Poly(ethylene glycol) (PEG)-mediated fusion between pure lipid bilayers: a mechanism in common with viral fusion and secretory vesicle release? (Review)

References

• Alvarez de Toledo, G.Fernandez-Chacon, R. and Fernandez, J. M. 1993, Release of secretory products during transient vesicle fusion. Nature, 363, 554–558.
   Google Scholar
• Arnold, K., Herrmann, A., Gawrisch, K. and Pratsch, L. 1985, Mechanisms of PEG-induced fusion. Studia Biophysica, 110, 135–141.
   Google Scholar
• Arnold, K., Pratsch, L. and Gawrisch, K. 1983, Effect of poly(ethylene glycol) on phospholipid hydration and polarity of the external phase. Biochimica et Biophysica Acta, 728, 121–128.
   PubMedGoogle Scholar
• Arnold, K., Zschoernig, O., Barthel, D. and Herold, W. 1990, Exclusion of poly(ethylene glycol) from liposome surfaces. Biochimica et Biophysica Acta, 1022, 303–310.
   PubMed Web of Science ®Google Scholar
• Bailey, F. E. and Koleske, J. V. 1967. Configuration and hydro- dynamic properties of the polyethylene chain in solution in nonionic surfactants. In Nonionic Surfactants, M.J. Schick (ed), NY, USA: Marcel Dekker pp. 794–822.
   Google Scholar
• Bentz, J., and Ellens, H. 1988, Membrane fusion: kinetics and mechanisms. Colloids and Surfaces, 30, 65–112.
   Google Scholar
• Blumenthal, R., Sarkar, D. P., Durell, S., Howard, D. E. and Morris, S. J. 1996, Dilation of the influenza hemagglutinin fusion pore revealed by the kinetics of individual cell-cell fusion events. Journal of Cell Biology, 135, 63–71.
   PubMed Web of Science ®Google Scholar
• Boni, L. T., Stewart, T. P., Alderfer, J. L. and Hui, S. W. 1981, Lipid- polyethylene glycol interactions: I. Induction of fusion between liposomes. Journal of Membrane Biology, 62, 65–70.
   PubMedGoogle Scholar
• Breckenridge, L. J. and Almers, W. 1987, Final steps in exocytosis observed in a cell with giant secretory granules. Proceedings of the National Academy of Sciences of the United States of America, 84, 1945-1949.
   PubMed Web of Science ®Google Scholar
• Bullough, P. A., Hughson, F. M., Skehel, J. J. and Wiley, D. C. 1994, Structure of influenza haemagglutinin at the pH of membrane fusion [see comments]. Nature, 371, 37–43.
   PubMed Web of Science ®Google Scholar
• Burgess, S. W., Massenburg, D., Yates, J. and Lentz, B. R. 1991, Poly(ethylene glycol)-induced lipid mixing but not fusion between synthetic phosphatidylcholine large unilamellar vesicles. Biochem- istry, 30, 4193–4200.
   Google Scholar
• Burgess, S. W., McIntosh, T. J. and Lentz, B. R. 1992, Modulation of poly(ethylene glycol)-induced fusion by membrane hydration: importance of interbilayer separation. Biochemistry, 31, 2653-2661.
   Google Scholar
• Carr, C. M. and Kim, P. S. 1993, A spring-loaded mechanism for the conformational change of influenza hemagglutinin. Cell, 73, 823- 832.
   PubMed Web of Science ®Google Scholar
• Carr, C. M., Chaudhry, C. and Kim, P. S. 1997, Influenza hemagglutinin is spring-loaded by a metastable native conforma- tion. Proceedings of the National Academy of Sciences (USA), 94, 14306–14313.
   PubMed Web of Science ®Google Scholar
• Chandler, D. E., and Heuser, J. 1979, Membrane fusion during secretion: cortical granule exocytosis in sex urchin eggs as studied by quick-freezing and freeze-fracture. Journal of Cell Biology, 83, 91–108.
   PubMed Web of Science ®Google Scholar
• Chanturiya, A., Chernomordik, L. V. and Zimmerberg, J. 1997, Flickering fusion pores comparable with initial exocytotic pores occur in protein-free phospholipid bilayers. Proceedings of the National Academy of Sciences (USA), 94, 14423–14428.
