Immunological diversity Gilbert Chu January 2004

Immunological diversity Gilbert Chu January 2004 www.phwiki.com

Immunological diversity Gilbert Chu January 2004

Laurie, Kim, Midday Host has reference to this Academic Journal, PHwiki organized this Journal Immunological diversity Gilbert Chu January 2004 Discovery of antibody diversity 430 BC Thucydides On bubonic plague: ”It was with those who had recovered from disease that the sick in addition to the dying found most compassion. These knew what it was from experience, in addition to had now no fear themselves; as long as the same man was never attacked twice – never at least fatally.” 1796 Jenner Noted that cowpox was rarely followed by smallpox Showed that cowpox innoculum protected from smallpox Pasteur coined “vaccine” from vacca, cow in Latin 1901 L in addition to steiner Discovered antibodies against ABO blood antigens Made antibodies against many organic molecules: specificity in addition to diversity Discovered antibodies against the red blood cell antigen in paroxysmal cold hemoglobinuria: autoimmunity The antibody molecule

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Mouse immunoglobulin genes V1-V500 D1-D12 J1-J4 H chain locus (Chr 12) Cm Cd Cg3 Cg1 Cg2b Cg2a Ce Ca V1-V250 J1-J5 Ck Vl2 Jl2 Cl2 k chain locus (Chr 6) l chain locus (Chr 16) Cl3 Cl1 Vl1 Jl3 Jl1 Mechanisms as long as generating antibody diversity V(D)J recombination Somatic hypermutation Class switch recombination Mechanisms as long as generating antibody diversity V(D)J recombination Hozumi in addition to Tonegawa, PNAS 1976

Mechanisms as long as generating antibody diversity V(D)J recombination Mechanisms as long as generating antibody diversity V(D)J recombination V1-V500 D1-D12 J1-J4 constant regions Mechanisms as long as generating antibody diversity V(D)J recombination D to J joining

Mechanisms as long as generating antibody diversity V(D)J recombination D to J joining Mechanisms as long as generating antibody diversity V(D)J recombination V to DJ joining Mechanisms as long as generating antibody diversity V(D)J recombination Somatic hypermutation Somatic mutations

Mechanisms as long as generating antibody diversity V(D)J recombination Somatic hypermutation Class switch recombination Class switch Mechanisms as long as generating antibody diversity V(D)J recombination Somatic hypermutation Class switch recombination Class switch Mechanisms as long as generating antibody diversity V(D)J recombination Somatic hypermutation Class switch recombination V(D)J recombination in addition to class switch recombination involve double-str in addition to breaks

Recombination signal sequence (RSS) direct V(D)J recombination V(D)J recombination involves DNA cleavage in addition to end-joining Cleavage is initiated by RAG1/RAG2 (recombination activating genes) van Gent, Gellert et al. Cell 1995

DNA ends are modified by addition in addition to deletion N-nucleotide addition by terminal deoxynucleotidyl transferase (TdT) P-nucleotide addition by asymmetric opening of hairpin coding ends Nucleotide deletion Addition at DNA ends Deletion at DNA ends

DNA pathways in V(D)J recombination Evolution of V(D)J recombination RAG1 in addition to RAG2 contain no introns in addition to are tightly linked on the same chromosome RAG1 in addition to RAG2 are conserved back to the evolution of jawed fish Evolutionary hypothesis: a transposon with RAG1, RAG2, in addition to associated RSSs infected a precursor of jawed fish RAG1 in addition to RAG2 do not exist in jawless fish

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Hypothetical RAG transposon Transposon integration Agrawal, Eastman in addition to Schatz Nature 1998 Origin of the immunoglobulin genes

The scid mouse Mouse with severe combined immunodeficiency, lacking mature B in addition to T cells Defective in the joining of coding ends Normal in the joining of signal ends Hypersensitive to ionizing radiation The scid mouse suggested a link between V(D)J recombination in addition to the repair of DNA double-str in addition to breaks Mutant nonlymphoid cells can be tested as long as V(D)J recombination Mutagenesis of Chinese hamster epithelial cells generated several X-ray sensitive cell lines These cells were co-transfected with RAG1, RAG2, in addition to V(D)J recombination substrates The cells were then assayed as long as either coding joint as long as mation or signal joint as long as mation V(D)J recombination substrates Lieber, Gellert et al. Cell 1998

Model as long as class switch recombination How does AID initiate CSR at one locus in addition to SHM at another (The C-terminus of AID is required as long as CSR, but not SHM.) Summary Diversity is generated by multiple mechanisms V(D)J recombination Somatic hypermutation Class switch recombination Some components are lymphocyte-specific RAG1/RAG2, TdT, AID Other components are ubiquitous Double-str in addition to break repair, base excision repair

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