WHY IS THIS IMPORTANT?

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1 CHAPTER 16 THE ADAPTIVE IMMUNE RESPONSE WHY IS THIS IMPORTANT? The adaptive immune system protects us from many infections The adaptive immune system has memory so we are not infected by the same pathogen a second time This is why vaccinations work OVERVIEW The Adaptive Immune Response COMPONENTS OF DEVELOPMENT OF COURSE OF THE IMMUNOLOGICAL OVERALL IMMUNE ADAPTIVE IMMUNITY LYMPHOCYTE POPULATIONS ADAPTIVE RESPONSE MEMORY RESPONSE ANTIGEN PRESENTATION CELLULAR (T CELL) RESPONSE HUMORAL (B CELL) RESPONSE 1

2 INTRODUCTION The adaptive immune response is the second level of immunity It involves lymphocytes It is a response to specific antigens It is extraordinarily specific It can adapt to any infection It has memory This confers potentially life-long immunity INTRODUCTION: Types of Response The innate response is a prerequisite for the adaptive immune response The adaptive immune response consists of two processes: Humoral production of antibodies Cellular killing of infected cells and regulation of other immune cells COMPONENTS OF ADAPTIVE IMMUNITY: Lymphatic System The adaptive response is associated with the lymphatic system It covers the entire body It involves lymphocytes and lymphoid structures 2

3 LYMPHATIC SYSTEM STRATEGIC LYMPHOID STRUCTURES Strategic lymphoid structures are found in places where pathogens enter MALT mucosa-associated lymphoid tissue Term given to lymphoid tissues associated with mucous membranes An important portal of entry Include several locations STRATEGIC LYMPHOID STRUCTURES GALT lymphoid tissue associated with the digestive tract Examples include the tonsils, adenoids, appendix, and Peyer s patches 3

4 STRATEGIC LYMPHOID STRUCTURES Peyer s patches are the most important part of GALT They contain M cells M cells are antigen collecting cells Under M cells are germinal centers Filled with B cells Surrounded by T cells STRATEGIC LYMPHOID STRUCTURES STRATEGIC LYMPHOID STRUCTURES BALT bronchus-associated lymphoid tissue Associated with the respiratory portal of entry Most available portal of entry NALT nasal-associated lymphoid tissue CALT conjunctival-associated lymphoid tissue 4

5 COMPONENTS ADAPTIVE IMMUNITY: Cells of Adaptive Immunity Cells involved in the adaptive response arise in the bone marrow They differentiate from stem cells Include the antigen presenting cells: dendritic cells, macrophages, and B cells Also include the lymphocytes: T cells and B cells DENDRITIC CELLS Dendritic cells are the important cells in the adaptive immune response. This is because: They have enormous surface areas They take up and process antigens They present antigens to T cells They are found in most tissues, especially associated with epithelial surfaces DENDRITIC CELLS David Scharf / Science Photo Library 5

6 DENDRITIC CELLS Immature dendritic cells migrate from bone marrow into blood and tissues They use phagocytosis and macropinocytosis to capture both self and nonself antigens Antigen recognition and processing causes a dendritic cell to mature into an antigen presenting cell (APC) DENDRITIC CELLS They eventually migrate to regional lymph nodes where they are responsible for: Presentation of antigens to lymphocytes Triggering of adaptive response DENDRITIC CELL Dr Olivier Schwartz, Institute Pasteur / Science Photo Library 6

7 ANTIGEN PRESENTING CELLS Several cell types can serve as antigen presenting cells and will display antigen on their surface Other immune cells, such as T cells, will be able to recognize and react to the antigen APCs also produce cytokines Include: Dendritic cells Macrophages B cells LYMPHOCYTES Common lymphoid precursor gives rise to: T cells B cells Natural Killer cells (part of innate immunity and covered in chapter 15) DIFFERENTIATION AND MATURATION OF LYMPHOCYTES Both B and T cells are formed in bone marrow B cells mature in the bone marrow There is always a supply of new B cells T cells mature in the thymus The thymus atrophies at puberty T cells are long lived The long-lived T cells can occasionally divide to maintain T cells to respond to infection 7

8 DIFFERENTIATION AND MATURATION OF LYMPHOCYTES When mature, both B and T cells circulate through the blood and tissues looking for antigens Mature lymphocytes that have not yet encountered their antigen are called naive lymphocytes They circulate as inactive cells DIFFERENTIATION AND MATURATION OF LYMPHOCYTES When B cells and T cells mature, they acquire specific antigen receptors The B cell receptor is an immunoglobulin molecule It has two antigen binding sites The T cell receptor is similar to Ig It has only one antigen binding site DIFFERENTIATION AND MATURATION OF LYMPHOCYTES 8

