한수지 50(3), , 2017 Original Article Korean J Fish Aquat Sci 50(3), ,2017 조피볼락 (Sebastes schlegelii) Interferon Regulatory Factor 8 (IRF8)
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1 한수지 50(3), , 2017 Original Article Korean J Fish Aquat Sci 50(3), ,2017 조피볼락 (Sebastes schlegelii) Interferon Regulatory Factor 8 (IRF8) 의분자유전학적특성및발현분석 양혜림 권혁재 이성도 S.D.N.K Bathige 김명진 이제희 제주대학교해양생명과학과 Molecular Characterization and Expression Analysis of Interferon Regulatory Factor 8 (IRF8) in the Black Rockfish Sebastes schlegelii Hyerim Yang, Hyukjae Kwon, Seongdo Lee, S.D.N.K Bathige, Myoung-Jin Kim, and Jehee Lee Department of Marine Life Sciences, Jeju National University, Jeju 63243, Korea Interferon regulatory factor 8 (IRF8) is essential for the development of B and T cells, as well as for the activity of dendritic cells and macrophages. We performed molecular characterization of IRF8 from rock fish, Sebastes schlegelii (Ss), and investigated the spatial and temporal profile of mrna expression after challenge with lipopolysaccharide (LPS), polyinosinic:polycytidylic acid (poly I:C), or Streptococcus iniae. The full-length cdna sequence of SsIRF8 was 1,657 bp, containing an ORF of 1,266 bp. The gene had a predicted molecular mass of 47.7 kda and an isoelectric point of The amino acid sequence coded by this gene showed the highest degree of identity (90.8%) and similarity (96.2%) with IRF8 from Oplegnathus fasciatus. The SsIRF8 mrna was expressed ubiquitously, at varying levels, with the highest level of expression observed in the spleen. To confirm the role of SsIRF8 in mediating the immune response, we measured SsIRF8 mrna expression in the splenic tissue at different time points after injection with LPS, poly I:C, or S. iniae. The qrt-pcr results showed that SsIRF8 mrna expression in the poly I:C-injected group was highly upregulated 6 hr after exposure (P<0.05). Expression of SsIRF8 mrna in the S. iniae-injected group peaked at 24 hr. These results suggest that SsIRF8 might be important in regulating the strength of the rockfish immune response to immunostimulatory agents. Key words: Black rockfish, IRF8, Immune regulator, mrna expression 서론 Interferon regulatory factor (IRF) family,,, (Barnes et al., 2002 ), 11 IRF family (Huang et al., 2010). IRF (interferon, IFN) interferon stimulated genes (ISGs) (Nguyen et al., 1997),,,, (Pfeffer et al., 1998). Type I Type II IFN (Inkpen et al., 2015), Type I IFNs (IFN IFN ) Type I IFNs (Mamane et al., 1999; Taniguchi et al., 2001; Tailor et al., 2007). IRF family N- ~120 amino acid 5 tryptophan (Trp) helixturn-helix motif DNA binding domain (DBD) (Nguyen et al., 1997). IRF DBD consensus DNA sequence IFN-stimulated response element (ISRE) GAAA, AANNNGAA ISRE (Honda and Taniguchi, 2006; Paun and Pitha, 2007). IRF-associated domain (IAD) IRF family C- IRF3-10 IAD1, IRF1 IRF2 IAD2 (Mamane Korean J Fish Aquat Sci 50(3) , June 2017 This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial Licens ( which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited. Received 1 June 2017; Revised 22 June 2017; Accepted 23 June 2017 Corresponding author: Tel: Fax: address: jehee@jejunu.ac.kr (J. Lee), mj.kim.lucky@gmail.com (M-J. Kim) Copyright 2017 The Korean Society of Fisheries and Aquatic Science 302 pissn: , eissn:
