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Antimicrobial Peptides in Mycobacterial Infection,Dong Min Shin and Eun Kyeong Jo. Figure 1 A schematic diagram for the role of antimicrobial peptides such as cathelicidins and defensins in host immune system Left diagram. shows biological effects of cathelicidin hCAP 18 LL 37 in immunity hCAP 18 LL 37 is synthesized and released from epithelial cells in response. to microbial infection or physical injury hCAP 18 LL 37 participates in the recruitment of neutrophils and other circulating cells including. monocytes macrophages at sites of infection by chemotaxis through secretion of several cytokines chemokines Release of hCAP 18 LL 37 from. keratinocytes results in induced wound healing Also they contribute to direct killing activity against invading pathogens and to indirect. antimicrobial activity by promoting autophagy activation maturation in monocytes macrophages Right diagram shows immunological functions. of defensins in various immune cells Defensins induced by various physiological sitimuli including TLRs or infection defensins are synthesized. and released from monocytes macrophages or neutrophils eosinophils whereas defensins are synthesized and released from not only their. cells but also DCs airway epitheliums or skin Released peptides have direct antimicrobial killing effects and they also have indirect killing. effects by interacting with various target cells and tissues to promote secondary responses that may be crucial for modulating inflammation. the recruitment of immune cells and activation maturation of several type of immune cell. to be clarified Thus understanding the molecular mecha activated macrophages and assist in elimination of ingested. nisms of expression of antimicrobial peptides and their role bacteria 4 5 Antimicrobial peptides such as cathelicidins. as immune modulators during the host innate response to and defensins play a crucial role in biological processes in. mycobacteria will help in the design of new therapies against cluding antimicrobial activities and immunomodulatory func. tuberculosis Due to the increasing global incidence of multi tions summarized in Fig 1 Cathelicidins are bipartite mole. drug resistant tuberculosis peptide derived microbicides rep cules consisting of an N terminal cathelin domain and a C ter. resent promising candidate therapeutics in the struggle against minal domain which has antimicrobial activity 12 13 The. resistant mycobacteria N terminal cathelin domain is known as a hallmark of the in. tracellular storage part of cathelicidins 12 13 Cathelicidins. GENERAL OVERVIEW OF ANTIMICROBIAL show constitutive and or inducible expression in various cells. PEPTIDES PROTEINS and tissues 12 13 Their tissue cell specific expression is. regulated by several stimuli including infection of microbes. Antimicrobial defense peptides proteins can be produced by inflammatory cytokines 13 15 Many studies reported that. 246 IMMUNE NETWORK http www ksimm or kr Volume 11 Number 5 October 2011. Antimicrobial Peptides in Mycobacterial Infection,Dong Min Shin and Eun Kyeong Jo. hCAP 18 LL 37 contributes to elimination of bacteria 14 sys striction of mycobacterial growth 28 Additionally the neu. temic protection against microbial invasion 13 chemo trophil peptides cathelicidin LL 37 and lipocalin2 Lcn2 also. taxis attraction through secretion of several cytokines chemo known as neutrophil gelatinase associated lipocalin NGAL. kines 16 wound healing 16 and autophagy activation ma contributed to inhibition of mycobacterial growth and im. turation 17 mune defense against tuberculosis 28 Earlier studies show. Defensins are antimicrobial cytotoxic peptides which con ed that the synthetic peptide LL 37 had broad antimicrobial. tain 29 35 amino acid residues including 6 invariant cys activity in airway epithelial cells of the lung 29 and in bron. teine residues 18 Defensins are expressed by various phys choalveolar lavage fluids 30 Recently we reported that M. iological biological stimuli including TLRs or infection 18 ulcerans infection significantly induces antimicrobial peptide. The antimicrobial spectrum of defensins include not only LL 37 in human primary keratinocytes via TLR2 and. gram positive microbes but also gram negative bacteria in Dectin 1 dependent pathways 31 These reports emphasize. cluding mycobacteria Treponema pallidum fungi and some a role for LL 37 in the early innate response to mycobacteria. viruses 18 20 Defensins exert nonspecific antimicrobial cy In mycobacterial