CHAPTER 9 BIOLOGY OF THE T LYMPHOCYTE
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1 CHAPTER 9 BIOLOGY OF THE T LYMPHOCYTE Coico, R., Sunshine, G., (2009) Immunology : a short course, 6 th Ed., Wiley-Blackwell 1
2 CHAPTER 9 : Biology of The T Lymphocytes Introduction Nature of the Antigen-Specific T Cell Receptor T Cell Genetics T Cell Differentiation in the Thymus Thymic Selection Single Positive T Cells Differentiation of Other Sets of Cells in Thymus 2
3 CONTENTS 1. Introduction 2. Nature of the Antigen-Specific T cell Receptor 2-1 Molecules that Interact with Antigen 2-2 Coreceptor Molecules 2-3 The T Cell Receptor Complex 2-4 Other Important Molecules Expressed on the T Cell Surface 2-5 γδ T cells 3. T Cell Genetics 3-1. Genes Coding for T Cell Receptors 3-2. T cell Receptor Gene Rearrangement 3-3. Generation of T Cell Receptor Diversity 4. T Cell Differentiation in the Thymus 4-1. Thymocytes Interact with Thymic Nonlymphoide Cells 4-2. Development of αβ T cells and γδ T cells 4-3. Thymocytes Maturation & Selection 5. Thymic Selection 5-1. Positive Selection 5-2. Negatie Selection 5-3. Role of AIRE Gene Product in Negative Selection 6. Single Positive T cells 7. Differentation of Other Sets of Cells in Thymus 3
4 1. Introduction 1. Location of the antigen - B cell & Ab ; Outside of the cell - T cells ; Inside of the cell (TcR cannot access to the antigens directly) 2. T Cell Receptor (TcR) Diversity ~10 18 different possible structures - The organization of the genes that TcR & BcR are strikingly similar 3. Differentiation of T cells & T cell education 4. Kinds of T Cell Receptors : - αβ TcR ; MHC restriction, (Th, CTL) - γδ TcR ; no MHC restriction 4
5 2. Nature of the Antigen-Specific T Cell Receptor CDRs on TcR 5
6 Differences between TcR & BcR Valence & conformation Antigen Secretion of the Receptor affinity maturation (somatic hypermutation) BcR bivalent & flexible at hinge region linear and conformational Yes Yes TcR mono-valent & rigid conformation only linear peptide associated with MHC molecules No No 6
7 [2-1] Molecules That Interact with Antigen Triple complex of MHC:pep:TcR 7
8 8
9 [2-2] Coreceptor Molecules α3 β2 Fig The TcR coreceptors and their interaction with MHC molecules 9
10 10
11 11
12 Figure 1. The life cycle of HIV presents several targets for inhibition. S. Nawale et al (2010) Int.J. ChemTech Res. 2(2),
13 13
14 [2-3] The T Cell Receptor Complex ITAM ITAM : Immunoreceptor tyrosine-based activation motif 14
15 [2-4] Other Important Molecules Expressed on the T cell Surface 1. Coreceptor Molecules : does not bind antigen but enhances T cell activation - CD4(1 chain) or CD8(2 chains) : CD4 + T cell and CD8 + T cell CD4 : MHC class II β 2 (professional APC and thymic epithlial cells) CD8 : MHC class I α 3 (all nucleated cells) 2. Costimulatory Ligands (for second signals) T cell APC Function CD28 B7-1(CD80), B7-2(CD86) IL-2 production CD40L (CD154) CD40 (professional APCs) CTLA4 (CD152) B7 Enhance CD28-B7 interaction For B cells isotype switching Negative signal to activated T cells (Terminate T cell activation) 15
16 [2-4] Other Important Molecules Expressed on the T cell Surface 3. Adhesion Molecules T cell APC Function CD2 LFA-3 (CD58) Adhesion LFA-1 (CD11aCD18) VLA-4 (CD49dCD29) 4. Homing Molecules ICAM-1 (CD54) VCAM-1 Early, naïve T cell Function Addressins at HEV of CD62L (L-selectin), α4β7 CCR7 To enter LNs peripheral Node mucosal Node Chemokine receptor 27 ; phagocytosis, 31 ; chemokine attraction 39 ; B/T 16