2 IRF8 의특징및발현분석 303 et al., 1999). IAD IRFs homodimer heterodimer (Eroshkin and Mushegian, 1999; Ozato et al., 2007). IRF family N- DBD C- IAD DBD IAD (Nehyba et al., 2002). IAD IRF family, (IRF1, IRF3, IRF7, IRF9), (IRF2, IRF8), (IRF2, IRF4, IRF5, IRF8) (Stellacci et al., 2004). IRF1, IRF2, IRF8 myeloid lymphoid cell (Wang and Morse, 2009; Marecki et al., 2015)., IRF8 IRF4 hematopoietic cell, myeloid progenitor cell B T (Lu, 2008), dendritic cells (DCs) plasmacytoid dendritic cells (pdcs) type I IFNs (Alter-Koltunoff et al., 2008). IRF8 Ets-IRFE motif ISRE motif (Lee et al., 2006). IRF8 LPS-TLR4 polyinosinic:polycytidylic acid (poly I:C)-TLR3 Toll-like receptor IFN- signaling pathways (Zhao et al., 2006), poly I:C-activated signaling pathways plasmacytoid DCs (pdcs) IFN DCs subtype IRF8 (Gabriele and Ozato, 2007; Tailor et al., 2007). IRF8 rainbow trout (Holland et al., 2010), turbot (Chen et al., 2012), rock bream (Bathige et al., 2012), Japanese flounder (Hu et al., 2013), tongue sole (Zhang et al., 2015), atlantic cod (Inkpen et al., 2015). in silico, IRF8, LPS, poly I:C, Streptococcus iniae. 재료및방법 Rockfish 의 IRF8 cdna sequence 분석 454 GS FLX sequencing (Roche 454 Life science, USA) cdna sequence database. DNAsist 2.2 software open reading frame (ORF). Expert Protein Analysis System ProtParam tool (SIB Swiss Institute of Bioinformatics, Swiss) ( expasy.org/protparam)., Basic Local Alignment Search Tool (BLAST) algorithm SsIRF8 IRF family member. IRF family domain motif proteins, motif scan Pfam hidden Markov models (Local models) (SIB Swiss Institute of Bioinformatics, Swiss) ( PFSCAN) IRF8 IRF8 homolog Matgat software ClustalW2 server (EMBL- EBI, UK) ( pairwise sequence alignment multiple sequence alignment strategies. Molecular Evolutionary Genetic Analysis (MEGA) software version 7.0 neighbor-joining (NJ) 1,000 bootstrap. 시험어준비및조직분리 (200 20g) L. SsIRF8. gill, heart, head kidney, spleen, liver, intestine, kidney, testis, ovary 9-80., 0.2% heparin sodium salt (USB, USA) (-1 ml/fish). (4, 3,000 g, 10 ) peripheral blood cells (PBCs) -80. 공격실험을통한면역자극비교및조직분리, SsIRF8 200 L 1 phosphate buffered saline (PBS) lipopolysaccharide (LPS; 1.25 g/ L) 200 L 1 PBS polyinosinic: polycytidylic acid (poly I:C; 1.5 g/ L, Sigma, USA), 200 L 1 PBS Streptococcus iniae ( colony-forming units/ L). 200 L 1 PBS 3, 6, 12, 24, 48, 72 5 RNA -80.
3 304 양혜림ㆍ권혁재ㆍ이성도ㆍ S.D.N.K Bathige ㆍ김명진ㆍ이제희 RNA 추출및 cdna 합성 qpcr 에의한발현분석 QIAzol (Qiagen) RNA. RNA purity 1.5% agarose gel, 260 nm Drop Plate (Thermo Scientific, Waltham, MA, USA). cdna RNA 2.5 g reaction mixture 20 L PrimeScript first-strand cdna synthesis kit (TaKaRa, Shiga, Japan)., cdna nuclease-free water 40 qpcr assay 20. quantitative real-time PCR (qpcr) IRF8 mrna. Reference gene elongation factor 1- (EF1 ) forward (5'-AACCTGACCACTGAGGTGAAGTCTG-3'), Reverse (5'-TCCTTGACGGACACGTTCTTGATGTT-3') oligonucleotide primer. SsIRF8 oligonucleotide primer forward (5'-CATCAATGCTTTCCCTCC- GCATCAAG-3'), reverse (5'-TGTGCATCAGCTTTCCTCC- GTAGT-3'). qpcr Real Time System TP800 Thermal Cycler Dice (TaKaRa). 