Lipopolysaccharide = Lipid + Polysaccharide. The Innate Immune Response is Conserved Throughout Evolution and is Triggered by Pattern Recognition
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1 Lipopolysaccharide = Lipid + Polysaccharide The Innate Immune Response is onserved Throughout volution and is Triggered by Pattern Recognition.coli ell wall organization Lipid A Lipopolysaccharide Outer membrane Peptidoglycan Inner membrane Architecture of LPS O-specific side chain ore region Lipid A From: Beutler and Rietschel, ature Reviews Immunology 3; (2003) Diversity of Pathogen-associated Molecular Patterns (PAMPs) Innate Immune Receptors for PAMPs Toll-like receptors (TLRs) omplement ollectins (e.g., Surfactant Protein-A) Scavenger receptors Pentraxins (e.g., RP) Lectins (e.g., Dectin-1) D14 OD-like receptors (LRs) RIG-1-like receptors From: Akira et al., ell 124:783, 2006 ollectins and Innate Immune Recognition Some Functions of ollectins Globular heads bind to: Ag/Ab complexes Apoptotic cells Pathogens -reactive protein ollagen-like tails bind to: 1q receptors on phagocytes (to promote phagocytosis) 1r/1s (to initiate the classical pathway) SP-A SP-D 1q Surfactant proteins From: Wright, ature Rev. Immunol. 5:58,
2 SoS o Soo L L The Scavenger Receptor Superfamily Recognizes PAMPs OxLDL: Oxidized LDL AG: Advanced glycation end-products KY TO DOMAIS. coli LTA S.aureus LPS Gram-positive bacteria Gram-negative bacteria Bacterial DA OxLDL AG -omplement control protein (P) -Somatomedin B --type Lectin -pidermal growth factor (GF) -Partial pidermal growth factor (GF) -Potential -linked glycosylation -Potential O-linked glycosylation Innate Immune Receptors Also Trigger a Systemic Response to Infection. coli S.aureus OxLDL HDL. coli S.aureus OxLDL * * SR-A I SR-A II SR-A III MARO D36 SR-B1 SR-A SR-B dsr- I SR- D68 LOX-1 SR SR-D SR- SR-F Innate Immune Functions of Reactive Protein (RP), an Acute Phase Protein Synthesized by Hepatocytes o-localization of RP and Activated omplement in Infarcted Human Myocardium 1q omplement activation RP MBL omplement activation Phosphocholine RP 4d RP acts as a bridge between phosphocholine on bacterial targets and 1q J = jeopardized; = normal Adapted from: Black et al., J. Biol. hem. 47:48487, 2004 From: ijmeijer et al., Am. J. Pathol. 163:269, 2003 Treatment of xperimental Myocardial Infarction with a RP-binding Analog of Phosphocholine Limits Infarct Size History of ndotoxin Research 3H/HeJ Primary mutation structure of reported LPS reported TLR2 is the receptor for PG Pasteur, Koch: the germ theory of disease Pfeiffer describes endotoxin ndotoxin = LPS TF mediates Discovery LPS-induced shock of D14 The post-microbial era began with the discoveries of Koch and Pasteur (1865). Four phases of discovery are depicted: the recognition that infection is poisonous (red); search for poisons culminating in the identification of endotoxin (green); the chemical and biological characteriztaion of endotoxin (orange); the identification of the endotoxin receptor and its role in promoting the immune response (purple) LPS = TLR4 From: Kitsis and Jialal, ew ngl. J. Med. 355:513, 2006 Modified from: Beutler and Rietschel, ature Reviews Immunology 3; (2003) 2
3 A Re-interpretation of the ndotoxin Research Timeline Discovery of the F-κB signaling pathway by Toll in Drosophila by Hoffman and colleagues Use of adjuvant to stimulate the immune response Molecular basis of adjuvant discovered by Medzhitov and Janeway Infectious-non-self model of immunity described by Janeway Primitive Specificity in Target Recognition by the Innate Immune System 3H/HeJ mutation reported Primary structure of LPS reported TLR2 is the receptor for PG ndotoxin = LPS TF mediates septic shock Discovery of D14 LPS = TLR4 Modified from: Beutler and Rietschel, ature Reviews Immunology 3; (2003) Recruitment of TLR2 to Yeast Phagosomes Ligand Specificity of TLRs TLR2 Phase ontrast TLR2 From: Underhill et al., ature 401:811,