   PubMed Web of Science ®Google Scholar
• Chen, J., Lee, K. H., Steinhauer, D. A., Stevens, D. J., Skehel, J. J. and Wiley, D. C. 1998, Structure of the hemagglutinin precursor cleavage site, a determinant of influenza pathogenicity and the origin of the labile conformation. Cell, 95, 409–417.
   PubMed Web of Science ®Google Scholar
• Chernomordik, L., Chanturiya, A., Green, J. and Zimmerberg, J. 1995a, The hemifusion intermediate and its conversion to complete fusion: regulation by membrane composition. Biophysi- cal Journal, 69, 922–929.
   PubMed Web of Science ®Google Scholar
• Chernomordik, L., Kozlov, M. M. and Zimmerberg, J. 1995b, Lipids in biological membrane fusion. Journal of Membrane Biology, 146, 1–14.
   PubMed Web of Science ®Google Scholar
• Chernomordik, L. V., Frolov, V. A., Leikina, E., Bronk, P. and Zimmerberg, J. 1998, The pathway of membrane fusion catalyzed by influenza hemagglutinin: restriction of lipids, hemifusion, and lipidic fusion pore formation. Journal of Cell Biology, 140, 1369-1382.
   PubMed Web of Science ®Google Scholar
• Chernomordik, L. V., Melikyan, G. B. and Chizmadzhev, Y. A. 1987, Biomembrane fusion: a new concept derived from model studies using two interacting planar lipid bilayers. Biochimica et Biophy- sica Acta, 906, 309–352.
   PubMed Web of Science ®Google Scholar
• Chernomordik, L. V., Vogel, S. S., Sokoloff, A., Onaran, H. O., Leikina, E. A. and Zimmerberg, J. 1993, Lysolipids reversibly inhibit Ca(2+)-, GTP- and pH-dependent fusion of biological membranes. FEBS Letters, 318, 71–76.
   PubMed Web of Science ®Google Scholar
• Chow, R. H., von Ruden, L. and Neher, E. 1992, Delay in vesicle fusion revealed by electrochemical monitoring of single secretory events in adrenal chromaffin cells. Nature, 356, 60–63.
   PubMed Web of Science ®Google Scholar
• Clague, M. J., Schoch, C., Zech, L. and Blumenthal, R. 1990, Gating kinetics of pH-activated membrane fusion of vesicular stomatitis virus with cells: stopped-flow measurements by dequenching of octadecylrhodamine fluorescence. Biochemistry, 29, 1303-1308.
   Google Scholar
• Cullis, P. R., Hope, M. J. and Tilcock, C. P. 1986, Lipid polymorphism and the roles of lipids in membranes. Chemical and Physical Lipids, 40, 127–144.
   PubMed Web of Science ®Google Scholar
• Cullis, P. R., Verkleij, A. J. and Ververgaert, P. H. 1978, Polymorphic phase behaviour of cardiolipin as detected by 31P NMR and freeze-fracture techniques. Effects of calcium, dibucaine and chlorpromazine. Biochimica et Biophysica Acta, 513, 11–20.
   Google Scholar
• Danieli, T., Pelletier, S. L., Henis, Y. I. and White, J. M. 1996, Membrane fusion mediated by the influenza virus hemagglutinin requires the concerted action of at least three hemagglutinin trimers. Journal of Cell Biology, 133, 559–569.
   PubMed Web of Science ®Google Scholar
• Davidson, R. L. and Gerald, P. S. 1977, Induction of mammalian somatic cell hybridization by polyethylene glycol. Methods in Cell Biology, 15, 325–338.
   PubMedGoogle Scholar
• De Kruijff, B. and Van Zoelen, E. J. 1978, Effect of the phase transition on the transbilayer movement of dimyristoyl phospha- tidylcholine in unilamellar vesicles. Biochimica et Biophysica Acta, 511, 105–115.
   Google Scholar
• Doms, R. W. and Helenius, A. 1986, Quaternary structure of influenza virus hemagglutinin after acid treatment. Journal of Virology, 60, 833–839.