5 -UTR(67) AAGAGCTCAACCGCTTATCACACTGCACAAGGAAACAATCCCCAAAACGCCCTTTCTCTTCATTAAG ATG TCA AAC ACT GGA GGT CGG AGA CTG AAG CAG TGG CTG GTG GAG CAG ATC CAG AGT GGC CAG TAC TCT GGA M S N T G G R R L K Q W L V E Q I Q S G Q Y S G CTG CAG TGG GAG GAT GAC AGC CGC ACT ATG TTC AGA ATC CCA TGG AAA CAC GCA GGC AAA CAA GAC TAC AAC L Q W E D D S R T M F R I P W K H A G K Q D Y N CAA GAA GTA GAC GCA TCC ATT TTC AAG GCC TGG GCT GTG TTT AAA GGC AAG TTT AAG GAG GGG GAG AAG GCT Q E V D A S I F K A W A V F K G K F K E G E K A GAG CCT GCC ACC TGG AAG ACC AGA CTC CGC TGT GCC CTG AAT AAA AGC CCT GAC TTT GAG GAG GTG ACT GAA E P A T W K T R L R C A L N K S P D F E E V T E AGG TCG CAG CTG GAC ATC TCT GAG CCC TAT AAA GTC TAC CGC ATT GTA CCT GAG GAA GAG CAG AAA AAC GGC R S Q L D I S E P Y K V Y R I V P E E E Q K N G AAA GGC GCA TTG ATG GCC ATG GCA GCC ACC ACC AGC TCC GGT GAT ATC AAT GAC ATG GAC TGC AGC CCT GCA K G A L M A M A A T T S S G D I N D M D C S P A GAG ATA GAG GAG CTC ATC AAA GAG GAG GAA GGC TGT AGT ATC CAG ACC AGT CCA GAG TAT TGG TCC CAT GGC E I E E L I K E E E G C S I Q T S P E Y W S H G AGC ATC AAT GCT TTC CCT CCG CAT CAA GAC CCT TTG CCA TCA GGC AAT CTC AGC ACA GCT TTC TCC CAG ATG S I N A F P P H Q D P L P S G N L S T A F S Q M ATG ATC AGT TTC TAC TAC GGA GGA AAG CTG ATG CAC AAC ACA CAT GTT GTT CAT CCT GAA GGC TGC CGA ATC M I S F Y Y G G K L M H N T H V V H P E G C R I GCC CCA CAA CAG CAC CTG GGC CGT GGC GCC CTA TAC AAT TCA GAC AGC ATG CAG AGT GTT CAC TTT CCT CCC A P Q Q H L G R G A L Y N S D S M Q S V H F P P GCT GAG CAC ATC GAG TAC GAC CGC CAG CGC CAT GTC ACA CGC AAG CTC CTG GGA CAC CTG GAG AGA GGT GTA A E H I E Y D R Q R H V T R K L L G H L E R G V CTG GTC CGT GCC AAC CAA GAG GGC ATC TTC ATC AAA AGG CTG TGC CAG AGC CGT GTC TTC TGG AGC GGG TTG L V R A N Q E G I F I K R L C Q S R V F W S G L GGA GAA GTG GGC TCA CAA TAT GGC TCC ATG CCT TGT AAA CTT GAG AGG GAT GCT GTA GTC AAG ATT TTT GAC G E V G S Q Y G S M P C K L E R D A V V K I F D ACA GGA AAG TTT CTT CAA GCT CTC CAG CTG TAC CAG GAG GGT CAG TTT CCA GCT CCT GAT CCG ACA GTG ACT T G K F L Q A L Q L Y Q E G Q F P A P D P T V T CTG TGT TTC GGA GAG GAG CTC CAT GAC CTC AGC AAT GCC AAG AGC AAG CTG ATC ATT GTG CAG ATC ACT GTG L C F G E E L H D L S N A K S K L I I V Q I T V GTG AAC TGC CAG CAG CTG CTG GAG GCA GTG AAC ATG CGT CGC TCC CAG CCT TAC TCC AAC AGC TCA AAC CTG V N C Q Q L L E A V N M R R S Q P Y S N S S N L GAG ATG TCT GAT AAT GTG GCC ATT GAC CAG ATG GCC CGC ATC TAC CAG GAC TTG TGC AGC TAT AGC GGC CCC E M S D N V A I D Q M A R I Y Q D L C S Y S G P TAC CAG GAC TTG TGC AGC TAT AGC GGC CCC CAG AGG CCA GCC TGC TAC AGG GAC AAC ATG CCC ATC ACT GCC Y Q D L C S Y S G P Q R P A C Y R D N M P I T A 3 -UTR(324) TGAGCCGTGAGCAGCAGCAAAACACCTGCCAGGAGCTCTTCAGCGAGTACCAGACTGACTGTACAACATATTAGGGACAGTCCTTATTTT CCTGACATACACTGACAAGGTGAATTCAGGTAAAAACCAGTATCACAATTAGTTTTGACTTTTTAAATGGAGCAAATATCTTACAGTAAGACT TTTACGCTGGAGCTCAGTATTAGGATTTTGTATACCAGTAATGTTTATGTGTCTTATATGTACAATGCTAACACAAACTCTGTGTCTAACTTC TGCTTTTTTACAATAAGAGAGTGTTATCATCCAGCTCGAGAGAAAAAG Fig. 1. The cdna and amino acid sequences of the black rockfish Sebastes schlegelii IRF8. The open reading frame from ATG to TGA is shown in the middle box. The amino acid sequence of the DNA binding domain (DBD) is shaded in gray and the IRF-association domain 1 (IAD1) is in bold, shaded in gray, and boxed. The five conserved tryptophan (W) residues in the DNA binding domain are boxed.