17 Fig Important molecules expressed at T-cell surface. Hatched bars indicate expression induced on activated cell.
18 18
19 19
20 20
21 [2-5] γδ T cells γδ T Cell not restricted by MHC - T cell receptor의 alternative form; αβ TcR 대신 γδ TcR - Found at mucosal epithelial sites (skin, gut, lung etc) - 기능, 항원의종류도전혀다름 mycobacteria, phospholipids, phosphoantigens, heat-shock proteins [2-6] NKT cells αβ T Cell not restricted by MHC - Bear NK cell surface marker (NKT란명칭의유래 ) - Do not recognize peptides displayed by APC recognize lipid antigens on CD1 molecule. - secretes cytokines 21
22 3. T Cell Genetics cock hen chicken 가족사진촬영 Scary time to be a Chicken
23 [3-1] GENES CODING FOR T CELL RECEPTORS Ch 7 Ch 14 Ch 7 23
24 [3-2] T Cell Receptor Gene Rearrangement 24
25 Ig gene expression 과특이하게다른점 : 1. No isotype switching : no changes in functions 2. No somatic mutation : no affinity maturations 3. TcR β chain has V-D-J-C and also V-J-C rearrangememt 25
26 [3-3] Generation of T Cell Receptor Diversity 26
27 27
28 4. T Cell Differentiation in the Thymus 1. Migration and proliferation: Bone marrow (precursor) Thymus (cortex medulla) Periphery 2. Differentiation (in thymus; immature T cell) TcR:CD3 accessory molecules (include CD4, CD8) 3. Selection (in thymus; mature T cell) Thymic Education Positive selection : became germ-line encoded self-mhc restricted T cell Negative selection : eliminates auto-reactive clones; the rests became self-tolerant and foreign Ag specific mature T cell 28
29 T CELL MIGRATION & DIFFERENTIATION 흉선은사춘기이후퇴화된다 - DiGeorge syndrom : Normal Ig, 세포성면역부전 반복감염, 귀, 코, 입기형유발 deletion of chromosome 22q11 absence of parathyroid gland no or very few T cells congenital thymic aplasia ( 선천성흉선무형성증 ) - Nude mouse(thymus not developed & lack of hair), SCID mouse 29
30 [4-1] Thymocytes Interact with Thymic Nonlymphoid Cells 30
31 Thymic epithelial cells express both MHC class I and II molecules Also produce IL-7 31
32 [4-2] Development of αβ T cells and γδ T cells Key stages in development of and T cells in the thymus. -TCR gene expression - Coreceptors CD4 and CD8. -The single line ( ) at the surface represents CD
33 [4-3] Thymocytes Maturation & Selection 95% of the T cells dies in the thymus during T cell education! 33
34 34
35 5. Thymic Selection General features ⑴ TcR must be expressed before selection ⑵ Random TcR expression give total repertoires for MHC and any antigen ⑶ Positive selection (self MHC recognition survive) ⑷ Negative selection (removal of self antigen recognition) * clonal deletion ; elimination clonal anergy ; inactivation (cf. no energy) ⑸ Double positive T cell 이 Positive & negative Selection 을거친후에야 Single Positive T cell 이된다. 35
36 Development of the self MHC-restricted T cell repertoire (1) The role of MHC, CD4 and CD8 in positive selection 1 Occurs during T cell passage through thymus by contacting with nurse cells. (T cell 의 maturation 과정중, 자신의 MHC 분자가아니라흉선상피세포의 MHC 분자에의한결합이며, 따라서 Thymocytes whose TcR has no affinity for self MHC will die) 2 Positive selection may be a result of the activation of thymocytes after the binding of their TcRs with self MHC molecules. without this stimulation, thymocyte will die (programmed cell death) 3 CD4 molecules recognize MHC classⅠ CD8 molecules recognize MHC classⅡ 4 Peptides also play a role in positive selection (TAP-1, TAP-2, 2C-pep) [5-1] Positive Selection (2) Mechanisms of Positive Selection in the Thymus Right combination of TcR may generate some signal to rescue of a thymocytes from programmed cell death. 36
