.
[20] REY T, LAPORTE P, BONHOMME M, et al. MtNFYA1, a central transcriptional regulator of symbiotic nodule development, is also a determinant of Medicago truncatula susceptibility toward a root pathogen[J].Frontiers in Plant Science,2016,7(7).
[21] MARCO ANTONIO LEYVAGONZ LEZ, IBARRALACLETTE E, ALFREDO CRUZRAM REZ, et al. Functional and Transcriptome Analysis Reveals an Acclimatization Strategy for Abiotic Stress Tolerance Mediated by Arabidopsis NF-YA Family Members[J]. Plos One,2012,7(10):e48138.
[22] LI W X, OONO Y, ZHU J, et al. The Arabidopsis NF-YA5 Transcription Factor Is Regulated Transcriptionally and Posttranscriptionally to Promote Drought Resistance[J]. The Plant Cell,2008,20(8):2238-2251.
[23] NI Z, HU Z, JIANG Q, et al. GmNFYA3, a target gene of miR169, is a positive regulator of plant tolerance to drought stress[J]. Plant Molecular Biology,2013,82(1-2):113-129.
[24] LUAN M, XU M, LU Y, et al. Family-Wide Survey of miR169s and NF-YAs and Their Expression Profiles Response to Abiotic Stress in Maize Roots[J]. PLOS ONE,2014,9.
[25] LI Y J, FANG Y, FU Y R, et al. NFYA1 Is Involved in Regulation of Postgermination Growth Arrest Under Salt Stress in Arabidopsis[J]. PLOS ONE,2013,8.
[26] FENG Z J, HE G H, ZHENG W J, et al. Foxtail Millet NF-Y Families:Genome-Wide Survey and Evolution Analyses Identified Two Functional Genes Important in Abiotic Stresses[J]. Frontiers in Plant Science,2015,6.
[27]黄锁,胡利芹,徐东北,等.谷子转录因子SiNF-YA5通过ABA非依赖途径提高转基因拟南芥耐盐性[J].作物学报,2016,42(12):1787-1797.
[28] KIBA T, KRAPP A. Plant Nitrogen Acquisition Under Low Availability:Regulation of Uptake and Root Architecture[J]. Plant&Cell Physiology,2016,57(4):707-714.
[29] ZHAO M, DING H, ZHU J K, et al. Involvement of miR169 in the nitrogen-starvation responses in Arabidopsis[J]. The New phytologist,2011,190(4):906-915.
[30]方广宁,胡利芹,王二辉,等.谷子转录因子SiNF-YA6的过表达提高转基因植株对低氮胁迫的抗性[J].中国农业科学,2015,48(20):3989-3997.
[31] MISSON J, RAGHOTHAMA K G, JAIN A, et al. A genome-wide transcriptional analysis using Arabidopsis thaliana Affymetrix gene chips determined plant responses to phosphate deprivation[J]. Proceedings of the National Academy of Sciences,2005,102(33):11934-11939.
[32] MULLER R, MORANT M, JARMER H, et al. GenomeWide Analysis of the Arabidopsis Leaf Transcriptome Reveals Interaction of Phosphate and Sugar Metabolism[J]. PLANT PHYSIOLOGY,2006,143(1):156-171.
[33] CONTRIBUTOR M T E, PAZARES J, WEIGEL D. A Collection of Target Mimics for Comprehensive Analysis of MicroRNA Function in Arabidopsis thaliana[J]. Plos Genetics,2010,6(7):e1001031.
[34] MARCO ANTONIO LEYVAGONZ LEZ, IBARRALACLETTE E, ALFREDO CRUZRAM REZ, et al. Functional and Transcriptome Analysis Reveals an Acclimatization Strategy for Abiotic Stress Tolerance Mediated by Arabidopsis NF-YA Family Members[J]. Plos One,2012,7(10):e48138.
[35] QU B, HE X, WANG J, et al. A Wheat CCAAT BoxBinding Transcription Factor Increases the Grain Yield of Wheat with Less Fertilizer Input[J]. Plant Physiology,2015,167(2):411-423.
[36] KAHLE J, BAAKE M, DOENECKE D, et al. Subunits of the Heterotrimeric Transcription Factor NF-Y Are Imported into the Nucleus by Distinct Pathways Involving Importin? and Importin 13[J]. Molecular and Cellular Biology,2005,25(13):5339-5354.
