(1.Animal Disease and Public Security Academician Workstation of Henan Province/The Key Lab of Animal Disease and Public Health，Henan University of Science and Technology，Luoyang 471003，China；2.Laboratory for Animal Health，ANSES，Maisons-Alfort 94706，France；3.Key Laboratory of Zoonisis of Ministry of Education，Institute of Zoonosis，Jilin University，Changchun 130062，China)
LIAO Cheng-shui，BOIREAU Pascal，LIU Ming-yuan，CHENG Xiang-chao. Research Progress on Extracellular Traps：the Third kinds of Defence Mechanisms of Innate Immune Cells[J]. ACTA VETERINARIA ET ZOOTECHNICA SINICA, 2016, 47(9): 1768-1774.
［1］DERETIC V.Autophagy in immunity and cell-autonomous defense against intracellular microbes［J］.Immunol Rev，2011，240(1)：92-104.
［2］BROWN E J.Phagocytosis［J］.Bioessays，1995，17(2)：109-117.
［3］BRINKMANN V，REICHARD U，GOOSMANN C，et al. Neutrophil extracellular traps kill bacteria［J］.Science，2004，303(5663)：1532-1535.
［4］VON KÖCKRITZ-BLICKWEDE M，BLODKAMP S，NIZET V.Interaction of bacterial exotoxins with neutrophil extracellular traps：Impact for the infected host［J］.Front Microbiol，2016，7：402.
［5］WANG Y，LI M，STADLER S，et al.Histone hypercitrullination mediates chromatin decondensation and neutrophil extracellular trap formation［J］.J Cell Biol，2009，184(2)：205-213.
［6］FUCHS T A，ABED U，GOOSMANN C，et al.Novel cell death program leads to neutrophil extracellular traps［J］.J Cell Biol，2007，176(2)：231-241.
［7］YIPP B G，PETRI B，SALINA D，et al.Infection-induced NETosis is a dynamic process involving neutrophil multitasking in vivo［J］.Nat Med，2012，18(9)：1386-1393.
［8］VON KÖCKRITZ-BLICKWEDE M，GOLDMANN O，THULIN P，et al.Phagocytosis-independent antimicrobial activity of mast cells by means of extracellular trap formation［J］.Blood，2008，111(6)：3070-3080.
［9］YOUSEFI S，GOLD J A，ANDINA N，et al.Catapult-like release of mitochondrial DNA by eosinophils contributes to antibacterial defense［J］.Nat Med，2008，14(9)：949-953.
［10］YOUSEFI S，MORSHED M，AMINI P，et al.Basophils exhibit antibacterial activity through extracellular trap formation［J］.Allergy，2015，70(9)：1184-1188.
［11］LIU P，WU X，LIAO C，et al.Escherichia coli and Candida albicans induced macrophage extracellular trap-like structures with limited microbicidal activity［J］.PLoS One，2014，9(2)：e90042.
［12］CHUAMMITRI P，OSTOJIC＇ J，ANDREASEN C B，et al.Chicken heterophil extracellular traps (HETs)：novel defense mechanism of chicken heterophils［J］.Vet ImmunolImmunopathol， 2009，129(1-2)：126-131.
［13］ALTINCICEK B，STÖTZEL S，WYGRECKA M，et al.Host-derived extracellular nucleic acids enhance innate immune responses，induce coagulation，and prolong survival upon infection in insects［J］.J Immunol， 2008，181(4)：2705-2712.
［14］GOLDMANN O，MEDINA E.The expanding world of extracellular traps：not only neutrophils but much more［J］. Front Immuno，2013，3：420.
［15］STOIBER W，OBERMAYER A，STEUNBACHER P，et al.The role of reactive oxygen species (ROS) in the formation of extracellular traps (ETs) in Humans［J］.Biomolecules，2015，5(2)：702-723.
［16］AWASTHI D，NAGARKOTI S，KUMAR A，et al.Oxidized LDL induced extracellular trap formation in human neutrophils via TLR-PKC-IRAK-MAPK and NADPH-oxidase activation［J］.Free Radic Biol Med，2016，93：190-203.
［17］BIANCHI M，HAKKIM A，BRINKMANN V，et al.Restoration of NET formation by gene therapy in CGD controls aspergillosis［J］.Blood，2009，114(13)：2619-2622.
［18］LIM M B，KUIPER J W，KATCHKY A，et al.Rac2 is required for the formation of neutrophil extracellular traps［J］.J Leukocyte Biol，2011，90(4)：771-776.
［19］RAAD H，PACLET M H，BOUSSETTA T，et al.Regulation of the phagocyte NADPH oxidase activity：phosphorylation of gp91phox/NOX2 by protein kinase C enhances its diaphorase activity and binding to Rac2，p67phox，and p47phox［J］.FASEB J，2009，23(4)：1011-1022.
［20］YOST C C，CODY M J，HARRIS E S，et al.Impaired neutrophil extracellular trap (NET) formation：a novel innate immune deficiency of human neonates［J］.Blood，2009，113(25)：6419-6427.
［21］PARKER H，DRAGUNOW M，HAMPTON M B，et al.Requirements for NADPH oxidase and myeloperoxidase in neutrophil extracellular trap formation differ depending on the stimulus［J］.J Leukocyte Biol，2012，92(4)：841-849.
［22］PILSCZEK F H，SALINA D，POON K K，et al.A novel mechanism of rapid nuclear neutrophil extracellular trap formation in response to Staphylococcus aureus［J］.J Immunol，2010，185(12)：7413-7425.