infection of human mononuclear cells. totoxic activity against mammalian target cells and micro LL 37 is induced in a vitamin D dependent manner and plays. organisms 18 In addition to their antimicrobial cytotoxic an important role in inhibition of intracellular mycobacteria. properties some defensins act as opsonins 18 contribute through NADPH oxidase 2 dependent mechanisms 6 32. to selective chemo attractants for monocytes 18 DCs and Similarly M bovis bacillus Calmette Gu rin BCG induced. T cells 21 and promote to wound healing 8 22 23 and up regulation of the antimicrobial peptide cathelicidin LL 37. regulation of inflammation 24 25 in human epithelial cells was dependent on NADPH ROS sig. naling pathways 33 Of note vitamin D is crucial for the. CATHELICIDIN hCAP 18 LL 37 regulation of LL 37 induction which can be expressed in hu. man monocytes and respiratory epithelial cells through con. The cathelicidin family a key member of host defense pep version of vitamin D into its active metabolites 6 7. tide families is derived from leukocytes and epithelial cells Recently emerging roles of autophagy in the regulation of. and has an important role in elimination of pathogenic mi innate immune functions have been reported 34 Impor. crobes 13 15 26 Various inflammatory or infectious stimuli tantly the autophagy pathway has been known to be a key. can induce cathelicidin LL 37 which then exhibits anti defensive mechanism to eliminate M tuberculosis through. microbial activity against a number of bacteria and fungi 12 phagosomal maturation 35 Our recent data have shown. 14 22 hCAP 18 LL 37 is currently the only identified human that vitamin D3 actively induces autophagy in human mono. cathelicidin and profoundly affects multiple biological and cytes and inhibits intracellular mycobacterial growth through. pathological conditions 14 26 Cathelicidins contain a con increased autophagosomal maturation 17 In this study we. served N terminal cathelin domain and a variable C terminal found that LL 37 plays an important role in the induction and. cationic antimicrobial domain that becomes active and has an maturation of autophagy pathways activated by vitamin D3. timicrobial activity The mature peptide LL 37 comprises the in human monocytes 17 LL 37 regulated the transcriptional. C terminal portion and is expressed in various cell and tissue expression of the autophagy related genes beclin 1 and atg 5. types including neutrophils monocytes keratinocytes lym via C EBP and MAPK activation 17 In addition both de. phocytes and epithelial cells of the skin testis and the gas fensin 4 and cathelicidin are induced by distinct pathways. trointestinal and respiratory tracts 12 15 in human monocytes but cooperate to activate the TLR2 1. Accumulating evidence supports an early defensive role for mediated antimycobacterial activity 9 Furthermore recent. LL 37 at various sites During mycobacterial infection the studies have shown that the mycobacterial lipoprotein LpqH. highest expression of LL 37 was observed in alveolar macro actively induces autophagy through functional vitamin D re. phages 27 Other studies showed the importance of neu ceptor signaling and following induction of LL 37 dependent. trophils in host defense against mycobacteria When the over pathways 36. all immunity of blood cells to mycobacterial infection was Besides autophagy regulation additional novel functions of. evaluated in tuberculosis contactors neutrophil counts were LL 37 have been reported regulation of chemoattraction in. associated with a high risk of tuberculosis infection and re hibition of apoptosis wound healing angiogenesis and re. IMMUNE NETWORK http www ksimm or kr Volume 11 Number 5 October 2011 247. Antimicrobial Peptides in Mycobacterial Infection,Dong Min Shin and Eun Kyeong Jo. lease of cytokines chemokines 15 16 LL 37 can also func 44 Moreover a protective role for defensin against my. tion as an immune regulator mediating M tuberculosis in cobacterial infection has been reported in human eosinophils. duced ROS release and production of pro inflammatory cyto 19 defensin is induced in eosinophils upon stimulation. kines and chemokines 32 In mice the only cathelicidin with M bovis BCG and lipomannan and shows a synergistic. CRAMP was reported and structurally similar but shorter effect with eosinophil cationic protein on mycobacterial. than human LL 37 12 Both human LL 37 and murine growth inhibition 19. CRAMP has a similar pattern of tissue distribution and bio The human defensins HBD 1 and HBD 2 are predom. logical