9 LYMPHOCYTES: B cells Activated mature B cells will differentiate into plasma cells Plasma cells make and secrete antibodies LYMPHOCYTES: T cells Activated mature T cells can differentiate into three types of T cells: Cytotoxic T cells kill infected host cells and pathogens Helper T cells activate B cells and other immune cells Regulatory T cells suppress the immune response once the antigen is reduced or gone LYMPHOTCYTES: Memory Cells Both B and T cells can differentiate into memory cells Memory cells are long lived lymphocytes responsible for immunological memory 9

10 CLONAL SELECTION OF LYMPHOCYTES Clonal selection is the process by which some lymphocytes are destroyed and others are maintained It takes place in the bone marrow for B cells and in the thymus for T cells CLONAL SELECTION OF LYMPHOCYTES Precursor cells produce lymphocytes Each lymphocyte is specific for a different antigen Specificity is generated through genetic rearrangement Clonal deletion is part of selection Lymphocytes that react with self are deleted (killed) CLONAL SELECTION OF LYMPHOCYTES Presentation of self antigens maintains tolerance Lymphocytes that ignore self antigens are selected and allowed to live Presentation of nonself antigens sends signals to T cells The adaptive response begins 10

11 CLONAL SELECTION OF LYMPHOCYTES Lymphocytes that remain after clonal deletion mature Each is specific for a nonself antigen If an antigen is encountered, the lymphocyte is activated It begins to divide and proliferate It forms a clone of cells specific for one antigen Lymphocytes that never encounter antigen eventually die CLONAL SELECTION OF LYMPHOCYTES Three important points about clonal selection: It generates a vast number of different antigen receptors Each receptor is specific for a different antigen All progeny of that lymphocyte will have the exact same receptor CLONAL SELECTION OF LYMPHOCYTES 11

12 SURVIVAL OF LYMPHOCYTE POPULATIONS: B Cells Millions of B cells are produced each day Their survival depends on signals from peripheral tissues These signals cause activation and proliferation or death B cells that never get stimulated undergo apoptosis Some activated B cells can differentiate into memory cells SURVIVAL OF LYMPHOCYTE POPULATIONS: T Cells T cells get survival signals from two places: Specialized epithelial cells in the thymus These occur during development Dendritic cells in peripheral lymphoid tissue SURVIVAL OF LYMPHOCYTE POPULATIONS: T Cells Once signaled, T cells in the lymph nodes stop migrating and become activated They become larger and multiply They multiply fourfold every 24 hours for 3-5 days Each one can differentiate into an armed effector T cell They can migrate into tissue or stay in the node 12

13 SURVIVAL OF LYMPHOCYTE POPULATIONS: T Cells Some activated effector T cells eventually die This is a safety precaution Some will become memory cells These respond very fast the second time an antigen is encountered LYMPHOID TISSUES Lymphocytes move through blood and lymph to the lymphoid tissues T and B cells are found in specific locations in a lymph node B cells are found in follicles These are in the outer cortex T cells surround follicles in paracortical areas LYMPHOID TISSUES 13

14 LYMPHOID TISSUES Lymphocytes circulate continuously through the lymphoid tissue They either find their specific antigen or die LYMPHOID TISSUES: B Cells B cells are constantly produced in the bone marrow Since most of them do not encounter their specific antigen, most of them die when they reach the peripheral tissue They are constantly replaced and therefore the number remains constant LYMPHOID TISSUES: T Cells T cells leave the thymus fully mature They are produced in smaller numbers than B cells T cells have a long life span They are thought to be self-renewing in peripheral tissue 14

15 ANTIGEN PRESENTATION: Types of Antigens There are two types of antigens: Self Nonself Self antigens are part of the immunological memory and cause tolerance Nonself antigens are any molecules that are not self ANTIGEN PRESENTATION: MHC For presentation to occur, the antigen must be bound to the major histocompatibility complex (MHC) The MHC is found on all nucleated cells ANTIGEN PRESENTATION: MHC There are two types of MHC: Class I is found on all nucleated body cells except immune cells and identify them as self Class II is found only on immune cells and serves to allow them communicate with each other 15

