PHM142 Lecture 3: Phagocytes + Granulocytes

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1 PHM142 Lecture 3: Phagocytes + Granulocytes 1

2 Constituents of Blood (erythrocytes) Cellular Elements (leukocytes) (~35-55%) (thrombocytes) 2

3 White Blood Cells: Division of Labour (60%) (1-4%) (2-9%) (20-40%) (0.5%) 3

4 Phagocytosis Ilya IIyich (Élie) Metchnikoff ( ): the father of natural immunity Published a key paper in 1883 describing phagocytic cells in frogs The traits of the phagocyte have been retained most completely in the mesoderm where a large number of amoeboid cells occur to ingest the body s own dead or weak as well as foreign particles such as senescent red blood cells Awarded the Nobel Prize in Physiology and Medicine 1908 The key to immunity: stimulate the phagocytes 4

5 Macrophages Big Eaters Precursors: Monocytes blood, bone marrow, and spleen (storage) non-proliferative chemokine and pathogen recognition receptors; inflammatory cytokine production differentiate into macrophages and dendritic cells Tissue-specific populations: Mosser & Edwards Nat Rev Immunol

6 Roles of Macrophages 1. Tissue Homeostasis Removal of cellular debris from: tissue remodelling apoptosis necrosis: release of endogenous danger signals (heatshock proteins, nuclear proteins, histones, DNA) -> bind pattern recognition receptors -> production of inflammatory mediators 6

7 Roles of Macrophages 2. Host Defence Tissue surveillance: amoeboid movement; PRRs (TLRs, CLRs, NLRs) recognize signals associated with invading pathogens; Opsonin receptors (C3b and FC domain receptors) Microbicidal capacity: Macrophage activation occurs in response to innate and adaptive immune response signals (IFN and TNF) > secrete pro-inflammatory cytokines, superoxide, and oxygen radicals Natural killer cells first; TH1 cells after Phagocytosis 7

8 Roles of Macrophages 3. Wound Healing Basophils and Mast cells: release IL-4 in response to injury/ infection (chitin) TH2 cells: release IL-4 and IL-13 IL-4 -> arginase -> arginine to ornithine -> polyamines,.collagen -> extracellular matrix 4. Immune Regulation Switch: Anti-inflammatory activity to dampen the immune response 8

9 Neutrophils Synthesized from myeloid precursor cells in the bone marrow billion/day; half-life approx 8 hours (longer during inflammation) Production stimulated by IL-17A (NK cells) IL-17A released in response to IL-23 (macrophages + dendritic cells) Have a three to four-lobed nucleus + a granular cytoplasm 9

10 Mechanisms of Neutrophil Killing Kolaczkowska & Kubes Nat Rev Immunol

11 Mechanisms of Neutrophil Killing 1. Phagocytosis NADPH oxidase-dependent mechanisms (ROS) Anti-bacterial proteins (eg. cathepsins, defensins, lactoferrin, and lysozyme) released into the phagosome from neutrophil granules 11

12 Mechanisms of Neutrophil Killing 1. Phagocytosis: NADPH oxidase-dependent generation of ROS NADPH Oxidase: 2O 2 + NADPH -> 2O NADPH + + H + Multi-unit transmembrane electron transport chain Flavocytochrome b558: a heterodimeric heme protein (gp91 and p22) 2O H + -> O2 + H 2 O 2 iii. Nitric Oxide Synthase: 12

13 Mechanisms of Neutrophil Killing 1. Phagocytosis: NADPH oxidase-dependent generation of ROS NADPH Oxidase: 2O 2 + NADPH -> 2O NADPH + + H + Phagosome ii.superoxide Dismutase: 2O H + -> O2 + H 2 O 2 iii. Nitric Oxide Synthase: Cyt b558 13

14 Mechanisms of Neutrophil Killing 2. Degranulation Azurophilic (primary) granules: contain proteinases, cathepsins, elastase, azurocidin, defensins, lysozyme myeloperoxidase: H 2 O 2 + Cl - + H + -> HOCl (antimicrobial) + H 2 O; green pus Specific (secondary) granules: contain lysozyme, lactoferrin, and cytochrome B 558 Gelatinase (tertiary) granules: contain lysozyme, cytochrome B 558, matrix metalloproteinase 14

15 Mechanisms of Neutrophil Killing 3. Neutrophil Extracellular Traps composed of a core DNA element to which histones, proteins, and enzymes released from neutrophil granules attach directly kill, immobilize, promote phagocytosis of pathogens Kolaczkowska & Kubes Nat Rev Immunol

16 Neutrophil Extravasation Kolaczkowska & Kubes Nat Rev Immunol

17 Neutrophil Extravasation Step 1: Tethering endothelial cell activation (histamine/cytokines/ppr) -> expression of P-selectin and E-selectin -> bind P-selectin glycoprotein 1 ligand on neutrophil Step 2: Rolling adhesive bond breakage at the rear is balanced with adhesive bond formation at the leading edge Step 3: Adhesion GPCR binding of chemokine -> conformational change in surface integrins -> increased affinity for endothelial cell ligand 17