37 [5-2] Negative Selection (1) Development of self-tolerance - deletion or inactivation of self-reactive clones of T cells during their maturation in the thymus ; T cell tolerance to self. - Bone marrow derived cells within thymus, such as dendritic cells, and MΦ can present self antigens to and cause the deletion of self antigen reactive T cell clones. (2) Mechanism of Negative Selection in the thymus. - Clonal deletion in the thymus : apoptosis [cell shrinkage, DNA fragmentation into nucleosome and nuclear fragmentation] - This apoptotic cell death is turned on by TcR-generated signals (activation-induced cell death). * Certain Vβ gene segment(vβ17a) in a TcR β chain 에결합하는 superantigen (peptide + I-E) 이있어서이러한 mouse 의 cortex 에는 TcR with Vβ17a 가존재하나 medulla 및 periphral organ 에는 delete 되어나타나지않음. [CD4 와 Vβ17a+TcR 및 I-E classii MHC molecule 들에의해 Vβ17a 가 deletion 된다.] 37
38 [5-3] Negative Selection-Role of AIRE gene Product How do we develop self-tolerance to molecules normally found outside the thymus? Certain Vβ gene segment(vβ17a) in a TcR β chain 에결합하는 superantigen (peptide + I-E) 이있어서이러한 mouse 의 cortex 에는 TcR with Vβ17a 가존재하나 medulla 및 peripheral organ 에는 delete 되어나타나지않음. CD4 와 Vβ17a+TcR 및 I-E classii MHC molecule 들에의해 Vβ17a 가 deletion 된다. Insulin(pancrease), thyroglobulin(thyroid), MBP(CNS) are expressed by thymic medullary epithelial cells. Autoimmune Regulator (AIRE) gene product encodes a transcription factor regulator allowing peripheral tissue antigens to be expressed in the thymus. 38
39 39
40 These topics are unclear and the subject of intense research activities! 40
41 41
42 FIGURE 9.7. Positive and negative thymic selection: positive selection, showing interaction of TCR + CD4 + CD8 + double-positive cell with cortical epithelial cells; negative selection, showing double-positive cell interacting with thymic dendritic cell.
43 6. Single Positive T Cells 1) Single Positive T cells ; Negative selection in thymus down regulated the expression of CD4 or CD8 resulting in CD4 + or CD8 + T cells. Leave the Thymus via blood stream to secondary lymphoid organ utilizing adhesion molecules and chemokin receptors. Naïve T cells may live for a long time(up to years) in a resting state. Mature T cells in ther periphery that have not encountered antigen need signals (including IL-7) to keep them alive. 2) Characteristics of αβ T Cells Emerging from Thymus ; (a) Self-MHC restricted (b) Self-tolerant 3) By antigen activation, T cells become effector T cell or memory T cell 43
44 7. Differentiation of Other Sets of Cells in Thymus Other sets of cells developed in the thymus and found at outside of the thymus (1) NK Cell - double negative cell 단계에서분리되는 cell line 으로 do not express TcR - part of the innate immune defenses against virus-infected and tumor cells (2) NK like cell expressing TcR - respond to glycolipid not by MHC but by CD1d - have features of both the innate and adaptive defenses. (3) Treg Cells - a subset of CD4+ T ells (~10% of peripheral helper T cells) - characteristically express CD25 and transcription factor Foxp3. - recognize combination of self MHC and self peptide, but have survived negative selection in the thymus. - Inhibitory response to self and foreign antigens to maintain and regulate self tolerance. 44
45 END of CH9 45
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