16 ANTIGEN PRESENTATION: Antigen Delivery by MHC Class I MHC molecules associate with cytoplasmic antigens in infected body cells They present the class I MHC-Antigen complex to cytotoxic T cells Cytotoxic T cells will proliferate, look for and kill host cells displaying this class I MHC- Antigen combination ANTIGEN PRESENTATION: Antigen Delivery by MHC Class II MHC is found exclusively on immune cells APCs combine collected antigen with class II MHC and present them to helper T cells The Helper T cells are activated and go on to activate other immune cells ANTIGEN PRESENTATION: Antigen Delivery by MHC 16

17 T CELL RESPONSE TO SUPERANTIGENS T cells can respond to superantigens These are distinct classes of antigens produced by many pathogens They provoke a response beneficial to the pathogen T CELL RESPONSE TO SUPERANTIGENS Superantigens bind to the outside of MHC molecules that are already bound to another antigen They cause massive overproduction of cytokines by helper T cells They cause inflammatory illness and toxic shock THE CELLULAR (T CELL) RESPONSE The cellular immune response is generated by T cells Cytotoxic T cells Helper T cells Regulatory T cells T cells that have not seen antigen are considered to be naive After encountering antigen, they become activated effector T cells 17

18 PRODUCTION OF ACTIVATED EFFECTOR T CELLS The activation of naive T cells can be done by: Dendritic cells Macrophages B cells ACTIVATION BY DENDRITIC CELLS Dendritic cells secrete chemokines Chemokines attract naive T cells Dendritic cells present viral, fungal, and bacterial antigens Dendritic cells produce co-stimulatory molecules to activate naive T cells They also present antigens that cause transplantation rejection and allergic reactions ACTIVATION BY MACROPHAGES Resting macrophages have few or no class II MHC molecules After ingesting a microorganism, they produce more They then degrade the microorganism and present antigen-class II MHC complex to T cells This causes the release of co-stimulatory molecules These allow the activation of T cells to continue 18

19 ACTIVATION OF T CELLS BY B CELLS Surface immunoglobulin on B cells is an antigen receptor It binds free antigen and is internalized by endocytosis Antigen is then bound to MHC class II molecules The complex is moved back to the surface of the B cell It is presented to T cells This causes the release of co-stimulatory molecules ACTIVATION OF T CELLS FUNCTIONS OF ACTIVATED EFFECTOR T CELLS There are three classes of activated effector T cells: Cytotoxic (CD8) T cells Helper (CD4) T cells Regulatory T cells Most of these move into the blood when they are activated 19

20 FUNCTIONS OF ACTIVATED EFFECTOR T CELLS Cytotoxic T cells release cytotoxins which kill infected host cells or pathogens Helper T cells release cytokines that activate other immune cells Regulatory T cell turn off the immune response when the antigen is gone CYTOTOXIC T CELLS: Antigen Encounter Some antigens are degraded in the cytoplasm of infected cells They are carried to the cell surface by class I MHC molecules They are then presented to cytotoxic (CD8) T cells Cytotoxic T cells proliferate and look for any cells also expressing that antigen They will kill those cells CYTOTOXIC T CELLS All viruses and some bacteria multiply in the cytoplasm of infected cells Once inside a cell, pathogens are no longer susceptible to antibodies 20

21 CYTOTOXIC T CELLS Cytotoxic T cells kill infected cells by inducing apoptosis in the infected cell They kill through cytotoxic granule release Perforins punch holes in the cell membrane Granulysin and granzyme enter the holes Granulysin induces apoptosis in the target cell Granzyme is a protease that kills cells and viruses CYTOTOXIC T CELLS CYTOTOXIC T CELLS Cytotoxic T cells are selective and repetitive killers of target cells They cells also release cytokines (interferon and TNF) These are part of the innate immune response 21

22 HELPER T CELLS: Antigen Encounter Antigens from pathogens are taken to the surface of the APC They are coupled with MHC class II molecules They are then presented to naive helper (CD 4) T cells Naive Helper T cells then proliferate and become immature effector cells Upon further stimulation, these immature effector T cells differentiate into several types of helper T cells TYPES OF HELPER T CELLS In the next slides we will talk about four types of helper T cells Each of these types has a distinct function in adaptive immunity The four type of helper T cells include: T H 1 cells T H 2 cells T H 17 cells T FH cells TYPES OF HELPER T CELLS: T H 1 cells Some bacteria proliferate in macrophages They can be eliminated if the macrophage becomes activated T H 1 cells provide the activation signal Activation increases the number of class II molecules This enhances antigen presentation and increases the adaptive response 22