4 IRF8 의특징및발현분석 305 qpcr 3 L cdna, 5 L 2 TaKaRa Ex Taq SYBR premix, 0.4 L 10 M forward, reverse primer, 1.2 L H 2 O 10 L. qpcr cycle, 95 5, 58 10, cycle, 95 15, 60 30, cycle 3. 통계학적분석 SPSS version 18 program (IBM, USA) one-way ANOVA Table 1. The percentages of identity and similarity of SsIRF8 amino acid sequence with that of other species. Oplegnathus fasciatus IRF8 protein sequence showed the highest values Scientific Name Accession Number Sebastes schlegelii Oplegnathus fasciatus Epinephelus coioides Epinephelus coioides Miichthys miiuy Dicentrarchus labrax Scophthalmus maximus Paralichthys olivaceus Larimichthys crocea Gadus morhua Oncorhynchus mykiss Ctenopharyngodon idella Gymnocypris przewalskii Danio rerio Scleropages formosus Gallus gallus Cathartes aura Sebastes schlegelii Oplegnathus fasciatus Epinephelus coioides Epinephelus coioides Similarity (%) Identity (%) AFU AND AND Miichthys miiuy AHB Dicentrarchus labrax Scophthalmus maximus Paralichthys olivaceus Larimichthys crocea AKC AFE AFE KKF Gadus morhua AJR Oncorhynchus mykiss Ctenopharyngodon idella Gymnocypris przewalskii ALS AMT AMB Danio rerio NP_ Scleropages formosus KPP Gallus gallus NP_ Cathartes aura KFP
5 306 양혜림ㆍ권혁재ㆍ이성도ㆍ S.D.N.K Bathige ㆍ김명진ㆍ이제희, mrna (Mean) (Standard deviation) IRF8 mrna. P<0.05. 결과및고찰 IRF8 full-length cdna sequence mrna. SsIRF8 full-length cdna sequence 1,658 bp, 5 -untranslated region (UTR) 67 bp, 3 -untranslated region (UTR) 324 bp. Open reading frame (ORF) 1,266 bp 422 (Fig. 1) kda SsIRF8 bioinformatics tool in silico IRF family DBD, tryptophan, IAD. DBD IRF family ISRE/IRF-E consensus sequence. 5 tryptophan DBD IRF motif (Eroshkin and Mushegian, 1999; Nehyba et al., 2002). SsIRF8 DNA-binding domain N aa IRF3-10 IRF-association domain (IAD) C aa. 5 tryptophan 12W, 27W, 39W, 59W, 77W (Fig. 1). IRF8 IRF8, Oplegnathus fasciatus IRF8 96.2%, 90.8% Epinephelus coioides, Miichthys miiuy, Dicentrarchus labrax 95.5%, 95.3%, 95.7% (Table 1). Phylogenetic tree SsIRF8. (Miichthys miiuy), (Larimichthys crocea), (Dicentrarchus labrax), (Oplegna Miichthys miiuy Larimichthys crocea Dicentrarchus labrax Oplegnathus fasciatus 100 Epinephelus coioides Epinephelus coioides(2) Scophthalmus maximus Paralichthys olivaceus Sebastes schlegelii Gadus morhua Oncorhynchus mykiss Scleropages formosus Danio rerio Ctenopharyngodon idella Gymnocypris przewalskii Gallus gallus 100 Cathartes aura 0.05 Fig. 2. Neighbor-joining phylogenetic tree of IRF8 constructed with Mega7. The bootstrap confidence values shown at the nodes of the tree are based on 1000 bootstrap replications. Black rock fish Sebastes schlegelii IRF8 is indicated with