Tolerance and memory

In part, the clonal selection hypothesis was necessitated by the need to account for two fundamental hallmarks of the immune response: the capacity to respond to nonself or foreign antigens while not responding to self antigens; and the long-term retention of the capacity to mount accelerated and vigorous responses to previously encountered antigens. As anticipated by the theory, numerous studies have demonstrated the inactivation of developing B and T cells by antigens present in their environment, thus validating negative clonal selection as the basis for self tolerance. Although most B cells in an unim-munized animal have the capacity only to generate antibody-forming cell clones and not memory B cells, Linton, Decker and Klinman have identified a sub-population of precursor cells that do give rise to memory B cells and that fulfill, in detail, Burnet's predictions: 1) 'that a first contact with antigen sets some sort of tooling-up process in train, while secondary contact is needed to evoke actual antibody liberation into the circulation'; and 2) that 'the combination of frequent minor mutation and a highly effective selective process would rapidly improve the accuracy of the complementary relationship to new antigenic determinations'. The process of somatic mutation and antigen selection of these mutations has been described by numerous investigators including Tonegawa, Weigert, Gefter, Rajewsky, Berek and Milstein.

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