   Google Scholar
• Düzgüneş, N., Wilschut, J. and Papahadjopoulos, D. 1985, Control of membrane fusion by divalent cations, phospholipid headgroups and proteins. In Physical Methods on Biological Membranes and Their Model Systems, F. Conti, W. E. Blumberg, J. de Gier and F. Pocchiari (eds), (NY, USA: Plenum Press Publishing Corp.), pp. 193–218
   Google Scholar
• Ellens, H., Bentz, J., Mason, D., Zhang, F. and White, J. M. 1990, Fusion of influenza hemagglutinin-expressing fibroblasts with glycophorin-bearing liposomes: role of hemagglutinin surface density. Biochemistry, 29, 9697–9707.
   PubMed Web of Science ®Google Scholar
• Ellens, H., Bentz, J. and Szoka, F. C. 1986, Destabilization of phosphatidylethanolamine liposomes at the hexagonal phase transition temperature. Biochemistry, 25, 285–294.
   PubMed Web of Science ®Google Scholar
• Ellens, H., Siegel, D. P., Alford, D., Yeagle, P. L., Boni, L., Lis, L. J., Quinn, P. J. and Bentz, J. 1989, Membrane fusion and inverted phases. Biochemistry, 28, 3692–3703.
   PubMed Web of Science ®Google Scholar
• Evans, E. and Metcalfe, M. 1984, Free energy potential for aggregation of mixed phosphatidylcholine/phosphatidylserine lipid vesicles in glucose polymer (dextran) solutions. Biophysical Journal, 45, 715–720.
   Google Scholar
• Evans, E. and Needham, D. 1988. Intrinsic colloidal attraction/ repulsion between lipid bilayers and strong attraction induced by non-adsorbing polymers. In Molecular Mechanisms of Mem- brane Fusion, S. Ohki, D. Doyle, T. D. Flanagan, S. H. Hui and E. Mayhew (eds), (NY, USA: Plenum Press Publishing Corp.), pp. 83–99.
   Google Scholar
• Faist, S. and Seibicke, S. 1987, A hypothesis on the mechanism of PEG-mediated membrane fusion. Cell Biology Internatlonal Reports, 11, 563.
   Google Scholar
• Fernandez, J. M., Neher, E. and Gomperts, B. D. 1984, Capacitance measurements reveal stepwise fusion events in degranulating mast cells. Nature, 312, 453–455.
   PubMed Web of Science ®Google Scholar
• Godley, L, Pfeifer, J. Steinhauer, D., Ely, B., Shaw, G., Kaufmann, R., Suchanek, E., Pabo, C., Skehel, J. J. and Wiley, D. C., 1992, Introduction of intersubunit disulfide bonds in the membrane-distal region of the influenza hemagglutinin abolishes membrane fusion activity. Cell, 68, 635–645.
   Google Scholar
• Gray, C. and Tamm, L. K. 1997, Structural studies on membrane- embedded influenza hemagglutinin and its fragments. Protein Science, 6, 1993-2006.
   Google Scholar
• Gruner, S. M. 1985, Intrinsic curvature hypothesis for biomembrane lipid composition: a role for nonbilayer lipids. Proceedings of the National Academy of Sciences (USA), 82, 3665–3669.
   PubMed Web of Science ®Google Scholar
• Harter, C., Bachi, T., Semenza, G. and Brunner, J. 1988, Hydrophobic photolabeling identifies BHA2 as the subunit mediat- ing the interaction of bromelain-solubilized influenza virus hemag- glutinin with liposomes at low pH. Biochemistry, 27, 1856-1864.
   PubMed Web of Science ®Google Scholar
• Hernandez, L. D., Hoffman, L. R., Wolfsberg, T. G. and White, J. M. 1996, Virus-cell and cell-cell fusion. Annals of Reviews in Cell Development Biology, 12, 627–661.
   PubMed Web of Science ®Google Scholar
• Herrmann, A., Pratsch, L., Arnold, K. and Lassmann, G. 1983, Effect of poly(ethylene glycol) on the polarity of aqueous solutions and on the structure of vesicle membranes. Biochimica et Biophysica Acta, 733, 87–94.