23 TYPES OF HELPER T CELLS: T H 2 cells T H 2 cells promote antiparasitic activity Activate eosinophils and mast cells at tissue surfaces Activate plasma cells to produce antibody with antiparasitic action TYPES OF HELPER T CELLS: T H 17 cells and T FH cells T H 17 cells stimulate neutrophils and protect against extracellular bacteria and fungi T FH cells provide signals required to activate B cells that then produce antibody TYPES OF HELPER T CELLS 23

24 REGULATORY T CELLS Regulatory T cells produce cytokines that inhibit dendritic cells and T cell proliferation Suppress the immune response after the antigen is gone THE HUMORAL (B CELL) RESPONSE The humoral response is carried out by B lymphocytes It involves the production of the antibody B cells can be directly activated by antigen but in most cases, activation of B cells requires help from T cells THE HUMORAL RESPONSE: Activation After activation: Some B cells proliferate and differentiate into plasma cells Plasma cells produce massive amounts of antibody Some B cells become memory cells 24

25 THE HUMORAL RESPONSE: Antibodies Antibodies are also known as immunoglobulins They are found in the blood and in extracellular spaces They contribute to the adaptive response in three ways: Neutralization Neutralizes toxins and viruses Prevents bacterial attachment (staying in) Opsonization Facilitates uptake of pathogens by phagocytic cells Complement Activates the classical pathway THE HUMORAL RESPONSE: Antibodies THE IMMUNOGLOBULIN MOLECULE Antigen receptors on B cells are called immunoglobulins (Ig) All Ig molecules have a Y shape They are composed of four polypeptide chains Two light chains Two heavy chains The four amino terminal ends make up the antigenbinding site Remainders of the heavy chains make up the constant region 25

26 THE IMMUNOGLOBULIN MOLECULE THE IMMUNOGLOBULIN MOLECULE: Binding Antibodies are generally made against epitopes Epitopes are small surface regions of antigens Antibody binding involves hydrophobic and electrostatic forces Hydrogen bonds are also involved IMMUNOGLOBULIN ISOTYPES There are five different types of constant regions known as isotypes: IgG IgM IgA IgE IgD 26

27 IMMUNOGLOBULIN ISOTYPES The constant region of any immunoglobulin has three main functions: Recognition by specialized receptors on phagocytic cells Forming antigen-antibody complexes that initiate classical complement pathway Delivering antibody to tissues and secretions IMMUNOGLOBULIN ISOTYPES DISTRIBUTION OF IMMUNOGLOBULINS Most antibodies are distributed by diffusion Some are distributed by specialized transport mechanisms These are used to bring the antibody to the epithelial surfaces including: Digestive tract Lungs 27

28 IMMUNOGLOBULIN ISOTYPES: IgM IgM is the first antibody to be produced IgM can be in a pentamer structure This has ten binding sites and great binding strength It is usually found in blood It is an excellent activator of the complement system It is the primary response to bloodborne pathogens It is also found in pleural spaces This protects against environmental pathogens IMMUNOGLOBULIN ISOTYPES: IgG IgG antibodies are monomers They can diffuse easily in the fluids of the body IgG can be found in the blood and the extracellular fluid of the tissues IMMUNOGLOBULIN ISOTYPES: IgG IgG is very effective for opsonization, complement activation, blocking viral infection, neutralizing toxins, and preventing attachment of pathogens Newborn babies are born with their mother s IgG in their blood since it can easily cross the placenta This provides for early protection against infection 28

29 IMMUNOGLOBULIN ISOTYPES: IgA IgA is a dimer It is the principle antibody in secretions It is found in the respiratory and digestive tracts It is not very effective in activating the complement system IMMUNOGLOBULIN ISOTYPES: IgA IgA serves to block viral and bacterial attachment at mucosal surfaces Immunoglobulins can move across epithelial barriers through a process called transcytosis It occurs most frequently with IgA It is held together by a J chain and a secretory piece Allows for movement through epithelial cells IMMUNOGLOBULIN ISOTYPES: IgA and Transcytosis 29

30 IMMUNOGLOBULIN ISOTYPES: IgA IgA is found in colostrum It is transferred from mother to child It protects against newly encountered bacteria IMMUNOGLOBULIN ISOTYPES: IgE IgE is found in low levels in the blood and extracellular fluids It is found bound tightly to mast cells and basophils just below the skin and mucosa After antigen binding, powerful chemical mediators including histamine are released They cause coughing, sneezing and vomiting ACTIVATION OF BASOPHILS AND MAST CELLS BY IgE Photo courtesy of Ann Dvorak 30