4 Specificity of TLR Transcriptional Programs TLR Signaling: Two Major Pathways artoon of major signal transduction pathways following engagement of TLRs. TLR4 is the major sensor of LPS. TLR3 recognizes dsra and is important in the anti-viral response. The IRF pathway leading to production of Type I IFs (i.e., IFα/β) is particularly prominent in a minor subset of dendritic cells (called plasmacytoid Ds ) that are the major source of these IFs in response to viral infections. From: Luster, urr. Opin. Immunol. 14:129, 2002 Do not memorize this cascade but rather appreciate that it consists of two parallel pathways, one that activates F-κB, leading to production of pro-inflammatory proteins, and one that activates the IRF pathway, leading to production of Type I IFs. From: Moynagh,Trends Immunol. 26:469, 2005 TLR Signaling: Two Major Pathways TLRs Sense Microbial Pathogens and Trigger xpression of Pro-inflammatory ytokines and hemokines artoon of major signal transduction pathways following engagement of TLRs. TLR4 is the major sensor of LPS. TLR3 recognizes dsra and is important in the anti-viral response. The IRF pathway leading to production of Type I IFs (i.e., IFα/β) is particularly prominent in a minor subset of dendritic cells (called plasmacytoid Ds ) that are the major source of these IFs in response to viral infections. Do not memorize this cascade but rather appreciate that it consists of two parallel pathways, one that activates F-κB, leading to production of most pro-inflammatory proteins, and one that activates the IRF pathway, leading to production of Type I IFs. From: Moynagh,Trends Immunol. 26:469, 2005 Adapted from: reagh and O eill, Trends Immunol. 27:352, 2006 OD Proteins: Intracellular Peptidoglycan Sensors OD-1 OD-2 ewly Recognized omponents of the Innate Immune System od Protein ARD LRR; Ligand Recognition RIK I-κB p50 p65 F-κB Polymorphisms in od-2 are associated with up to 30-40% of cases of rohn s disease (an inflammatory bowel disease) ARD, caspase-recruitment domain; LRR, leucine-rich repeat; RIK, a ARD-containing protein kinase 4
5 ytosolic Bacterial Recognition Systems and the Inflammasome ARD = caspase-recruitment domain LRR = leucine-rich repeat OD = OD2-like domain MDP = muramyl dipeptide Mutations in Pyrin, Another ARD-containing Innate Immune-like Protein, is Responsible for Familial Mediterranean Fever From: Akira et al., ell 124:783, 2006 ontrast-enhanced abdominal T from a 31 year-old patient with Familial Mediterranean Fever suffering an acute attack of abdominal pain, nausea, vomiting, and arthritis. ote mesenteric vessel with thickened mesenteric fold (white arrow). Histopathology demonstrated neutrophilic infiltrate and associated vasculitis. Treatment with an IL-1 receptor antagonist (Anakinra) resulted in prompt cessation of symptoms. Pathogenesis of Gout Uncovered in 2006: Monosodium urate crystals activate the inflammasome od-like Receptors (LRs) Sense Microbial Products, Activate the Inflammasome, and Trigger Maturation of IL-1 From: Martinon and Glimcher J. lin. Invest. 116:2073, 2006 Adapted from: reagh and O eill, Trends Immunol. 27:352, 2006 Dendritic ell Maturation The Dendritic ell and Development of The Primary Immune Response: Wisdom Through Maturity From: Mellman & Steinman, ell 106:255,
6 The Innate Immune Response Orchestrates D Trafficking to Secondary Lymphoid Organs Functional Differences Between Immature and Mature Ds xpression of R7 Migration towards T-cell zone of lymph node From: Luster, urr. Opin. Immunol. 14:129, 2002 The (Primary) Acquired Immune Response is Initiated by Innate Immune Recognition hemokines Direct Trafficking of Immune ells From: Luster, urr. Opin. Immunol. 