function 15 For example both LL 37 and CRAMP inately expressed at epithelial sites and less well defined than. have been shown to be chemotactic for various immune cells defensin family 23 25 Both HBD 1 and HBD 2 has bac. including neutrophils monocytes macrophages and T cells tericidal activity against both Gram positive and Gram neg. 37 38 Further increasing evidence indicates that various cy ative acteria 23 25 At least six HBD 1 isoforms range in. tokines and signals affect induction of cathelicidin expression length from 36 to 47 amino acids have been identified in. 16 The Th1 cytokine IFN up regulates whereas the Th2 urine whereas a single isoform of HBD 2 41 amino acids. cytokine IL 4 down regulates TLR2 1 mediated induction of in length has been isolated from respiratory epithelial secre. cathelicidin 39 In this way cell mediated immune re tions and saliva 23 While HBD 1 is constitutively ex. sponses can cross talk with innate immune pathways via cath pressed HBD 2 is upregulated during bacterial infection or. elicidin and other AMPs 39 Collectively these data suggest in response to endogenous inflammatory cytokines 23 45. that cathelicidin LL 37 exerts not only direct antimicrobial ac 46 suggesting a role for HBD 2 in regulation of antimicrobial. tivity but is also an important immune modulator of autoph and inflammatory responses During mycobacterial infection. agy regulation during mycobacterial infection HBD 2 can be transported into mycobacteria containing mac. rophage phagosomes to exert mycobactericidal and myco. DEFENSINS bacteristatic activity 20 In airway epithelial cells HBD 2. mRNA is induced by M bovis BCG infection and is upregu. Human defensins constitute a large portion of the pulmonary lated by TNF produced by M bovis BCG infected cells 46. innate host defense system Earlier studies showed that de HBD 2 expression is also triggered by bacterial LPS TLR4. fensins are present in high concentrations on respiratory epi stimulation through a CD14 dependent mechanism and ulti. thelia and selectively target microbial structures 40 Additio mately results in activation of NF B 24. nally defensins function as signaling molecules which link Recent studies have emphasized the role of HBD4 in the. the adaptive immune system to invader microorganisms 40 innate immune defense against mycobacteria 9 35 TLR2 1. Similar to cathelicidins precursors of defensins that contain mediated IL 1 is required for up regulation of HBD4 ex. a characteristic sheet rich fold and a framework of six di pression which has antimicrobial activity against intracellular. sulfide linked cysteines require proteolytic cleavage for anti mycobacterial infection 9 Moreover intratracheal admin. microbial activity 41 The small 3 5 kDa human cationic istration of l isoleucine into mice infected with the antibiotic. defensins are a delineated family of effectors of host defense sensitive strain H37Rv and a multidrug resistant clinical isolate. inflammation and cytotoxicity 18 Defensins are divided in significantly up regulated defensins 3 and 4 and inhibited. to three subfamilies and defensins 25 Six bacillary loads 47 M bovis BCG mediated expression of. defensins four human defensins HBD1 4 reviewed in HBD2 was found to be regulated by the protein kinase C. Ref 25 and additional defensin gene clusters have been PKC JNK and PI3K Akt pathways in airway epithelial cells. identified by computational analysis 42 46 Of interest more highly virulent M bovis strains exhibit. High throughput studies using microarray analyses of gene lower murine defensin 4 expression and vice versa during. expression profiles of PBMCs from patients with tuberculosis early infection 48 In experimental tuberculosis initial ex. and M tuberculosis infected healthy donors found that the ef pression of murine defensin 3 and defensin 4 in airway. fector molecules defensin 1 3 and 4 are upregulated in epithelial cells was correlated with temporary control of my. patients with tuberculosis 43 Human defensins show syn cobacterial growth 49 Additionally high and stable pro. ergy with antituberculous drugs thus suggesting that they duction of mouse defensins mBD3 and mBD4 during la. may be a promising adjunct to antituberculous chemotherapy tent infection is associated with long term control of myco. 248 IMMUNE NETWORK http www ksimm or kr Volume 11 Number 5 October 2011. Antimicrobial Peptides in Mycobacterial Infection,Dong Min Shin and Eun Kyeong Jo. Table I The roles of AMPs in immune system,AMPs Expressed cells organs Functions References.