18 Neutrophil Extravasation Step 4: Crawling directed by shear flow NOT chemotactic gradient Step 5: Transmigration paracellular transcellular Chemokine hierarchy 18

19 Disorders of Neutrophil Function 1. Neutrophil Quantity: Neutropenia mild (ANC cells/μl); moderate (ANC cells/μl); severe (ANC<500 cells/μl) lack of pus at infection sites; susceptible to mouth sores + invasive bacterial infections genetic basis of severe congenital neutropenias: A. autosomal dominant mutation in ELANE B. mutations in GFI1 transcription factor C. autosomal recessive mutation in HAX1 chemotherapy-induced neutropenia 19

20 Disorders of Neutrophil Function 2. Neutrophil Granules Chediak-Higashi Syndrome: defects in granule morphogenesis; delayed and incomplete degranulation; associated with neurologic diseases + increased risk of bacterial infection Neutrophil-Specific Granule Deficiency: autosomal recessive mutations in C/EBP increased propensity for infections in the skin and mucous membranes 20

21 Disorders of Neutrophil Function 3. Neutrophil Chemotaxis: Leukocyte Adhesion Deficiencies autosomal recessive mutations in integrin components early onset soft tissue and invasive bacterial infections; delayed umbilical cord separation; peridontal disease; intellectual disabilities 4. Neutrophil Killing: Chronic Granulomatous Disease 1: 200,000 people; most present <5 y.o. recurrent, severe fungal + bacterial infections - skin, lungs, liver, lymph nodes genetic basis of CGD: A. X-linked mutation in gp91 B. Autosomal recessive mutations in p22 C. Autosomal recessive mutations in p47/p67 21

22 Eosinophils First described in 1879 by Paul Ehrlich Formed in the bone marrow, reside in the thymus and GI tract Exhibit antimicrobial activity towards bacterial, viral, and protozoan pathogens Contain large granules -> store and secrete cationic eosinophil-derived graunle proteins Form extracellular DNA traps -> meshwork of mitochondrial DNA fibers + EDGPs that trap and kill bacteria Rosenberg et al Nat Rev Immunol

23 Eosinophil-Derived Granule Proteins alters smooth muscle contaction generates ROS causing oxidative stress and cell death 23

24 Effects of Eosinophils 1. Eosinophil-Derived Granule Proteins 2. Cytokine Production (IL-2, -4, -6, -10, -12, -13, -16) 3. Lipid mediators: leukotrienes, platelet activating factor secreted by regulated exocytosis and degranulation ( piecemeal degranulation) 24

25 Basophils + Mast Cells produced in the bone marrow - basophils enter peripheral blood as mature cells; mast cells enter as immature cells granules containing histamine, cytokines, inflammatory chemokines -> released by IgE binding lifespan: ~60 hours for basophils; weeks-months for mast cells 25

26 Allergy Allergy: a hypersensitivity reaction associated with the production of allergen-specific IgE and with the expansion of allergen-specific T-cells that react against what are typically harmless, non-infectious environmental substance Allergen: non-infectious environmental substance that induces IgE production so later re-exposure induces an allergic reaction Anaphylaxis: a systemic allergic reaction 26

27 IgE Antibodies + Receptors IgE Structure: two heavy chains (extra domain vs IgG) + two light chains High Affinity Fc Receptor for IgE (FcεRI) αβγ 2 tetramer; expressed on mast cells + basophils αγ 2 trimer; expressed on eosinophils, APCs, monocytes, platelets, + smooth muscle cells Low Affinity CD23 Receptor for IgE three lectin head domains + a triple α-helix coiled-coil; expressed on activated B cells + epithelial cells Gould & Sutton Nat Rev Immunol

28 Sensitization allergen-mhc II complex presented by dendritic cells -> activate helper T-cells IL-4 (mast cells, basophils, eosinophils) -> Type 2 Helper T-cell characteristics IL-4 and IL-13 (TH2 cells, mast cells, and basophils) -> IgE production from plasma cells IgE -> systemic distribution via blood -> binds high affinity receptor -> sensitized mast cell Galli et al. Nature

29 Allergy in Three Phases 1. Early Phase Reactions (seconds - minutes) re-exposure to allergen -> crosslinking of IgE -> aggregation of FcεRI on mast cells + basophils -> degranulation result: vasodilation, increased vascular permeability, smooth muscle contraction, increased mucus secretion, nociceptor stimulation 2. Late Phase Reactions (hours) release of newly synthesized inflammatory mediators recruitment of T-cells, monocytes, eosinophils, neutrophils 3. Chronic Reactions Innate immune cells (eosinophils, basophils, neutrophils, monocyte/ macrophages) and adaptive immune cells (T cells, B cells) take up residence in the tissue structural changes/tissue remodelling 29

30 Chronic allergic inflammation + tissue remodeling in Asthma Galli et al. Nature

31 Treatment for Allergy 1. Allergen avoidance 2. Corticosteroids 3. Short/Long acting β2 adrenoceptor agonists 4. Anti-histamines 5. α adrenoceptor agonists 6. Anti-IgE Therapy (omalizumab) 31

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