a black dot ( ) and the scientific name of the fish is in bold. GenBank accession numbers are as follows: Miichthys miiuy, AHB ; Larimichthys crocea, KKF ; Dicentrarchus labrax, AKC ; Oplegnathus fasciatus, AFU ; Epinephelus coioides, AND , AND ; Scophthalmus maximus, AFE , Paralichthys olivaceus AFE ; Gadus morhua, AJR ; Oncorhynchus mykiss, ALS ; Scleropages formosus, KPP ; Danio rerio, NP_ ; Ctenopharyngodon idella, AMT ; Gymnocypris przewalskii, AMB ; Gallus gallus, NP_ ; Cathartes aura, KFP
6 IRF8 의특징및발현분석 307 Fig. 3. Multiple sequence alignment of SsIRF8 amino acid sequence with other known orthologs from GenBank. The alignment was conducted using ClustalW. The DNA binding domain (DBD) and IRF-association domain1 (IAD1) are shown with arrows. The conserved tryptophan (W) residues in the DBD are boxed and shaded in white. thus fasciatus), (Epinephelus coioides), (Scophthalmus maximus), (Paralichthys olivaceus), (Gadus morhua), (Oncorhynchus mykiss), (Scleropages formosus), (Danio rerio), (Ctenopharyngodon idella), Gymnocypris przewalskii, (Gallus gallus), (Cathartes aura) IRF8, SsIRF8 cluster (Fig. 2). Multiple alignment IRF8 DBD IAD1 homology
7 308 양혜림ㆍ권혁재ㆍ이성도ㆍ S.D.N.K Bathige ㆍ김명진ㆍ이제희 Relative mrna expression level of SsIRF SP SK IT GL BL HK KD HT LV St MS Tissue type Relative mrna expression of SsIRF Poly I:C S.iniae LPS control 3hr 6hr 12hr 24hr 48hr 72hr Fig. 4. SsIRF8 mrna expression in different tissues (SP- spleen, SK-skin, IT-intestine, GL-gill, BL-blood, HK-head kidney, LVliver, St-stomach, MS-muscle) of the healthy black rockfish Sebastes schlegelii as measured using real time qpcr. The relative expressions of the IRF8 are shown as a ratio to IRF8 expression in the MS. Sebastes schlegelii EF1α was used as a reference gene. Data are shown as the mean (n=3) and the error bars represent the standard deviation.. SsIRF8 DBD Oplegnathus fasciatus, Gadus morhua, Oncorhynchus mykiss, Gallus gallus, Mus musculus, Homo sapiens IAD1 (Fig. 3). IRF8 lymphomyeloid-rich (Lehtonen et al., 2003; Wang and Morse, 2009). lymphomyeloid-rich,, IRF8,.,, (Holland et al., 2010; Bathige et al., 2012; Chen et al., 2012; Hu et al., 2013). SsIRF8 9 (gill, heart, head kidney, spleen, liver, intestine, kidney, testis, ovary) PBCs cdna qpcr. Elongation factor 1- (EF1 ) reference gene. qpcr mrna. 256,,,,,, (Fig. 4). myeloid progenitor cell B, T IRF8 (Lu, 2008) SsIRF8, lymphomyeloid-rich, Relative IRF8 expression post Poly I:C, S.iniae, LPS challenge Fig. 5 Gene expression of SsIRF8 in the spleen after challenge with either poly I:C or S. iniae or LPS. All data were normalized to that of Sebastes schlegelii EF1α. The time course of IRF8 mrna expression in the spleen at 0, 3, 6, 12, 24, 48, 72 hr post injection is shown. Data are presented as the mean (n=3) and the error bars represent the standard deviation. Asterisks () represent significant differences (P<0.05).,,, (Hu et al., 2013). mrna PBS, S. iniae, poly I:C, LPS qrt-pcr. Poly I:C (P<0.05). S. iniae 12 12, 24, (P<0.05). LPS 6 12 (P<0.05). SsIRF8 (Fig. 5)., SsIRF8 LPS- TLR4 polyinosinic:polycytidylic acid (poly I:C)-TLR3 Toll-like receptor IFN- signaling pathways IRF8 (Zhao et al., 2006). poly I:C,. poly I:C pathogen associated molecular pattern (PAMP) (Perdiguero and Esteban, 2009; Hu et al., 2013), poly I:C-activated signaling pathways SsIRF8 (Gabriele and Ozato, 2007; Tailor et al., 2007). SsIRF8
