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ADDITIONAL T LYMPHOCYTE INTEGRINSThe extracellular matrix components collagen, fibronectin, laminin, vitronectin, and others are deposited and assembled into basement membranes and into three-dimensional fibrillar networks forming roads, boundaries and signposts in tissues that are probably used to guide lymphocyte migration. T lymphocytes express a number of integrins besides LFA-1, most of which are knownto interact with different extracellular matrix components. Resting T lymphocytes express VLA-4(~4fll), and T lymphocyte subpopulations express high levels of VLA-5(~5//1) and VLA-6(~6j~l) (63) (Table 2).

Cells do not crawl on planar membraneswithout proteins. In contrast, B lymphoblastoid cells become anchored to planar membranescontaining LFA-1, and despite dramatic motility of the cell body, no locomotion is observed over 12 hr of observation. The ability of cell surface LFA-1and other integrins to undergo cycles of avidity regulation maythus be critical for cell migration over the surface of other cells or through the extracellular matrix. One context for LFA-l-dependent lymphocyte migration is during extravasation in which lymphocytescrawl over the surface of endothelial cells expressing ICAM-1(Figure 2).

However,quantitative study of spontaneousdeadhesion of T lymphocytesfrom antigen bearing cells has been incomplete, possibly because all the required ingredients for spontaneous de-adhesion have not been present in most in vitro experiments (see below). However, it is clear that CTLcan "recycle" and kill manytarget cells with intervening migration (109-112), and that clusters of helper T lymphocytes with antigen-presenting cells disperse upon decay of antigen (108). The strength of adhesion between CTLsand target cells during the adhesion/ recognition stage is so great that when adherent cells are pulled apart the cells are dramatically deformed before they separate (113).

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