[37] LIU J X, HOWELL S H. bZIP28 and NF-Y Transcription Factors Are Activated by ER Stress and Assemble into a Transcriptional Complex to Regulate Stress Response Genes in Arabidopsis[J]. The Plant Cell,2010,22(3):782-796.
[38] CERIBELLI M, DOLFINI D, MERICO D, et al. The Histone-Like NF-Y Is a Bifunctional Transcription Factor[J]. Mol.cell.biol,2008,28(6):2047-2058.
[39] FLEMING J D, PAVESI G, BENATTI P, et al. NF-Y coassociates with FOS at promoters, enhancers,repetitive elements, and inactive chromatin regions,and is stereo-positioned with growth-controlling transcription factors[J]. Genome Research, 2013,23(8):1195-1209.
[40] CAO S, KUMIMOTO R W, GNESUTTA N, et al. A distal CCAAT/NUCLEAR FACTOR Y complex promotes chromatin looping at the FLOWERING LOCUS T promoter and regulates the timing of flowering in Arabidopsis[J]. The Plant Cell,2014,26(3).
[41] BATTAGLIA M, R PODAS C, CL A J, et al. A nuclear factor Y interacting protein of the GRAS family is required for nodule organogenesis, infection thread progression, and lateral root growth[J]. Plant Physiology,2014,164(3):1430-1442.
[42] JUNKER A, GUDRUN M?NKE, RUTTEN T, et al.Elongation-related functions of LEAFY COTYLEDON1 during the development of Arabidopsis thaliana[J]. Plant Journal,2012,71(12):427-42.
[43] SUN X, LING S, LU Z, et al. OsNF-YB1, a rice endosperm-specific gene, is essential for cell proliferation in endosperm development[J]. Gene,2014,551(2):214-221.
[44] STEPHENSON T J, MCINTYRE C L, COLLET C, et al.TaNF-YB3, is involved in the regulation of photosynthesis genes in Triticum aestivum[J].Functional&Integrative Genomics,2011,11(2):327-340.
[45] KUMIMOTO R W, ADAM L, HYMUS G J, et al. The Nuclear Factor Y subunits NF-YB2 and NF-YB3 play additive roles in the promotion of flowering by inductive long-day photoperiods in Arabidopsis[J].Planta,2008,228(5):709-723.
[46] LI C, DISTELFELD A, COMIS A, et al. Wheat flowering repressor VRN2 and promoter CO2 compete for interactions with NUCLEAR FACTOR-Y complexes[J]. The Plant Journal,2011,67(5):11.
[47] HOU X, ZHOU J, LIU C, et al. Nuclear factor Y-mediated H3K27me3 demethylation of the SOC1 locus orchestrates flowering responses of Arabidopsis[J].Nature Communications,2014,5(8).
[48] TIWARI S B, SHEN Y, CHANG H C, et al. The flowering time regulator CONSTANS is recruited to the FLOWERING LOCUS T promoter via a unique ciselement[J]. New Phytologist,2010,187(1):57-66.
[49]付蕊欣.油菜BnNF-YB2/3/4/5/6在植物花期调节以及非生物胁迫中的功能研究[D].南京:南京农业大学,2016.
[50] MAR A EUGENIA ZANETTI, BLANCO F A, MAR A P A BEKER, et al. A C subunit of the plant nuclear factor NF-Y required for rhizobial infection and nodule development affects partner selection in the common bean-Rhizobium etli symbiosis.[J]. Plant Cell,2010,22(12):4142-4157.
[51] SOYANO T, KOUCHI H, HIROTA A, et al. NODULE INCEPTION Directly Targets NF-Y Subunit Genes to Regulate Essential Processes of Root Nodule Development in Lotus japonicus[J]. Plos Genetics,2013,9(3):e1003352.
[52] MAR A EUGENIA ZANETTI, BLANCO F A, MAR A P A BEKER, et al. A C subunit of the plant nuclear factor NF-Y required for rhizobial infection and nodule development affects partner selection in the common bean-Rhizobium etli symbiosis.[J]. Plant Cell,2010,22(12):4142-4157.
[53] BATTAGLIA M, R PODAS C, CLoA J, et al. A nuclear factor Y interacting protein of the GRAS family is required for nodule organogenesis, infection thread progression, and lateral root growth[J]. Plant Physiology,2014,164(3):1430-1442.