hCAP 18 LL 37 Neutrophails monocytes macrophages Elimination of pathogenic microbes e g Mycobacteria fungi Ref 14 26. keratinov cytes lymphocytes epithelial Induction and maturation of auotphagy Ref 17. cells and bronchoalveolar lavage fluids Inhibition of apoptosis Ref 16. Regulation of chemoattraction Ref 16,Release of cytokines chemokines Ref 16. Wound healing angiogenesis Ref 16, Up regulation of Th1 cytokine e g IFN down refulation of Ref 37. Th2 cytokine e g IL 4,Defensin Eosinophils PBMCs DCs T cells. monocytes macrophages airway,epithelial cells,defensin Up regulation in TB patients Ref 41. Protective effects against mycobacteria in eosinophils Ref 19. defensin Mycobactericidal and Mycobacteristatic activity Ref 20. Regulation of Inflammation e g TNF Ref 44,Activation of NF B Ref 24.
Chemotatic effects in immature DCs and T cells Ref 21. defensin Still unknown yet, Hepcidin hepatocytes macrophages Homeostatic regulation of iron absorption iron recycling and Ref 55 56. iron mobilization, Inhibition of invasion of microbes and tumor cells Ref 59. Direct antimicrobial activity Ref 10,bacterial proliferation 49 HEPCIDIN. Similar to hCAP 18 LL 37 HBD 2 plays roles other than di. rect antimicrobial action These include chemotactic roles for Hepcidin is a cationic amphipathic bactericidal peptide pri. immature dendritic cells and memory T cells through a che marily produced in the liver and acts as a homeostatic regu. mokine receptor CCR6 dependent mechanism This mecha lator of iron absorption recycling and mobilization Its ex. nism promotes adaptive immune responses by recruiting im pression is markedly induced during infectious and in. mune cells to sites of microbial invasion 21 Unlike and flammatory conditions 57 58 Hepcidin is synthesized by iron. defensins defensins are found in some non human pri loading and cytokine IL 6 and decreased by anemia and. mates but not in humans gorillas bonobos or chimpanzees hypoxia The major mechanism of hepcidin function is. 50 Tang et al first isolated a trisulfide containing anti thought to be related to regulation of transmembrane iron. microbial peptide termed rhesus theta defensin 1 RTD 1 transport through binding with ferroportin an iron exporter. from granules of neutrophils and monocytes of the rhesus expressed in hepatocytes and macrophages 59 60 The re. macaque 51 Although defensins have antimicrobial ac sulting decrease in extracellular iron concentrations probably. tivity against diverse pathogens 51 53 especially viruses makes less available for invading microorganisms and tumor. 54 56 there is as yet no evidence that defensins are cells thereby contributing to host defense and controlling. involved in defense against mycobacterial infection The roles chronic diseases 57 58 61 As a novel mediator of innate im. of antimicrobial peptides in mycobacterial infection are sum munity hepcidin and related therapeutics are promising candi. marized in Table I Future studies will reveal the multiple dates for the treatment of various diseases such as hemochro. roles of various human defensins in the regulation of immune matosis and anemia from chronic inflammation 57 58 61. responses and host defense against mycobacterial infection Hepcidin is expressed in macrophages after infection with. the intracellular pathogens M avium and M tuberculosis. IMMUNE NETWORK http www ksimm or kr Volume 11 Number 5 October 2011 249. Antimicrobial Peptides in Mycobacterial Infection,Dong Min Shin and Eun Kyeong Jo. Stimulation of macrophages with mycobacteria and IFN CONFLICTS OF INTEREST. synergistically induced hepcidin mRNA and protein which lo. calized to the mycobacteria containing phagosomes 10 The author have no financial conflict of interest. Additionally hepcidin possesses direct antimicrobial activity. and causes damage to M tuberculosis 10 Further inves REFERENCES. tigation of the signaling mechanisms responsible for hepcidin. 1 WHO Global Tubercalosis Control 2010 Geneva WHO, mRNA expression showed that STAT1 and NF B activation.
