Chapter 5: Antigen Recognition by T Lymphocytes

Contents

  • T-cell receptor diversity

    • 5-1 The T-cell receptor resembles a membrane-associated Fab fragment of immunoglobulin

    • 5-2 T-cell receptor diversity is generated by gene rearrangement

    • 5-3 Expression of the T-cell receptor on the T-cell surface requires association with additional proteins

    • 5-4 A distinctive population of T cells expresses a second class of T-cell receptor with γ and δ chains

  • Antigen processing and presentation

    • 5-5 T-cell receptors recognize peptide antigens bound to MHC molecules

    • 5-6 Two classes of MHC molecule present peptide antigens to two types of T cell

    • 5-7 MHC class I and class II molecules have similar structures

    • 5-8 MHC class I binds shorter and more precisely defined peptides than MHC class II

    • 5-9 MHC class I and class II bind peptides in different intracellular compartments

    • 5-10 Peptides produced in the cytosol are transported to the endoplasmic reticulum for binding to MHC class I

    • 5-11 MHC class I binds peptides in the context of a highly specific peptide-loading complex

    • 5-12 All cells express MHC class I, whereas MHC class II is mainly expressed by professional antigen-presenting cells

    • 5-13 Invariant chain prevents MHC class II from binding peptides in the endoplasmic reticulum

    • 5-14 Cross-presentation enables extracellular antigens to be presented by MHC class I

  • The major histocompatibility complex

    • 5-15 Human MHC diversity is the product of gene families and genetic polymorphisms

    • 5-16 HLA class I and class II genes occupy separate regions of the HLA complex

    • 5-17 Proteins involved in antigen processing and presentation are encoded by genes in the HLA class II region

    • 5-18 Some MHC class I and class II genes are highly polymorphic

    • 5-19 Selection by infectious disease is a likely major cause of HLA class I and class II diversity

    • 5-20 Human populations all maintain a diversity of HLA class I and class II alleles

Key Terms and Processes Covered

  • Types of TCRs and associated proteins

  • Generation of TCR diversity

  • Process of antigen processing, presentation and recognition (TCR, MHC I and II)

  • MHC = Major Histocompatibility Complex (also called HLA in humans)

  • Transplantation

T-cell Receptor Diversity

  • 5-1 The T-cell receptor resembles a membrane-associated Fab fragment of immunoglobulin

    • TCRs (T-cell Receptors) are similar to BCRs (B-cell Receptors):

    • They possess an antigen-binding site, a constant region and a variable region.

    • They have complementarity-determining regions (CDR regions).

    • They are membrane-bound and comprise two polypeptide chains: alpha and beta.

    • TCR undergoes a transition from germline to rearranged DNA during development.

    • All TCRs on a single T cell are identical, while different T cells express different TCRs.

  • 5-2 T-cell receptor diversity is generated by gene rearrangement

    • TCR is produced similarly to BCR prior to encountering an antigen.

    • TCR consists of two polypeptide chains:

    • The alpha chain (analogous to the light chain of a BCR) is formed by V and J gene segment joining.

    • The beta chain (analogous to the heavy chain of a BCR) is formed by V, D, and J gene segment joining.

    • Rearrangement occurs in the thymus, producing a fully functional polypeptide with a leader peptide region and correctly assembled variable (V), constant (C), and membrane regions.

  • 5-3 Expression of the T-cell receptor on the T-cell surface requires association with additional proteins

    • CD3 complex: Composed of ε, γ, δ, and ζ chains, fulfilling crucial roles such as:

    • Facilitating transport of TCR to the cell surface

    • Enabling signal transduction, allowing extracellular signals to be transmitted into the cell

  • 5-4 A distinctive population of T cells expresses a second class of T-cell receptor with γ and δ chains

    • TCRα (TCR alpha) and TCRβ (TCR beta) represents 95% of T cells while TCRγ (TCR gamma) and TCRδ (TCR delta) account for 5%.

    • TCRγ and TCRδ are predominant in epithelial tissues and have the ability to recognize antigens beyond those presented by MHC molecules.