[54] R PODAS, CAROLINA, CL A, JOAQU N, BATTAGLIA M, et al. Transcriptional regulators of legume-rhizobia symbiosis[J]. Plant Signaling&Behavior,2014,9(5):e28847.
[55] LIU X, LIU X, LI Y, et al. Arabidopsis NF-YCs Mediate the Light-Controlled Hypocotyl Elongation via Modulating Histone Acetylation[J]. Molecular Plant,2017,10(2):260-273.
[56] MYERS Z A, KUMIMOTO R W, SIRIWARDANA C L,et al. NUCLEAR FACTOR Y, Subunit C(NF-YC)Transcription Factors Are Positive Regulators of Photomorphogenesis in Arabidopsis thaliana[J]. Plos Genetics,2016,12(9):e1006333.
[57] HUANG M, HU Y, LIU X, et al. Arabidopsis Leafy Cotyledon1 Mediates Postembryonic Development via Interacting with Phytochrome-Interacting Factor4[J].The Plant Cell,2015,27(11):3099-3111.
[2] LALOUM T, DE MITA, STEPHANE, GAMAS P, et al.Erratum to:‘CCAAT-box binding transcription factors in plants:Y so many?’[J]. Trends in Plant Science,2013,18(10):594-595.
[3] PETRONI K, KUMIMOTO R W, GNESUTTA N, et al.The Promiscuous Life of Plant NUCLEAR FACTOR Y Transcription Factors[J]. Plant Cell,2012,24(12):4777-4792.
[4] LIU J X, HOWELL S H. bZIP28 and NF-Y Transcription Factors Are Activated by ER Stress and Assemble into a Transcriptional Complex to Regulate Stress Response Genes in Arabidopsis[J]. The Plant Cell, 2010,22(3):782-796.
[5] HACKENBERG D, WU Y, VOIGT A, et al. Studies on Differential Nuclear Translocation Mechanism and Assembly of the Three Subunits of the Arabidopsis thaliana Transcription Factor NF-Y[J]. Molecular Plant,2012,5(4):876-888.
[6] MANTOVANI R. The molecular biology of the CCAATbinding factor NF-Y[J]. Gene,1999,239(1):15.
[7] XING Y, ZHANG S, OLESEN J T, et al. Subunit interaction in the CCAAT-binding heteromeric complex is mediated by a very short alpha-helix in HAP2.[J]. Proceedings of the National Academy of Sciences,1994,91(8):3009-3013.
[8] MAITY S N, DE C B. Biochemical analysis of the B subunit of the heteromeric CCAAT-binding factor. A DNA-binding domain and a subunit interaction domain are specified by two separate segments.[J]. Journal of Biological Chemistry,1992,267(12):8286-92.
[9] SIEFERS N, DANG K K, KUMIMOTO R W, et al.Tissue-specific expression patterns of arabidopsis NFY transcription factors suggest potential for extensive combinatorial complexity.[J]. Plant Physiology, 2009,149(2):625-641.
[10] SORIN C, DECLERCK M, CHRIST A, et al. A miR169isoform regulates specific NF-Y targets and root architecture in Arabidopsis[J]. New Phytologist, 2014,202(4):1197-1211.
[11] SIRIWARDANA C L, KUMIMOTO R W, JONES D S,et al. Gene Family Analysis of the Arabidopsis NF-Y Transcription Factors Reveals Opposing Abscisic Acid Responses During Seed Germination[J]. Plant Molecular Biology Reporter, 2014, 32(5):971.
[12] MU J, TAN H, HONG S, et al. Arabidopsis Transcription Factor Genes NF-YA1, 5, 6, and 9 Play Redundant Roles in Male Gametogenesis, Embryogenesis, and Seed Development[J]. Molecular Plant,2013,6(1):188-201.
[13] FORNARI M, CALVENZANI V, MASIERO S, et al.The Arabidopsis NF-YA3 and NF-YA8 Genes Are Functionally Redundant and Are Required in Early Embryogenesis[J]. Plos One,2013,8(11):e82043.
[14] LALOUM T, BAUDIN M, FRANCES L, et al. Two CCAAT box-binding transcription factors redundantly regulate early steps of the legume-rhizobia endosymbiosis[J]. Plant Journal for Cell&Molecular Biology,2015,79(5):757-768.