8 IRF8 의특징및발현분석 309 SsIRF8 SsIRF8 IRF8 myeloid lymphoid cell. 사사 2016 ( ). References Alter-Koltunoff M, Goren S, Nousbeck J, Feng CG, Sher A, Ozato K, Azriel A and Levi BZ Innate immunity to intraphagosomal pathogens is mediated by Interferon Regulatory Factor 8 (IRF-8) that stimulates the expression of macrophage-specific Nramp1 through antagonizing repression by c-myc. J Biol Chem 283, org/ /jbc.m Barnes B, Lubyova B and Pitha PM On the role of IRF in host defense. J Interf cytokine Res 22, org/ / Bathige SDNK, Whang I, Umasuthan N, Lim BS, Park MA, Kim E, Park HC and Lee J Interferon regulatory factors 4 and 8 in rock bream, Oplegnathus fasciatus: Structural and expressional evidence for their antimicrobial role in teleosts. Fish Shellfish Immunol 33, org/ /j.fsi Chen X, Hu G, Dong X, Liu Q and Zhang S Molecular cloning and expression analysis of interferon regulatory factor 8 (IRF8) in turbot, Scophthalmus maximus. Vet Immunol Immunopathol 149, vetimm Eroshkin A and Mushegian A Conserved transactivation domain shared by interferon regulatory factors and Smad morphogens. J Mol Med 77, org/ /s Gabriele L and Ozato K The role of the interferon regulatory factor (IRF) family in dendritic cell development and function. Cytokine Growth Factor Rev 18, dx.doi.org/ /j.cytogfr Holland JW, Karim A, Wang T, Alnabulsi A, Scott J, Collet B, Mughal MS, Secombes CJ and Bird S Molecular cloning and characterization of interferon regulatory factors 4 and 8 (IRF-4 and IRF-8) in rainbow trout, Oncorhynchus mykiss. Fish Shellfish Immunol 29, org/ /j.fsi Honda K and Taniguchi T IRFs: master regulators of signalling by Toll-like receptors and cytosolic pattern-recognition receptors. Nat Rev Immunol 6, org/ /nri1900. Hu G, Chen X, Gong Q, Liu Q, Zhang S and Dong X Structural and expression studies of interferon regulatory factor 8 in Japanese flounder, Paralichthys olivaceus. Fish Shellfish Immunol 35, org/ /j.fsi Huang B, Qi ZT, Xu Z and Nie P Global characterization of interferon regulatory factor (IRF) genes in vertebrates: glimpse of the diversification in evolution. BMC Immunol 11, Inkpen SM, Hori TS, Gamperl AK, Nash GW and Rise ML Characterization and expression analyses of five interferon regulatory factor transcripts (Irf4a, Irf4b, Irf7, Irf8, Irf10) in Atlantic cod (Gadus morhua). Fish Shellfish Immunol 44, Lee CH, Melchers M, Wang H, Torrey TA, Slota R, Qi CF, Kim JY, Lugar P, Kong HJ, Farrington L, van der Zouwen B, Zhou JX, Lougaris V, Lipsky PE, Grammer AC and Morse HC Regulation of the germinal center gene program by interferon (IFN) regulatory factor 8/IFN consensus sequence-binding protein. J Exp Med 203, dx.doi.org/ /jem Lehtonen A, Lund R, Lahesmaa R, Julkunen I, Sareneva T and Matikainen S IFN-α and IL-12 activate IFN regulatory factor 1 (IRF-1), IRF-4, and