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lies have been reported to exhibit antimicrobial activities and active vitamin D to its active form potential effects on host. immunomodulatory functions These peptides are produced defense J Immunol 181 7090 7099 2008. in different types of innate immune cells such as macro 8 Cole AM Waring AJ The role of defensins in lung biology. and therapy Am J Respir Med 1 249 259 2002, phages neutrophils keratinoyctes and epithelial cells As a 9 Liu PT Schenk M Walker VP Dempsey PW Kanchana. bridge between the innate and adaptive immune responses poomi M Wheelwright M Vazirnia A Zhang X Steinmeyer. cathelicidin may contribute to host antibacterial defenses and A Z gel U Hollis BW Cheng G Modlin RL Convergence. of IL 1beta and VDR activation pathways in human, dampen harmful inflammation Furthermore antimycobac TLR2 1 induced antimicrobial responses PLoS One 4 e5. terial immune defense is linked to cathelicidin expression and 810 2009. its role in mediating autophagy IL 1 a crucial cytokine in 10 Sow FB Florence WC Satoskar AR Schlesinger LS. antimycobacterial defense is required for defensin expre Zwilling BS Lafuse WP Expression and localization of hep. cidin in macrophages a role in host defense against. ssion which is critical for innate immunity to mycobacteria tuberculosis J Leukoc Biol 82 934 945 2007. The accumulating data will enable development of ther 11 Beisswenger C Bals R Functions of antimicrobial peptides. apeutic options and innovative antibiotics derived from host in host defense and immunity Curr Protein Pept Sci 6. 255 264 2005,antimicrobial peptides, 12 Lehrer RI Ganz T Cathelicidins a family of endogenous. antimicrobial peptides Curr Opin Hematol 9 18 22 2002. ACKNOWLEDGEMENTS 13 Zanetti M The role of cathelicidins in the innate host de. fenses of mammals Curr Issues Mol Biol 7 179 196 2005. 14 Fahy RJ Wewers MD Pulmonary defense and the human. We thank coworkers and students for many fruitful cathelicidin hCAP 18 LL 37 Immunol Res 31 75 89 2005. discussions This work was supported by the Korea Science 15 Zanetti M Cathelicidins multifunctional peptides of the in. Engineering Foundation through the Infection Signaling nate immunity J Leukoc Biol 75 39 48 2004. Network Research Center R13 2007 020 01000 0 at Chung 16 Lai Y Gallo RL AMPed up immunity how antimicrobial. peptides have multiple roles in immune defense Trends. nam National University We apologize to colleagues whose Immunol 30 131 141 2009. work and publications could not be referenced owing to 17 Yuk JM Shin DM Lee HM Yang CS Jin HS Kim KK Lee. space constraints ZW Lee SH Kim JM Jo EK Vitamin D3 induces autophagy. 250 IMMUNE NETWORK http www ksimm or kr Volume 11 Number 5 October 2011. Antimicrobial Peptides in Mycobacterial Infection,Dong Min Shin and Eun Kyeong Jo. in human monocytes macrophages via cathelicidin Cell 32 Yang CS Shin DM Kim KH Lee ZW Lee CH Park SG. Host Microbe 6 231 243 2009 Bae YS Jo EK NADPH oxidase 2 interaction with TLR2 is. 18 Lehrer RI Lichtenstein AK Ganz T Defensins anti required for efficient innate immune responses to mycobac. microbial and cytotoxic peptides of mammalian cells Annu teria via cathelicidin expression J Immunol 182 3696 3705. Rev Immunol 11 105 128 1993 2009, 19 Driss V Legrand F Hermann E Loiseau S Guerardel Y 33 M ndez Samperio P P rez A Torres L Role of reactive.
