References Alon, R., D.A. Hammer, and T.A. Springer. 1995. Lifetime of the P-selectin carbohydrate bond and its response to tensile force in hydrodynamic flow. Nature. 374:539–542. Alon, R., R.C. Fuhlbrigge, E.B. Finger, and T.A. Springer. 1996. Interactions through L-selectin between leukocytes and adherent leukocytes nucleate rolling adhesions on selectins and VCAM-1 in shear flow. J. Cell Biol. 135: 849–865. Alon, R., S. Chen, D.K. Puri, E.B. Finger, and T.A. Springer. 1997. The kinetics of L-selectin tethers and the mechanics of selectin-mediated rolling. J. Cell Biol. 138:1169–1180. Bahara, P., and G.B. Nash. 1998. Sparsely adherent platelets support capture and immobilization of flowing neutrophils. J. Lab. Clin. Med. 132:223–228. Bell, G.I. 1978. Models for the specific adhesion of cells to cells. Science. 200: 618–627. Bruehl, R.E., K.L. Moore, D.E. Lorant, N. Borregaard, G.A. Zimmerman, R.P. McEver, and D.F. Bainton. 1997. Leukocyte activation induces surface re- distribution of P-selectin glycoprotein ligand-1. J. Leuk. Biol. 61:489–499. Buttrum, S.M., R. Hatton, and G.B. Nash. 1993. Selectin-mediated rolling of neutrophils on immobilized platelets. Blood. 82:1165–1174. Chen, S.Q., and T.A. Springer. 1999. An automatic braking system that stabi- lizes leukocyte rolling by an increase in selectin bond number with shear. J. Cell Biol. 144:185–200. Diacovo, T.G., S.J. Roth, J.M. Buccola, D.F. Bainton, and T.A. Springer. 1996. Neutrophil rolling, arrest, and transmigration across activated, surface- adherent platelets via sequential action of P-selectin and the b2-integrin CD11b/CD18. Blood. 88:146–157. Evans, S., D. Berk, and A. Leung. 1991. Detachment of agglutinin-bonded red blood cells. I. Forces to rupture molecular-point attachments. Biophys. J. 59: 838–848. Fritz, J., A.G. Katopodis, F. Kolbinger, and D. Anselmetti. 1998. Force-medi- ated kinetics of single P-selectin/ligand complexes observed by atomic force microscopy. Proc. Natl. Acad. Sci. USA. 95:12283–12288. Goldman, A.J., R.G. Cox, and H. Brenner. 1967. Slow viscous motion of a sphere parallel to a plane wall: couette flow. Chem. Eng. Sci. 22:653–660. Hagberg, I.A., H.E. Roald, T. Lyberg. 1998. Adhesion of leukocytes to growing arterial thrombi. Thromb. Haemost. 80:852–858. Hamburger, S.A., and R.P. McEver. 1990. GMP-140 mediates adhesion of stim- ulated platelets to neutrophils. Blood. 75:550–554. Kostantopoulos, K., J.C. Grotta, C. Sills, K.K. Wu, and J.D. Hellums. 1995. Shear-induced platelet aggregation in normal subjects and stroke patients. Thromb. Haemost. 74:1329–1334. Lalor, P., and G.B. Nash. 1995. Adhesion of flowing leucocytes to immobilized platelets. Br. J. Haematol. 89:725–732. Lawrence, M.B., and T.A. Springer. 1991. Leukocytes roll on a selectin at phys- iologic flow rates: distinction from and prerequisite for adhesion through in- tegrins. Cell. 65:859–873. Lawrence, M.B., G.S. Kansas, E.J. Kunkel, and K. Ley. 1997. Threshold levels of fluid shear promote leukocyte adhesion through selectins (CD62L,P,E). J. Cell Biol. 136:711–727. Lim, Y.C., K. Snapp, G.S. Kansas, R. Camphausen, H. Ding, and F.W. Luscin- skas. 1998. Important contributions of P-selectin glycoprotein ligand-1-medi- ated secondary capture to human monocyte adhesion to P-selectin, E-selec- tin, and TNF-alpha-activated endothelium under flow in vitro. J. Immunol. 161:2501–2508. Mehta, P., R.D. Cummings, and R.P. McEver. 1998. Affinity and kinetic analy- sis of P-selectin binding to P-selectin glycoprotein ligand-1. J. Biol. Chem. 273:32506–32513. Mickelson, J.K., N.M. Lakkis, G. Villarreal-Levy, B.J. Hughes, and C.W. Smith. 1996. Leukocyte activation with platelet adhesion after coronary angio- plasty: a mechanism for recurrent disease? J. Am. Coll. Cardiol. 28:345–353. Moore, K.L., K.D. Patel, R.E. Bruehl, F. Li, D.A. Johnson, H.S. Lichenstein, R.D. Cummings, D.F. Bainton, and R.P. McEver. 1995. P-selectin glycopro- tein ligand-1 mediates rolling of human neutrophils on P-selectin. J. Cell Biol. 128:661–671. Neumann, F.J., N. Marx, M. Gawaz, K. Brand, I. Ott, C. Rokitta, C. Sticherling, C. Meinl, A. May, and A. Schömig. 1997. Induction of cytokine expression in leukocytes by binding of thrombin-stimulated platelets. Circulation. 95: 2387–2394. Rinder, C.S., J.L. Bonan, H.M. Rinder, J. Mathew, R. Hines, and B.R. Smith. 1992. Cardiopulmonary bypass induces leukocyte-platelet adhesion. Blood. 79:1201–1205. Shao, J.Y., and R.M. Hochmuth. 1996. Micropipette suction for measuring pi- conewton forces of adhesion and tether formation from neutrophil mem- branes. Biophys. J. 71:2892–2901. Shao, J.Y., and R.M. Hochmuth. 1999. Mechanical anchoring strength of L-selec- tin, b2 integrins, and CD45 to neutrophil cytoskeleton and membrane. Bio- phys. J. 77:587–596. Shao, J.Y., H.P. Ting-Beall, and R.M. Hochmuth. 1998. Static and dynamic lengths of neutrophil microvilli. Proc. Natl. Acad. Sci. USA. 95:6797–6802. Smith, M.J., E.L. Berg, and M.B. Lawrence. 1999. A direct comparison of selec- tin-mediated transient, adhesive events using high temporal resolution. Bio- phys. J. 77:3371–3383. Walcheck, B., K.L. Moore, R.P. McEver, and T.K. Kishimoto. 1996. Neu- trophil-neutrophil interactions under hydrodynamic shear stress involve L-selectin and PSGL-1. J. Clin. Invest. 98:1081–1087. Weber, C., and T.A. Springer. 1997. Neutrophil accumulation on activated, sur- face-adherent platelets in flow is mediated by interaction of Mac-1 with fi- brinogen bound to aIIbb3 and stimulated by platelet-activating factor. J. Clin. Invest. 100:2085–2093. Yeo, E.L., J.A. Sheppard, and J.A. Feuerstein. 1994. Role of P-selectin and leu- kocyte activation in polymorphonuclear cell adhesion to surface adherent activated platelets under physiologic shear conditions (an injury vessel wall model). Blood. 83:2498–2507. Schmidtke and Diamond Neutrophil Tethering under Flow 729