   Google Scholar
• Honda, K., Maeda, Y., Sasakawa, S., Ohno, H. and Tsuchida, E. 1981, Activities of cell fusion and lysis of the hybrid type of chemical fusogens. (I). Structure and function of the promotor of cell fusion. Biochemical & Biophysical Research Communications, 100, 442–448.
   PubMed Web of Science ®Google Scholar
• Huang, S. K. and Hui, S. W. 1986, Chemical co-treatments and intramembrane particle patching in the poly(ethylene glycol)- induced fusion of turkey and human erythrocytes. Biochimica et Biophysica Acta, 860, 539–548.
   Google Scholar
• Hui, S. W., Isac, T., Boni, L. T. and Sen, A. 1985, Action of polyethylene glycol on the fusion of human erythrocyte mem- branes. Journal of Membrane Biology, 84, 137–146.
   PubMedGoogle Scholar
• Jones, J. D. and Thompson, T. E. 1990, Mechanism of spontaneous, concentration-dependent phospholipid transfer between bilayers [published errata appear in Biochemistry, 29, 4746 and 30, 10818]. Biochemistry, 29, 1593–1600.
   Google Scholar
• Kanaseki, T., Kawasaki, K., Murata, M., Ikeuchi, Y. and Ohnishi, S. 1997, Structural features of membrane fusion between influenza virus and liposome as revealed by quick-freezing electron microscopy. Journal of Cell Biology, 137, 1041–1056.
   PubMed Web of Science ®Google Scholar
• Kemble, G. W., Bodian, D. L., Rose, J., Wilson, I. A. and White, J. M. 1992, Intermonomer disulfide bonds impair the fusion activity of influenza virus hemagglutinin. Journal of Virology, 66, 4940-4950.
   Google Scholar
• Kemble, G. W., Danieli, T. and White, J. M. 1994, Lipid-anchored influenza hemagglutinin promotes hemifusion, not complete fusion. Cell, 76, 383–391.
   PubMed Web of Science ®Google Scholar
• Knutton, S. 1979, Studies of membrane fusion. III. Fusion of erythrocytes with polyethylene glycol. Journal of Cell Science, 36, 61–72.
   Google Scholar
• Knutton, S. and Pasternak, C. A. 1979, The mechanism of cell-cell fusion. TIBS, 4, 220–223.
   Web of Science ®Google Scholar
• Kozlov, M. M. and Chernomordik, L. V. 1998, A mechanism of protein-mediated fusion: coupling between refolding of the influenza hemagglutinin and lipid rearrangements. Biophysics Journal, 75, 1384–1396.
   PubMed Web of Science ®Google Scholar
• Krahling, H. 1981, Discrimination of two fusogenic properties of aqueous polyethylene glycol solutions. Zeitschrift fur Natur- forschung - Section C - Biosciences, 36, 593–596.
   Google Scholar
• Lee, J. and Lentz, B. R. 1997a, Evolution of lipidic structures during model membrane fusion and the relation of this process to cell membrane fusion. Biochemistry, 36, 6251–6259.
   PubMed Web of Science ®Google Scholar
• Lee, J. and Lentz, B. R. 1997b, Outer leaflet-packing defects promote poly(ethylene glycol)-mediated fusion of large unilamellar vesicles. Biochemistry, 36, 421–431.
   Google Scholar
• Lee, J. and Lentz, B. R. 1998, Secretory and viral fusion may share mechanistic events with fusion between curved lipid bilayers. Proceedings of the National Acadamy of Sciences (USA), 95, 9274–9279.
   Google Scholar
• Lentz, B. R. 1994, Polymer-induced membrane fusion: potential mechanism and relation to cell fusion events. Chemical and Physical Lipids, 73, 91–106.
   PubMed Web of Science ®Google Scholar
• Lentz, B. R., Carpenter, T. J. and Alford, D. R. 1987, Spontaneous fusion of phosphatidylcholine small unilamellar vesicles in the fluid phase. Biochemistry, 26, 5389–5397.
   PubMed Web of Science ®Google Scholar
• Lentz, B. R., McIntyre, G. F., Parks, D. J., Yates, J. C. and Massenburg, D. 1992, Bilayer curvature and certain amphipaths promote poly(ethylene glycol)-induced fusion of dipalmitoylpho- sphatidylcholine unilamellar vesicles. Biochemistry, 31, 2643-2653.
