Regulation of Type 2 Immunity by Basophils Prof. Dr. David Voehringer
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1 Regulation of Type 2 Immunity by Basophils Department of Infection Biology Institute of Clinical Microbiology, Immunology and Hygiene Outline of the presentation The concept of type 2 immunity Basophil development, turnover and homeostasis Basophil effector functions 2 Type 1 vs. Type 2 immunity Bacteria Viruses Worms Allergens 3 Type 1 immune response Th1 cells (IFN-g, TNF-a) Opsonizing Ab (IgG2a) CTL, NK cells Type 2 immune response Th2 cells (IL-4, IL-5, IL-13), IgG1 Basophils, eosinophils, mast cells 1
2 Type 2 immunity a double-edged sword Positive Protective immunity against helminths >2 billion people infected worldwide 1 million people die every year No vaccines yet Negative Allergies / Asthma ~15% of people in westernized countries suffer from allergies Increasing prevalence High costs for the health care system 4 Development of a type 2 immune response Lymph node / spleen Tissue TCR MHC II naive T cell APC antigen Th2 IgG1 IL-4 IL-13 B cell IgG1 Th2 activation IL-5 Basophils/ Mast cells FcR Eosinophils Degranulation Cytokine secretion 5 Helminths Allergens Elimination Allergy / Asthma Discovery of innate type 2 effector cells Paul Ehrlich ( ) Basophils Eosinophils Mast cells 6 2
3 Comparison between basophils, mast cells and eosinophils High affinity receptor (FcεRI) Basophils Mast cells Eosinophils h Mobilization IL-3 IL-3, SCF IL-5 7 Histamine Carboxypeptidase A IL-4 expression Condensed nucleus Maturation in bone marrow Lifespan of few days h in human but not mouse Developmental relationship Bone marrow/ Spleen GATA-2 Eosinophil precursor Eosinophil c/ebpa c/ebpa GATA-2 c/ebpa c/ebpa Basophil precursor Basophil Granulocyte/ Monocyte precursor (GMP) Basophil/Mast cell common precursor (BMCP) Mast cell precursor Tissue 8 Arinobu et al., (29) Allergology International 58: Mast cell Detection of basophils by CD49b/ staining Blood Spleen Bone marrow CD49b
4 Nippostrongylus brasiliensis infection Hookworm-like life cycle 1µm Day 1-4 Lung L3 => L5 N. brasiliensis in small intestine Day 4-1 Small Intestine L6, adult 5 L3 larvae 1 Cell count (x1 5 ) Days after infection Voehringer et al., (24) Immunity2: 267 N. brasiliensis infected lung Eosinophils MBP/DAPI Basophils CD2R3/DAPI Mast cells c-kit/b22 11 Ohnmacht, C. and Voehringer, D. (28) Blood 113: 2816 N. brasiliensis infected small intestine c-kit W-sh/W-sh mice FceRI/DAPI 2mm 12 Ohnmacht, C. and Voehringer, D. (28) Blood 113:
5 A B % Basophils of total 1.69 Analysis of basophil turnover in vivo with BrdU naive infected BM SP BL LU control e BrdU C naive BM SP infected BM SP BL LU 15h 36h 84h Ohnmacht, C. and Voehringer, D. (28) Blood 113: 2816 Activating receptors on basophils 14 Voehringer (29) Trends in Parasitology 25: 551 Activation of basophils by Antigen Affinity for : 1-1 M Isoforms: abg 2 : Mast cells, basophils ag 2 : Monocytes, eosinophils, neutrophils FceRIa b g 2 ITAM Degranulation Cytokine release 15 5
6 Cytokine instruction of basophils IL-18 IL-33 IL-3 GM-CSF TSLP IL-18Ra ST2 IL-18Rb IL1-RAcP b FcRg c 2 b c IL-3Ra TSLPR GM-CSFRa IL-7Ra MyD88 JAK2 JAK2 JAK2? 16 NFkB ERK p38 JNK Degranulation Lipid mediators Cytokines Syk PI3KHck/Lyn IL-4 IL-6 SHIP STAT5 Proliferation Survival Activation Negative feedback Pim-1 Lipid metabolism Cell adhesion Cell communication SOCS1/ SOCS3/ CIS Basophil effector functions IgG1- mediated anaphylaxis PAF -mediated chronic allergic inflammation MHC-II low Worm expulsion 17 Voehringer (29) Trends in Parasitology 25: 551 Generation and analysis of basophil-deficient mice 18 6