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OM Oppenheim JJ Beta defensins linking innate and fected macrophages Cell 119 753 766 2004. adaptive immunity through dendritic and T cell CCR6 36 Shin DM Yuk JM Lee HM Lee SH Son JW Harding CV. Science 286 525 528 1999 Kim JM Modlin RL Jo EK Mycobacterial lipoprotein acti. 22 De Smet K Contreras R Human antimicrobial peptides de vates autophagy via TLR2 1 CD14 and a functional vitamin. fensins cathelicidins and histatins Biotechnol Lett 27 D receptor signalling Cell Microbiol 12 1648 1665 2010. 1337 1347 2005 37 De Yang Chen Q Schmidt AP Anderson GM Wang JM. 23 O Neil DA Porter EM Elewaut D Anderson GM Eckmann Wooters J Oppenheim JJ Chertov O LL 37 the neutrophil. L Ganz T Kagnoff MF Expression and regulation of the granule and epithelial cell derived cathelicidin utilizes. human beta defensins hBD 1 and hBD 2 in intestinal formyl peptide receptor like 1 FPRL1 as a receptor to che. epithelium J Immunol 163 6718 6724 1999 moattract human peripheral blood neutrophils monocytes. 24 Becker MN Diamond G Verghese MW Randell SH CD14 and T cells J Exp Med 192 1069 1074 2000. dependent lipopolysaccharide induced beta defensin 2 ex 38 Kurosaka K Chen Q Yarovinsky F Oppenheim JJ Yang. pression in human tracheobronchial epithelium J Biol D Mouse cathelin related antimicrobial peptide chemo. Chem 275 29731 29736 2000 attracts leukocytes using formyl peptide receptor like. 25 Oppenheim JJ Biragyn A Kwak LW Yang D Roles of anti 1 mouse formyl peptide receptor like 2 as the receptor and. microbial peptides such as defensins in innate and adaptive acts as an immune adjuvant J Immunol 174 6257 6265. immunity Ann Rheum Dis 62 Suppl 2 ii17 21 2003 2005. 26 Kai Larsen Y Agerberth B The role of the multifunctional 39 Edfeldt K Liu PT Chun R Fabri M Schenk M Wheelwright. peptide LL 37 in host defense Front Biosci 13 3760 3767 M Keegan C Krutzik SR Adams JS Hewison M Modlin. 2008 RL T cell cytokines differentially control human monocyte. 27 Rivas Santiago B Hernandez Pando R Carranza C Juarez antimicrobial responses by regulating vitamin D meta. 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Antibacterial components in bronchoalveolar lavage fluid and disease caused by Mycobacterium tuberculosis J Mol. from healthy individuals and sarcoidosis patients Am J Med Berl 85 613 621 2007. Respir Crit Care Med 160 283 290 1999 44 Kalita A Verma I Khuller GK Role of human neutrophil. 31 Lee HM Shin DM Choi DK Lee ZW Kim KH Yuk JM peptide 1 as a possible adjunct to antituberculosis chemo. Kim CD Lee JH Jo EK Innate immune responses to therapy J Infect Dis 190 1476 1480 2004. Mycobacterium ulcerans via toll like receptors and dectin 1 45 Harder J Meyer Hoffert U Teran LM Schwichtenberg L. in human keratinocytes Cell Microbiol 11 678 692 2009 Bartels J Maune S Schr der JM Mucoid Pseudomonas aer. IMMUNE NETWORK http www ksimm or kr Volume 11 Number 5 October 2011 251. Antimicrobial Peptides in Mycobacterial Infection,Dong Min Shin and Eun Kyeong Jo. uginosa TNF alpha and IL 1beta but not IL 6 induce hu cytes isolation synthesis antimicrobial activities and bac. man beta defensin 2 in respiratory epithelia Am J Respir