［23］GABRIEL C，MCMASTER W R，GIRARD D，et al.Leishmania donovani promastigotes evade the antimicrobial activity of neutrophil extracellular traps［J］.J Immunol，2010，185(7)：4319-4327.
［24］LI P，LI M，LINDBERG M R，et al.PAD4 is essential for antibacterial innate immunity mediated by neutrophil extracellular traps［J］.J Exp Med，2010，207(9)：1853-1862.
［25］NEELI I，KHAN S N，RADIC M.Histone deimination as a response to inflammatory stimuli in neutrophils［J］.J Immunol，2008，180(3)：1895-1902.
［26］GUPTA A K，JOSHI M B，PHILIPPOVA M，et al.Activated endothelial cells induce neutrophil extracellular traps and are susceptible to NETosis-mediated cell death［J］.FEBS Lett，2010，584(14)：3193-3197.
［27］MORSHED M，HLUSHCHUK R，SIMON D，et al.NADPH oxidase-independent formation of extracellular DNA traps by basophils［J］.J Immunol，2014，192(11)：5314-5323.
［28］UEKI S，MELO R C，GHIRAN I，et al.Eosinophil extracellular DNA trap cell death mediates lytic release of free secretion-competent eosinophil granules in humans［J］.Blood，2013，121(11)：2074-2083.
［29］BRINKMANN V，ZYCHLINSKY A.Beneficial suicide：why neutrophils die to make NETs［J］.Nat Rev Microbiol，2007，5(8)：577-582.
［30］CLARK S R，MA A C，TAVENER S A，et al.Platelet TLR4 activates neutrophil extracellular traps to ensnare bacteria in septic blood［J］.Nat Med，2007，13(4)：463-469.
［31］MCDONALD B，URRUTIA R，YIPP B G，et al.Intravascular neutrophil extracellular traps capture bacteria from the bloodstream during sepsis［J］.Cell Host Microbe，2012，12(3)：324-333.
［32］URBAN C F，REICHARD U，BRINKMANN V，et al.Neutrophil extracellular traps capture and kill Candida albicans yeast and hyphal forms［J］.Cell Microbiol，2006，8(4)：668-676.
［33］BYRD A S，O’BRIEN X M，JOHNSON C M，et al.An extracellular matrix-based mechanism of rapid neutrophil extracellular trap formation in response to Candida albicans［J］.JImmunol，2013，190(8)：4136-4148.
［34］BRUNS S，KNIEMEYER O，HASENBERG M，et al.Production of extracellular traps against Aspergillus fumigatus in vitro and in infected lung tissue is dependent on invading neutrophils and influenced by hydrophobin RodA［J］.PLoS Pathog，2010，6(4)：e1000873.
［35］DOUDA D N，JACKSON R，GRASEMANN H，et al.Innate immune collectin surfactant protein D simultaneously binds both neutrophil extracellular traps and carbohydrate ligands and promotes bacterial trapping［J］.J Immunol，2011，187(4)：1856-1865.
［36］HIRSCH J G.Bactericidal action of histone［J］.J Exp Med，1958，108(6)：925-944.
［37］LI G H，MINE Y，HINCKE M T，et al.Isolation and characterization of antimicrobial proteins and peptide from chicken liver［J］.J Pept Sci，2007，13(6)：368-378.
［38］LEHRER R I，GANZ T.Defensins of vertebrate animals［J］.Curr Opin Immunol，2002，14(1)：96-102.
［39］LEHRER R I，GANZ T.Cathelicidins：a family of endogenous antimicrobial peptides［J］.Curr Opin Hematol，2002，9(1)：18-22.
［40］ZANETTI M.Cathelicidins，multifunctional peptides of the innate immunity［J］.J Leukocyte Biol，2004，75(1)：39-48.
［41］JAILLON S，PERI G，DELNESTE Y，et al.The humoral pattern recognition receptor PTX3 is stored in neutrophil granules and localizes in extracellular traps［J］.J Exp Med，2007，204(4)：793-804.
［42］URBAN C F，ERMERT D，SCHMID M，et al.Neutrophil extracellular traps contain calprotectin，a cytosolic protein complex involved in host defense against Candida albicans［J］.PLoS Pathog，2009，5(10)：e1000639.
［43］MULCAHY H，CHARRON-MAZENOD L，LEWENZA S.Extracellular DNA chelates cations and induces antibiotic resistance in Pseudomonas aeruginosa biofilms［J］.PLoS Pathog，2008，4(11)：e1000213.
［44］MENEGAZZI R，DECLEVA E，DRI P.Killing by neutrophil extracellular traps：fact or folklore?［J］.Blood，2012，119(5)：1214-1216.
［45］WEISS S J，CURNUTTE J T，REGIANI S.Neutrophil-mediated solubilization of the subendothelial matrix：oxidative and nonoxidative mechanisms of proteolysis used by normal and chronic granulomatous disease phagocytes［J］.J Immunol，1986，136(2)：636-641.
［46］RICE W G，WEISS S J.Regulation of proteolysis at the neutrophil-substrate interface by secretory leukoprotease inhibitor［J］.Science，1990，249(4965)：178-181.
［47］MCCORMICK A，HEESEMANN L，WAGENER J，et al.NETs formed by human neutrophils inhibit growth of the pathogenic mold Aspergillus fumigatus［J］.Microbes Infect，2010，12(12-13)：928-936.