14:129, 2002 The arly Antiviral Response: ytokines of the Innate Immune System Innate Immune Receptors for dsra ooperate to Initiate the Immune Response to RA Viruses Pathogen IL-12 IF-γ IF-γ Double-stranded RA products of virus infection bind to RIG-I or MDA5, which in turn bind to IPS-1 via ARD domain interactions. This complex then signals the activation of IKK-ε and TBK1 or other kinases to phosphorylate IRF-3, possibly through direct recruitment of signaling effectors, leading to IRF-3 dimerization, nuclear translocation and assembly onto the IF-β enhancer. IPS-1 might also signal the activation of the IKK complex via direct binding of IKK components or through recruitment of RIP-1, FADD and/or TRAF6, causing the phosphorylation of IκB, the inhibitor of FκB. Phosphorylated IκB is then ubiquitinated and targeted to the proteosome for degradation, releasing the active F-κB complex to translocate to the nucleus. During virus infection, dsra products can signal through TLR3 to activate IRF-3 and F-κB by the actions of the TRIF adaptor protein and RIP-1, respectively. From: Johnson and Gale, Trends Immunol. 27:1, 2006.B.: Do not memorize this cartoon, but appreciate how cytosolic dsra receptors (RIG-1, MDA5) and plasma membraneassociated dsra receptors (TLR3) cooperate to activate IRF- and F-κBdependent gene expression. 6
7 The Antiviral Response: a ascade of Transcriptional vents Some targets of IRFs RIG-1-like Receptors (RLRs) Sense Viral Products, Activate the IRF Pathway, and Trigger and Trigger Production of Antiviral Proteins Gene p21 IL-15 FasL IL-12 Function ell cycle arrest K cell maturation ell death Th1 immune response Multiphasic induction of murine type I IF genes can be divided into three phases. (a) The immediate early phase. Virus infection stimulates a phosphorylation cascade, leading to the activation of at least three families of transcription factors, including F-κB, AP-1 and IRF3. Activation of the IF-α promoter requires all three transcription factors. (b) IRF7 induction phase. Secretion of early IF produces an autocrine response through stimulation of the JAK-STAT pathway. Among the pathway s target genes is IRF7, itself. (c) Delayed early (amplification) phase. Many members of the IF-α gene family possess promoter binding sites for activated IRF7 and become transcriptionally active. Adapted from: reagh and O eill, Trends Immunol. 27:352, 2006 Summary 1. Innate immunity is conserved throughout evolution and is triggered by recognition of pathogenassociated molecular patterns (e.g., LPS) by pattern recognition receptors. 2. ollectins (e.g., SP-A, 1q, MBP) recognize carbohydrates on pathogen surfaces and perform multiple anti-microbial functions (e.g., opsonization). ollectins are essential for innate immunity, but also help clear apoptotic debris. 3. Members of the Scavenger Receptor superfamily recognize bacteria as well as glucose-modified proteins and oxidized lipoproteins. They are implicated in the response to infection as well as atherosclerosis and other degenerative diseases. 4. TLR4 is the major LPS receptor in mammalian cells. TLR4 triggers activation of F-κB (leading to production of TF-α, for example). Other TLRs recognize additional microbial products. OD-like receptors (LRs) are intracellular sensors of bacterial products that activate the inflammasome, triggering caspase-dependent maturation of IL Dendritic cells undergo a maturation program: immature Ds, which traffic to the periphery, capture antigen, and mature Ds, which traffic to the lymph node, present antigen. Innate immune stimuli trigger D maturation, which upregulates co-stimulatory molecules and facilitates antigen presentation. Thus, the innate immune response ushers in the acquired immune response. 6. K cells, a component of innate immunity, especially to viruses, represent an early source of IF-γ and serve to stimulate macrophages and Ds in inflammatory sites. Additional components of the antiviral response include intracellular dsra sensors (RIG-like proteins) that activate the IRF pathway to signal antiviral gene expression. 7
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