   PubMed Web of Science ®Google Scholar
• Lentz, B. R., Talbot, W., Lee, J. and Zheng, L. X. 1997, Transbilayer lipid redistribution accompanies poly(ethylene glycol) treatment of model membranes but is not induced by fusion. Biochemistry, 36, 2076–2083.
   Google Scholar
• Lentz, B. R., Wu, J. R., Zheng, L. and Prevratil, J. 1996, The interfacial region of dipalmitoylphosphatidylcholine bilayers is perturbed by fusogenic amphipaths. Biophysical Journal, 71, 3302–3310.
   Google Scholar
• Lindau, M. and Almers, W. 1995, Structure and function of fusion pores in exocytosis and ectoplasmic membrane fusion. Current Opinion in Cell Biology, 7, 509–517.
   PubMed Web of Science ®Google Scholar
• Massenburg, D. and Lentz, B. R. 1993, Poly(ethylene glycol)- induced fusion and rupture of dipalmitoylphosphatidylcholine large, unilamellar extruded vesicles. Biochemistry, 32, 9172-9180.
   PubMed Web of Science ®Google Scholar
• Melikyan, G. B., White, J. M. and Cohen, F. S. 1995, GPI-anchored influenza hemagglutinin induces hemifusion to both red blood cell and planar bilayer membranes. Journal of Cell Biology, 131, 679- 691.
   Google Scholar
• Monck, J. R. and Fernandez, J. M. 1992, The exocytotic fusion pore. Journal of Cell Biology, 119, 1395–1404.
   PubMed Web of Science ®Google Scholar
• Monck, J. R., Oberhauser, A. F. and Fernandez, J. M. 1995, The exocytotic fusion pore interface: a model of the site of neuro- transmitter release. Molecular Membrane Biology, 12, 151–156.
   PubMed Web of Science ®Google Scholar
• Moore, B. M., Lentz, B. R. and Meissner, G. 1978, Effects of sarcoplasmic reticulum Ca2 +-ATPase on phospholipid bilayer fluidity: boundary lipid. Biochemistry, 17, 5248–5255.
   PubMed Web of Science ®Google Scholar
• Nanavati, C., Markin, V. S., Oberhauser, A. F. and Fernandez, J. M. 1992, The exocytotic fusion pore modeled as a lipidic pore. Biophysical Journal, 63, 1118–1132.
   PubMed Web of Science ®Google Scholar
• Nir, S., Bentz, J., Wilschut, J. and Düzgüneş, N. 1983, Aggregation and fusion of phospholipid vesicles. Progress in Surface Science, 13, 1–124.
   Web of Science ®Google Scholar
• Nussler, F., Clague, M. J. and Herrmann, A. 1997, Meta-stability of the hemifusion intermediate induced by glycosylphosphatidylino- sitol-anchored influenza hemagglutinin. Biophysical Journal, 73, 2280–2291.
   PubMed Web of Science ®Google Scholar
• Oberhauser, A. F., Monck, J. R. and Fernandez, J. M. 1992, Events leading to the opening and closing of the exocytotic fusion pore have markedly different temperature dependencies. Kinetic analysis of single fusion events in patch-clamped mouse mast cells. Biophysical Journal, 61, 800–809.
   PubMedGoogle Scholar
• Ohki, S. 1984, Effects of divalent cations, temperature, osmotic pressure gradient, and vesicle curvature on phosphatidylserine vesicle fusion. Journal of Membrane Biology, 77, 265–275.
   PubMedGoogle Scholar
• Ohki, S. and Arnold, K. 1990, Surface dielectric constant, surface hydrophobicity and membrane fusion. Journal of Membrane Biology, 114, 195–203.
   PubMed Web of Science ®Google Scholar
• Ohno, H., Maeda, Y. and Tsuchida, E. 1981, 1H-NMR study of the effect of synthetic polymers on the fluidity, transition temperature and fusion of dipalmitoyl phosphatidylcholine small vesicles. Biochimica et Biophysica Acta, 642, 27–36.
   PubMedGoogle Scholar
• Perin, M. S. and MacDonald, R. C. 1989, Interactions of liposomes with planar bilayer membranes. Journal of Membrane Biology, 109, 221–232.