[15] LAPORTE P, LEPAGE A, FOURNIER, JO LLE, et al. The CCAAT box-binding transcription factor NF-YA1 controls rhizobial infection[J]. Journal of Experimental Botany,2014,65(2):481-494.
[16] SIRIWARDANA C, GNESUTTA N, KUMIMOTO R, et al. NUCLEAR FACTOR Y, subunit A(NF-Y)proteins positively regulate flowering and act through FLOWERING LOCUS T[J]. Plos Genetics,2016,12(12):e1006496.
[17] ALAM M M, TANAKA T, NAKAMURA H, et al.Overexpression of a rice heme activator protein gene(OsHAP2E)confers resistance to pathogens, salinity and drought, and increases photosynthesis and tiller number[J]. Plant Biotechnology Journal,2015,13(1):85-96.
[18] HANEMIAN M, BARLET X, SORIN C, et al. Arabidopsis CLAVATA1 and CLAVATA2 receptors contribute to Ralstonia solanacearum pathogenicity through a miR169-dependent pathway[J]. New Phytol,2016,211(2):502-515.
[19] SINGH K, TALLA A, QIU W. Small RNA profiling of virus-infected grapevines:evidences for virus infection-associated and variety-specific miRNAs[J].Functional&Integrative Genomics,2012,12(4):659-669.
[20] REY T, LAPORTE P, BONHOMME M, et al. MtNFYA1, a central transcriptional regulator of symbiotic nodule development, is also a determinant of Medicago truncatula susceptibility toward a root pathogen[J].Frontiers in Plant Science,2016,7(7).
[21] MARCO ANTONIO LEYVAGONZ LEZ, IBARRALACLETTE E, ALFREDO CRUZRAM REZ, et al. Functional and Transcriptome Analysis Reveals an Acclimatization Strategy for Abiotic Stress Tolerance Mediated by Arabidopsis NF-YA Family Members[J]. Plos One,2012,7(10):e48138.
[22] LI W X, OONO Y, ZHU J, et al. The Arabidopsis NF-YA5 Transcription Factor Is Regulated Transcriptionally and Posttranscriptionally to Promote Drought Resistance[J]. The Plant Cell,2008,20(8):2238-2251.
[23] NI Z, HU Z, JIANG Q, et al. GmNFYA3, a target gene of miR169, is a positive regulator of plant tolerance to drought stress[J]. Plant Molecular Biology,2013,82(1-2):113-129.
[24] LUAN M, XU M, LU Y, et al. Family-Wide Survey of miR169s and NF-YAs and Their Expression Profiles Response to Abiotic Stress in Maize Roots[J]. PLOS ONE,2014,9.
[25] LI Y J, FANG Y, FU Y R, et al. NFYA1 Is Involved in Regulation of Postgermination Growth Arrest Under Salt Stress in Arabidopsis[J]. PLOS ONE,2013,8.
[26] FENG Z J, HE G H, ZHENG W J, et al. Foxtail Millet NF-Y Families:Genome-Wide Survey and Evolution Analyses Identified Two Functional Genes Important in Abiotic Stresses[J]. Frontiers in Plant Science,2015,6.
[27]黄锁,胡利芹,徐东北,等.谷子转录因子SiNF-YA5通过ABA非依赖途径提高转基因拟南芥耐盐性[J].作物学报,2016,42(12):1787-1797.
[28] KIBA T, KRAPP A. Plant Nitrogen Acquisition Under Low Availability:Regulation of Uptake and Root Architecture[J]. Plant&Cell Physiology,2016,57(4):707-714.
[29] ZHAO M, DING H, ZHU J K, et al. Involvement of miR169 in the nitrogen-starvation responses in Arabidopsis[J]. The New phytologist,2011,190(4):906-915.
[30]方广宁,胡利芹,王二辉,等.谷子转录因子SiNF-YA6的过表达提高转基因植株对低氮胁迫的抗性[J].中国农业科学,2015,48(20):3989-3997.
[31] MISSON J, RAGHOTHAMA K G, JAIN A, et al. A genome-wide transcriptional analysis using Arabidopsis thaliana Affymetrix gene chips determined plant responses to phosphate deprivation[J]. Proceedings of the National Academy of Sciences,2005,102(33):11934-11939.
