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The immune system has a huge stake in keeping mature T lymphocytes in a resting state until needed. Inappropriate T cell activation has devastating consequences for the host, including a variety of autoimmune diseases (1, 2). It is possible that the T cell maintains its quiescent state through overwhelming negative regulation, but this is unlikely because the T cell has to be ready to react with great sensitivity to MHC-peptide complexes on the surface of the antigenpresenting cell (APC). As few as three agonist MHC-peptide complexes on the surface of the APC are sufficient to activate a thymocyte for negative selection, whereas 300 are necessary to activate a naive T cell, with an intermediate number required for an effector T cell (3). Because only a fraction of these complexes will be found in the interface with a given thymocyte or T cell, it appears that thymocytes can be triggered by one MHC-peptide complex and that naive T cells require only about a dozen. Both thymocytes and mature T cells must measure interactions of T cell receptors (TCRs) with self-peptides that may be more numerous but that have interactions so weak as to be beyond reliable biochemical

Membrane domains and the immunological synapse: keeping T cells resting and ready