Antigen Processing and Presentation

  • 5-5 T-cell receptors recognize peptide antigens bound to MHC molecules

    • Terms to Know:

    • Antigen Processing: The process in which antigens are broken down into peptide fragments that are displayed on the cell surface.

    • Antigen Presentation: The display of antigenic peptides on MHC molecules at the surface of antigen-presenting cells (APCs).

    • Professional APC: Dendritic cells, macrophages, and B cells that are particularly effective in presenting antigens.

  • 5-6 Two classes of MHC molecule present peptide antigens to two types of T cell

    • MHC Class I and Class II have structural similarities but functionally differ in their source of peptides and interaction with T cells.

  • 5-7 MHC class I and class II molecules have similar structures

    • Both MHC classes have similar overall structure, including a peptide-binding groove and a stable region composed of β2-microglobulin (specific to MHC class I).

  • 5-8 MHC class I binds shorter and more precisely defined peptides than MHC class II

    • MHC Class I typically binds peptides that are 8-10 amino acids in length, whereas MHC Class II can present peptides of 13-25 amino acids long.

    • Each MHC molecule binds a single peptide at a time but can potentially accommodate numerous different peptide types over time.

  • 5-9 MHC class I and class II bind peptides in different intracellular compartments

    • The vesicular system involves vesicles and organelles such as the Golgi apparatus and the endoplasmic reticulum, serving as distinct compartments for peptide loading of MHC Class I and II.

  • 5-10 Peptides produced in the cytosol are transported to the endoplasmic reticulum for binding to MHC class I

    • Both self and foreign proteins are translated in the cytosol. Mismatched proteins are typically directed towards the proteasome for degradation.

    • During an infection, the cellular machinery modifies proteasomes to catalyze the production of peptides optimized for MHC Class I loading, transforming from standard proteasome to immunoproteasome.

  • 5-11 MHC class I binds peptides in the context of a highly specific peptide-loading complex

    • Steps include IFN-gamma inducing changes to proteasomes, resulting in the formation of immunoproteasomes that produce MHC I-binding peptides, which are then transported to the ER by TAP (Transporter Associated with Antigen Processing). Chaperone proteins facilitate proper folding and loading of the peptide onto MHC I.

  • 5-12 All cells express MHC class I, whereas MHC class II is mainly expressed by professional antigen-presenting cells

    • Presence of MHC class I across all cell types enables them to present endogenous antigens to CD8 T cells.

    • MHC class II is predominantly found on specialized APCs, increasing their antigen-presenting capabilities.

  • 5-13 Invariant chain prevents MHC class II from binding peptides in the endoplasmic reticulum

    • The invariant chain blocks premature peptide binding to MHC class II within the ER, facilitating appropriate antigen presentation only after the MHC class II has reached endocytic vesicles.

  • 5-14 Cross-presentation enables extracellular antigens to be presented by MHC class I

    • Cross-presentation allows professional APCs to present exogenous antigens to CD8 T cells by loading these antigens onto MHC class I molecules, crucial for initiating cytotoxic responses.

The Major Histocompatibility Complex

  • 5-15 Human MHC diversity is the product of gene families and genetic polymorphisms

    • MHC diversity is characterized by the presence of multiple alleles resulting from gene family expansion and varying polymorphisms among populations.

  • 5-16 HLA class I and class II genes occupy separate regions of the HLA complex

    • HLA genes are organized on a specific region of chromosome 6. Grouping allows for coordinated expression of MHC genes.

  • 5-17 Proteins involved in antigen processing and presentation are encoded by genes in the HLA class II region

    • This region has additional genes that directly influence the antigen processing pathway, impacting the immune response triggered by antigen presentation.

  • 5-18 Some MHC class I and class II genes are highly polymorphic

    • High polymorphism in MHC class I and II genes results in a broad range of peptide binding capabilities, enhancing immune defense

  • 5-19 Selection by infectious disease is a likely major cause of HLA class I and class II diversity

    • Increased diversity likely offers a selective advantage for populations under various infectious pressures, resulting in a greater number of alleles maintained across populations.

  • 5-20 Human populations all maintain a diversity of HLA class I and class II alleles

    • The maintenance of a diverse set of HLA alleles is crucial for effective immune responses against pathogens, supporting survival and health.