[32] MULLER R, MORANT M, JARMER H, et al. GenomeWide Analysis of the Arabidopsis Leaf Transcriptome Reveals Interaction of Phosphate and Sugar Metabolism[J]. PLANT PHYSIOLOGY,2006,143(1):156-171.
[33] CONTRIBUTOR M T E, PAZARES J, WEIGEL D. A Collection of Target Mimics for Comprehensive Analysis of MicroRNA Function in Arabidopsis thaliana[J]. Plos Genetics,2010,6(7):e1001031.
[34] MARCO ANTONIO LEYVAGONZ LEZ, IBARRALACLETTE E, ALFREDO CRUZRAM REZ, et al. Functional and Transcriptome Analysis Reveals an Acclimatization Strategy for Abiotic Stress Tolerance Mediated by Arabidopsis NF-YA Family Members[J]. Plos One,2012,7(10):e48138.
[35] QU B, HE X, WANG J, et al. A Wheat CCAAT BoxBinding Transcription Factor Increases the Grain Yield of Wheat with Less Fertilizer Input[J]. Plant Physiology,2015,167(2):411-423.
[36] KAHLE J, BAAKE M, DOENECKE D, et al. Subunits of the Heterotrimeric Transcription Factor NF-Y Are Imported into the Nucleus by Distinct Pathways Involving Importin? and Importin 13[J]. Molecular and Cellular Biology,2005,25(13):5339-5354.
[37] LIU J X, HOWELL S H. bZIP28 and NF-Y Transcription Factors Are Activated by ER Stress and Assemble into a Transcriptional Complex to Regulate Stress Response Genes in Arabidopsis[J]. The Plant Cell,2010,22(3):782-796.
[38] CERIBELLI M, DOLFINI D, MERICO D, et al. The Histone-Like NF-Y Is a Bifunctional Transcription Factor[J]. Mol.cell.biol,2008,28(6):2047-2058.
[39] FLEMING J D, PAVESI G, BENATTI P, et al. NF-Y coassociates with FOS at promoters, enhancers,repetitive elements, and inactive chromatin regions,and is stereo-positioned with growth-controlling transcription factors[J]. Genome Research, 2013,23(8):1195-1209.
[40] CAO S, KUMIMOTO R W, GNESUTTA N, et al. A distal CCAAT/NUCLEAR FACTOR Y complex promotes chromatin looping at the FLOWERING LOCUS T promoter and regulates the timing of flowering in Arabidopsis[J]. The Plant Cell,2014,26(3).
[41] BATTAGLIA M, R PODAS C, CL A J, et al. A nuclear factor Y interacting protein of the GRAS family is required for nodule organogenesis, infection thread progression, and lateral root growth[J]. Plant Physiology,2014,164(3):1430-1442.
[42] JUNKER A, GUDRUN M?NKE, RUTTEN T, et al.Elongation-related functions of LEAFY COTYLEDON1 during the development of Arabidopsis thaliana[J]. Plant Journal,2012,71(12):427-42.
[43] SUN X, LING S, LU Z, et al. OsNF-YB1, a rice endosperm-specific gene, is essential for cell proliferation in endosperm development[J]. Gene,2014,551(2):214-221.
[44] STEPHENSON T J, MCINTYRE C L, COLLET C, et al.TaNF-YB3, is involved in the regulation of photosynthesis genes in Triticum aestivum[J].Functional&Integrative Genomics,2011,11(2):327-340.
[45] KUMIMOTO R W, ADAM L, HYMUS G J, et al. The Nuclear Factor Y subunits NF-YB2 and NF-YB3 play additive roles in the promotion of flowering by inductive long-day photoperiods in Arabidopsis[J].Planta,2008,228(5):709-723.
[46] LI C, DISTELFELD A, COMIS A, et al. Wheat flowering repressor VRN2 and promoter CO2 compete for interactions with NUCLEAR FACTOR-Y complexes[J]. The Plant Journal,2011,67(5):11.
[47] HOU X, ZHOU J, LIU C, et al. Nuclear factor Y-mediated H3K27me3 demethylation of the SOC1 locus orchestrates flowering responses of Arabidopsis[J].Nature Communications,2014,5(8).
[48] TIWARI S B, SHEN Y, CHANG H C, et al. The flowering time regulator CONSTANS is recruited to the FLOWERING LOCUS T promoter via a unique ciselement[J]. New Phytologist,2010,187(1):57-66.
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