31 IMMUNOGLOBULIN ISOTYPES: IgD IgD is found in very small amounts in the blood It is found on the surface of B cells On B cells it is involved as an early antigen receptor It has no know function in serum TIMING OF IMMUNOGLOBULIN RELEASE: Primary Response Naive B cells express both IgM and IgD on their surface After activation, IgD disappears IgM is the first isotype of antibody produced in large amounts This is the primary response TIMING OF IMMUNOGLOBULIN RELEASE: Primary Response Much later in the primary response, IgG is produced If the antigen producing the response is associated with a pathogen, the patient infected will show symptoms of the infection and feel ill IgG production will eventually rid the body of the infection, but comes too late to spare the patient the symptoms of the infection 31

32 TIMING OF IMMUNOGLOBULIN RELEASE: Secondary Response The secondary response occurs when the antigen is seen again Antibody production and isotype switching occurs quickly in the secondary response because memory cells are present for the antigen IgM gives way to IgG It is faster and more powerful than the primary response Helper T cells regulate the production and isotype of antibody TIMING OF IMMUNOGLOBULIN RELEASE: Secondary Response The secondary response is rapid enough that the pathogen is cleared from the body before causing the patient any symptoms The process of vaccination produces the primary response without causing real infection and symptoms If the patient encounters the pathogen in real life, the memory cells will already be present B CELL ACTIVATION BY T CELLS: Primary and Secondary Response 32

33 ANTIBODY ACTIVATION OF IMMUNE CELLS Antibody can activate many immune cells These include: Natural killer cells Basophils Mast cells Phagocytic cells B CELL ACTIVATION AND COOPERATION WITH T CELLS For the humoral response to work, each B cell must find a helper T cell This is accomplished by trapping Trapping occurs in the peripheral lymphoid tissue B CELL ACTIVATION AND COOPERATION WITH T CELLS Antigens are captured and processed by antigen-presenting cells (APCs) APCs migrate to lymph nodes and lodge in the paracortical areas Naive T cells move through the node If a T cell is presented with its antigen it is activated and remains trapped in the lymph node 33

34 B CELL ACTIVATION AND COOPERATION WITH T CELLS B cells also pass through the paracortical areas B cells are trapped if there is an activated T cell present T and B cells cooperate in the paracortical area B cells then move to follicular areas and differentiate into plasma cells B CELL ACTIVATION AND COOPERATION WITH T CELLS Plasma cells have a variety of life spans Some live for only a few days Others live for significantly longer periods They receive survival signals from bone marrow stromal cells COURSE OF THE ADAPTIVE RESPONSE 34

35 IMMUNOLOGICAL MEMORY Immunological memory is one of the most important properties of the adaptive immune response It can be seen in both T cells and B cells and is produced after infection or vaccination Memory is due to a persistent population of memory cells Most are at rest but a small percentage is dividing at all times Interleukins help to maintain the memory T cell population IMMUNOLOGICAL MEMORY After activation, the number of T cells increases As the infection subsides, the number decreases to a persistent level This level is times higher than the initial number before activation This increase is due to the increased number of memory cells NATURAL AND ARTIFICIAL IMMUNITY Adaptive immunity can be naturally acquired or artificially acquired It can also be active or passive 35

36 NATURAL AND ARTIFICIAL IMMUNITY Naturally acquired immunity is a result of normal life Naturally acquired active immunity would involve infection by a pathogen, production of an immune response to that pathogen and subsequent immunity to that pathogen Naturally acquired passive immunity would be the transfer of antibodies (IgG and IgA) from mother to infant NATURAL AND ARTIFICIAL IMMUNITY Artificially acquired immunity is the result of a medical treatment Artificially acquired active immunity occurs when a patient receives a vaccine, produces an immune response and is then immune to that pathogen Artificially acquired passive immunity occurs when a patient that is exposed to or infected by a pathogen is given antibodies or other immune produces from another individual THE OVERALL IMMUNE RESPONSE Innate and adaptive immune responses work together as fully integrated systems to defeat infection The end result is the control and elimination of infection and protection from re-infection 36

37 THE OVERALL IMMUNE RESPONSE The innate response works primarily in the early stages of the infection The adaptive immune response begins a few days after first exposure to an antigen Once pathogens have established an infection, it is only the adaptive response that can get rid of them THE OVERALL IMMUNE RESPONSE THE OVERALL IMMUNE RESPONSE 37

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