terial binding properties of the cyclic peptides J Biol Chem. Cell Mol Biol 22 714 721 2000 277 3079 3084 2002, 46 M ndez Samperio P Miranda E Trejo A Mycobacterium 54 Brandt CR Akkarawongsa R Altmann S Jose G Kolb AW. bovis Bacillus Calmette Gu rin BCG stimulates human be Waring AJ Lehrer RI Evaluation of a theta defensin in a. ta defensin 2 gene transcription in human epithelial cells Murine model of herpes simplex virus type 1 keratitis. Cell Immunol 239 61 66 2006 Invest Ophthalmol Vis Sci 48 5118 5124 2007. 47 Rivas Santiago CE Rivas Santiago B Le n DA Casta eda 55 Wang W Cole AM Hong T Waring AJ Lehrer RI Retrocy. Delgado J Hern ndez Pando R Induction of defensins clin an antiretroviral theta defensin is a lectin J Immunol. by l isoleucine as novel immunotherapy in experimental 170 4708 4716 2003. murine tuberculosis Clin Exp Immunol 164 80 89 2011 56 Yang C Boone L Nguyen TX Rudolph D Limpakarnjanar. 48 Aguilar Le n D Zum rraga MJ Jim nez Oropeza R Gioffr at K Mastro TD Tappero J Cole AM Lal RB. AK Bernardelli A Orozco Est vez H Cataldi AA Theta Defensin pseudogenes in HIV 1 exposed persistently. Hern ndez Pando R Mycobacterium bovis with different seronegative female sex workers from Thailand Infect. genotypes and from different hosts induce dissimilar im Genet Evol 5 11 15 2005. munopathological lesions in a mouse model of tuber 57 Ganz T Hepcidin and its role in regulating systemic iron. culosis Clin Exp Immunol 157 139 147 2009 metabolism Hematology Am Soc Hematol Educ Program. 49 Rivas Santiago B Sada E Tsutsumi V Aguilar Leon D 29 35 2006. Contreras JL Hernandez Pando R beta Defensin gene ex 58 Ganz T Hepcidin a peptide hormone at the interface of. pression during the course of experimental tuberculosis innate immunity and iron metabolism Curr Top Microbiol. infection J Infect Dis 194 697 701 2006 Immunol 306 183 198 2006. 50 Cole AM Wang W Waring AJ Lehrer RI Retrocyclins us 59 Ganz T Nemeth E Iron sequestration and anemia of. ing past as prologue Curr Protein Pept Sci 5 373 381 2004 inflammation Semin Hematol 46 387 393 2009. 51 Tang YQ Yuan J Osapay G Osapay K Tran D Miller CJ 60 Nemeth E Ganz T The role of hepcidin in iron meta. Ouellette AJ Selsted ME A cyclic antimicrobial peptide pro bolism Acta Haematol 122 78 86 2009. duced in primate leukocytes by the ligation of two trun 61 Atanasiu V Manolescu B Stoian I Hepcidin the link be. cated alpha defensins Science 286 498 502 1999 tween inflammation and anemia in chronic renal failure. 52 Leonova L Kokryakov VN Aleshina G Hong T Nguyen Rom J Intern Med 44 25 33 2006. T Zhao C Waring AJ Lehrer RI Circular minidefensins and 62 Sow FB Alvarez GR Gross RP Satoskar AR Schlesinger. posttranslational generation of molecular diversity J LS Zwilling BS Lafuse WP Role of STAT1 NF kappaB. Leukoc Biol 70 461 464 2001 and C EBPbeta in the macrophage transcriptional regu. 53 Tran D Tran PA Tang YQ Yuan J Cole T Selsted ME lation of hepcidin by mycobacterial infection and IFN. Homodimeric theta defensins from rhesus macaque leuko gamma J Leukoc Biol 86 1247 258 2009. 252 IMMUNE NETWORK http www ksimm or kr Volume 11 Number 5 October 2011.

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