HUMORAL IMMUNE RE- SPONSES: ACTIVATION OF B CELLS AND ANTIBODIES JASON CYSTER SECTION 13

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1 SECTION 13 HUMORAL IMMUNE RE- SPONSES: ACTIVATION OF B CELLS AND ANTIBODIES CONTACT INFORMATION Jason Cyster, PhD ( ) READING Basic Immunology: Functions and Disorders of the Immune System. Abbas, Abul K., and Andrew H. Lichtman. -- Chapter 8; pp OBJECTIVES Describe the key changes that occur in the B cell upon binding antigen via its B cell receptor (BCR) Understand the 2 major classes of antigen, noting how they differ in T helper cell requirement and in the type of antibody response they induce Explain what a B cell must do in order to receive T cell help Describe the major components of T cell help that promote B cell responses and appreciate how deficiencies in this process can cause immunodeficiency Explain how a conjugate vaccine works in promoting a T-dependent antibody response, and explain why T- dependent responses are better Describe how you can make an antibody response Explain how isotype switching occurs, noting what part of the antibody is changed and what remains the same (the purpose of this change will be described in the next lecture on humoral immunity) Describe the properties of plasma cells and how the form of antibody they make differs from that of B cells Describe in brief the process of antibody affinity maturation that occurs inside germinal centers, recognizing the 2 key processes involved and the cellular outputs Understand the main Ig isotypes made during memory responses and why these responses are faster and of greater magnitude 71

2 KEY WORDS: ANTIBODY B CELL ACTIVATION T CELL HELP CONJUGATE-VACCINE HAPTEN ISOTYPE SWITCHING PLASMA CELL AFFINITY MATURATION MEMORY RESPONSE MAIN IDEAS: Humoral immunity is mediated by antibodies, whose functions in host defense are to neutralize and eliminate extracellular microbes and microbial toxins. Antibodies are produced following activation of B cells and differentiation to plasma cells. B cells become activated upon binding antigen and they present antigen-derived peptides to CD4 helper T cells that help trigger their clonal expansion and differentiation. Plasma cells secrete antibody specific for the same antigen encountered by the parent B cell. During differentiation B cells often isotype switch to make antibodies with constant (effector or Fc) regions that are most appropriate for the given response. Antibody affinity maturation occurs in germinal centers and involves a process of mutation and selection that generates B cells producing antibodies with improved affinity for the antigen. THE B CELL RECEPTOR (BCR) AND B CELL ACTI- VATION: - membrane Ig molecules are noncovalently attached to transmembrane signaling molecules, Igα and Igβ, to form the BCR complex - when BCRs engage antigen, tyrosines in Igα and Igβ are phosphorylated and become docking sites for adapter proteins that then recruit a number of signaling molecules and transmit downstream events TYPES OF HUMORAL IMMUNE RESPONSES: - T-independent responses are induced by strongly BCR cross-linking (polyvalent) antigens, inducing 72

3 rapid responses that are not accompanied by induction of germinal centers. T-independent responses can provide early protection but they do not show affinity maturation or immunological memory. - T-dependent responses can be induced by antigens that only weakly cross-link the BCR and are unable to induce B cell proliferation and differentiation on their own. These responses require T cell help and the antigen must contain a protein component. Because of the cell-cell interaction requirement, these responses are slower to get underway but they are more robust because they can undergo affinity maturation and memory B cell formation. B lymphocytes express receptors for a protein of the complement system (C3b) that provide costimulatory signals for the activation of the cells. They also express TLRs and pathogen-derived molecules such as LPS can synergize with BCR signaling to promote B cell responses. DIFFERENTIATION OF HELPER T CELLS (TH) Helper (CD4 + ) T cells that have been activated to develop into effector cells interact with antigenstimulated B cells at the edges of follicles in spleen and lymph nodes. These helper CD4 T cells are referred to as follicular helper T cells or Tfh. PRESENTATION OF ANTIGEN BY B CELLS TO TH CELLS B lymphocytes that bind protein antigens by their specific antigen receptors (BCRs) endocytose these antigens, process them in endosomal vesicles, and display class II MHC-associated peptides for recognition by CD4 + helper T cells. MECHANISMS OF TH CELL MEDIATED ACTIVA- TION OF B LYMPHOCYTES Helper T lymphocytes that recognize antigen presented by B cells activate the B cells by expressing CD40 ligand (CD40L) and by secreting cytokines (e.g. IL-4, IL-21). CONJUGATE VACCINES AND THE CONCEPT OF LINKED HELP Many bacteria have polysaccharide capsules and antibodies to these polysaccharides are an important defense. However, infants make only poor T-independent antibody responses. Antibody responses to polysaccharides can be improved by conjugating the polysaccha- 73

4 rides to an immunogenic protein carrier, changing the response from T-independent to T-dependent. RESPONSES TO HAPTENS Small organic molecules do not provoke antibodies by themselves but can if attached to a protein carrier. Such molecules are termed haptens (from the Greek haptein, to fasten). Some drugs such as Penicillin can act as haptens and induce antibody-mediated allergic reactions. PLASMA CELL DEVELOPMENT After receiving appropriate BCR signals (and help from CD4 T cells), B cells differentiate into antibody secreting cells (Plasma cells). Many of these terminally differentiated cells migrate to the bone marrow or mucosal surfaces where they reside for many months secreting antibody. HEAVY CHAIN (ISOTYPE) CLASS SWITCHING Helper T cells stimulate the progeny of IgM + IgD expressing B lymphocytes to produce antibodies of different heavy chain classes (isotypes): IgG, IgA or IgE. Heavy chain class switching is initiated by CD40Lmediated signals, and switching to different classes is stimulated by different cytokines. This process is called switch recombination. Cytokines produced by helper T cells determine which heavy chain class is produced by influencing which heavy chain constant region gene participates in switch recombination. AFFINITY MATURATION AND THE GERMINAL CEN- TER RESPONSE Affinity maturation is the process by which the affinity of antibodies produced in response to a protein antigen increases with prolonged or repeated exposure to that antigen. Affinity maturation occurs in germinal centers (GCs), and is the result of (1) somatic hypermutation of Ig genes in dividing B cells followed by (2) the selection of high-affinity B cells by antigen. Within the germinal center antigen is displayed on the surface of follicular dendritic cells (FDCs). Helper T cells play a critical role in the selection process. The germinal center response gives rise to long-lived plasma cells and memory B cells. 74

5 SECONDARY (MEMORY) ANTIBODY RESPONSE Secondary or tertiary exposure to T-dependent antigens is followed by a response that is faster and of greater magnitude than the primary response, and dominated by IgG. This is due in part to the presence of memory B cells generated in germinal centers during the primary response. Compared to antigenspecific B cells present on primary exposure, these cells are present in elevated numbers and are often isotype switched and of higher affinity for the antigen. 75

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