IRF-8 gene expression in human NK and T cells. Cytokine 24, org/ /j.cyto Lu R Interferon regulatory factor 4 and 8 in B-cell development. Trends Immunol 29, org/ /j.it Mamane Y, Heylbroeck C, Génin P, Algarté M, Servant MJ, LePage C, DeLuca C, Kwon H, Lin R and Hiscott J Interferon regulatory factors: the next generation. Gene 237, Marecki S, Atchison ML and Fenton MJ Differential Expression and Distinct Functions of IFN Regulatory Factor 4 and IFN Consensus Sequence Binding Protein in Macrophages. J Immunol 163, Nehyba J, Hrdliková R, Burnside J and Bose HR A Novel Interferon Regulatory Factor (IRF), IRF-10, Has a Unique Role in Immune Defense and Is Induced by the v-rel Oncoprotein. Mol Cell Biol 22, /MCB Nguyen H, Hiscott J and Pitha PM The growing family of interferon regulatory factors. Cytokine Growth Factor Rev 8, Ozato K, Tailor P and Kubota T The interferon regulatory factor family in host defense: Mechanism of action. J Biol Chem 282, R Paun A and Pitha PM The IRF family, revisited. Biochimie
9 310 양혜림ㆍ권혁재ㆍ이성도ㆍ S.D.N.K Bathige ㆍ김명진ㆍ이제희 89, Perdiguero B and Esteban M The interferon system and vaccinia virus evasion mechanisms. J Interferon Cytokine Res 29, Pfeffer LM, Dinarello CA, Herberman RB, Williams BRG, Borden EC, Bordens EC, Bordens R, Walter MR, Nagabhushan TL, Trotta PP and Pestka S Biological properties of recombinant a-interferons: 40th anniversary of the discovery of interferons. Cancer Res 58, Stellacci E, Testa U, Petrucci E, Benedetti E, Orsatti R, Feccia T, Stafsnes M, Coccia EM, Marziali G and Battistini A Interferon regulatory factor-2 drives megakaryocytic differentiation. Biochem J 377, BJ Tailor P, Tamura T, Kong HJ, Kubota T, Kubota M, Borghi P, Gabriele L and Ozato K The Feedback Phase of Type I Interferon Induction in Dendritic Cells Requires Interferon Regulatory Factor 8. Immunity 27, org/ /j.immuni Taniguchi T, Ogasawara K, Takaoka A and Tanaka N IRF Family of transcription factors as regulatory of host defense. Annual Review Immunolo 19, /annurev.immunol Wang H and Morse HC IRF8 regulates myeloid and B lymphoid lineage diversification. Immunol Res 43, Zhang J, Li Y and Hu Y Molecular characterization and expression analysis of eleven interferon regulatory factors in half-smooth tongue sole, Cynoglossus semilaevis. Fish Shellfish Immunol 44, fsi Zhao J, Hee JK, Li H, Huang B, Yang M, Zhu C, Bogunovic M, Zheng F, Mayer L, Ozato K, Unkeless J and Xiong H IRF-8/interferon (IFN) consensus sequence-binding protein is involved in toll-like receptor (TLR) signaling and contributes to the cross-talk between TLR and IFN-γ signaling pathways. J Biol Chem 281, org/ /jbc.m
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Technical/Specs Sequencing 서비스안내 Standard Sequencing Plasmid/PCR product 를 primer 를이용하여고객이원하는 region 을분석하는서비스이며, 대부분의 normal 샘플에대해 최적화되어있습니다. Full Len
Technical/Specs Sequencing 서비스안내 Standard Sequencing Plasmid/PCR product 를 primer 를이용하여고객이원하는 region 을분석하는서비스이며, 대부분의 normal 샘플에대해 최적화되어있습니다. Full Length Sequencing (Primer walking) 길이가긴 insert 를포함한 plasmid
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