7 Mouse mast cell protease 8 is expressed in basophils Basophils Mast cells HPRT Mcpt-1 Mcpt-2 Mcpt-4 Mcpt-5 Mcpt-6 Mcpt-7 Mcpt-8 Mcpt-8 expression Ohnmacht et al., (21) Immunity33: 364 Constitutive deletion of basophils in Mcpt8Cre mice control Mcpt8Cre 2 CD49b e blood spleen bone marrow Ohnmacht et al., (21) Immunity33: 364 Mast cells are not deleted in Mcpt8Cre mice c-kit control Mcpt8Cre Peritoneum Mcpt8Cre Control c-kit W-sh Skin Small intestine 21 Ohnmacht et al., (21) Immunity33: 364 7
8 Effector function of basophils Th2 polarization Systemic anaphylaxis?? Chronic allergic dermatitis?? Worm expulsion 1µm 22 Are basophils required for Th2 polarization? 23 Are DCs or basophils required for Th2 polarization? Dendritic cells naive Allergen Th2 Th2? Basophils 24 8
9 Generation of DC-deficient mice CD11c-Cre R-DTA x CD11c-BAC (RP24 361C4) Cre BGHpA CD11c exon 1 Caton et al., (27) J Exp Med 24: 1653 DC mice 25 Ohnmacht et al., (29) J Exp Med 26: 549 Are DCs or basophils required for Th2 polarization? control CFSE +Transfer Papain +OVA Proliferation IL-4 expression OT-II (Thy1.1) Mcpt8Cre Footpad Popliteal LN 26 ΔDC Ohnmacht et al., (21) Immunity33: 364 DCs are required for Th2 polarization counts CFSE control Mcpt8Cre ΔDC % undivided OT-II P=.1 Thy 1.1 IL-4 control Mcpt8Cre DC % IL-4 + of OT-II P<.1 27 Ohnmacht et al., (21) Immunity33: 364 9
10 Role of innate IL-4 for Th2 polarization DC Naïve CD4+ T cell Innate helper cells Mast cell IL-4??? NKT Eos Th2 IL-4 IL-5 IL-13 IL-25 Nb IL-4eGFP Transfer naïve CD4 T cells TCR-Ca -/- Nb Analyze Th2 polarization 28 IL-4/IL-13/TCR-Ca -/- Innate IL-4/IL-13 regulates Th2 accumulation in the lung 4get CD4 in TCR-Ca -/- 4get CD4 in IL-4/IL-13/Ca -/- LYMPH NODE CD4 LUNG IL4eGFP 29 Basophils in anaphylactic responses 3 1
11 IgG1-mediated systemic anaphylaxis Basophils Tsujimura et al., (28) Immunity 28: 581 Neutrophils Jönsson et al., (211) J Clin Invest. 121: 1484 Macrophages Arias et al., (211) JACI127: Galli/Franco comment in Immunity28 Basophils are not required for systemic anaphylaxis -anti-tnp 32 Basophils in chronic allergic inflammation of the skin 33 11
12 Basophils are required for chronic allergic inflammation -anti-tnp i.v. TNP-OVA (left ear) OVA (right ear) d-1 d Analysis of ear swelling Ear swelling Wild-type mice Mcpt8Cre mice Baso-depleted mice FcRg-deficient mice Days after TNP-OVA Basophil transfer Mukai et al., (25) Immunity 23: 191 Obata et al., (27) Blood 11: 913 Ohnmacht et al., (21) Immunity33: 364 Basophil response against helminths and ticks 35 Cell counts (x1 6 ) Cell counts (x1 4 ) Primary response to N. brasiliensis Eosinophils Basophils Th2 cells days after infection control Mcpt8Cre Lung mln Ohnmacht et al., (21) Immunity33:
13 Expulsion kinetics of N. brasiliensis Number of worms in the intestine 1 5 Secondary infection Primary infection Days after infection 37 Protective immunity against N. brasiliensis Primary Infection Secondary Infection Worms / mouse nd nd Worms / mouse ND 38 Ohnmacht et al., (21) Immunity33: infection Long-lived Plasma cell Th2 IL-4 B IgG1 Basophils Ih2 IL-13 Activation of tissue cells IL-4/IL-13 FcR 39 Relm-b Mucus Smooth muscle activity Epithelial cell shedding Worm expulsion 13
14 Protective immunity against ticks 1 infestation 2 infestation 2 infestation Basophils present Basophils absent basophil Mast cell 4 Wada et al., (21) J. Clin. Invest. 12: 2867 Summary 1. Basophils have a lifespan of ~6h in steady-state conditions 2. In our hands Th2 polarization (helminths, papain, OVA/alum) depends on DCs and not on basophils 3. - or IgG1-mediated anaphylaxis occurs without basophils 4. Basophils are required for -mediated chronic allergic inflammation of the skin 5. Basophils contribute to protection against helminths and ticks 41 Acknowledgements Caspar Ohnmacht Christian Schwartz Marc Panzer Isabell Schiedewitz R. M. Locksley (UCSF) B. Reizis (Columbia University) J. Lee (Mayo Clinic Scottsdale) H. Karasuyama (University of Tokyo) David.voehringer@uk-erlangen.de 42 Universitätsklinikum Erlangen 14
15 43 15
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