   PubMedGoogle Scholar
• Portis, A., Newton, C., Pangborn, W. and Papahadjopoulos, D. 1979, Studies on the mechanism of membrane fusion: evidence for an intermembrane Ca2 + -phospholipid complex, synergism with Mg2+, and inhibition by spectrin. Biochemistry, 18, 780- 790.
   Google Scholar
• Rafalski, M., Ortiz, A., Rockwell, A., van Ginkel, L. C., Lear, J. D., DeGrado, W. F. and Wilschut, J. 1991, Membrane fusion activity of the influenza virus hemagglutinin: interaction of HA2 N-terminal peptides with phospholipid vesicles. Biochemistry, 30, 10211-10220.
   PubMed Web of Science ®Google Scholar
• Rand, R. P. and Parsegian, V. A. 1988. The influence of polar group identify on the interactions between phospholipid bilayers. In Molecular Mechanisms of Membrane Fusion, S. Ohki, editor (New York: Plenum Press Publishing Corporation), pp. 73–81.
   Google Scholar
• Rand, R. P. and Sengupta, S. 1972, Cardiolipin forms hexagonal structures with divalent cations. Biochimica et Biophysica Acta, 255, 484–492.
   PubMed Web of Science ®Google Scholar
• Rapaport, D., Nir, S. and Shai, Y. 1994, Capacities of pardaxin analogues to induce fusion and leakage of negatively charged phospholipid vesicles are not necessarily correlated. Biochemis- try, 33, 12615–12624.
   PubMed Web of Science ®Google Scholar
• Ravoo, B. J., Kevelam, J., Weringa, W. D. and Engberts, J. B. F. N. 1998, Evidence for the `cold fusion’ of bilayer membranes. Journal of Physical Chemistry, 102, 11001–11006.
   Google Scholar
• Rizo, J. and Sudhof, T. C. 1998, Mechanics of membrane fusion [news]. Natural Structure Biology, 5, 839–842.
   PubMedGoogle Scholar
• Ruigrok, R. W., Aitken, A., Calder, L. J., Martin, S. R., Skehel, J. J., Wharton, S. A., Weis, W. and Wiley, D. C. 1988, Studies on the structure of the influenza virus haemagglutinin at the pH of membrane fusion. Journal of Gene Virology, 69, 2785–2795.
   PubMed Web of Science ®Google Scholar
• Saez, R., Alonso, A., Villena, A. and Goni, F. M. 1982, Detergent-like properties of polyethyleneglycols in relation to model membranes. FEBS Letters, 137, 323–326.
   Google Scholar
• Sassaroli, M., Vauhkonen, M., Perry, D. and Eisinger, J. 1990, Lateral diffusivity of lipid analogue excimeric probes in dimyristoylpho- sphatidylcholine bilayers. Biophysical Journal, 57, 281–290.
   PubMed Web of Science ®Google Scholar
• Schullery, S. E., Schmidt, C. F., Felgner, P., Tillack, T. W. and Thompson, T. E. 1980, Fusion of dipalmitoylphosphatidylcholine vesicles. Biochemistry, 19, 3919–3923.
   PubMed Web of Science ®Google Scholar
• Shangguan, T., Siegel, D. P., Lear, J. D., Axelsen, P. H., Alford, D. and Bentz, J. 1998, Morphological changes and fusogenic activity of influenza virus hemagglutinin. Biophysics Journal, 74, 54–62.
   PubMed Web of Science ®Google Scholar
• Siegel, D. P. 1987, Membrane-membrane interactions via inter- mediates in lamellar-to-inverted hexagonal phase rransitions. In Cell Fusion, A. E. Sowers, editor (New York: Plenum Press Publishing Corporation), pp. 181–207.
   Google Scholar
• Siegel, D. P. 1993, Modelling protein-induced fusion mechanisms: Insights from the relative stability of lipidic structures. In Viral Fusion Mechanisms, J. Bentz, editor (Boca raton, FL: CRC Press), pp. 475–512.
   Google Scholar
• Skehel, J. J., Bayley, P. M., Brown, E. B., Martin, S. R., Waterfield, M. D., White, J. M., Wilson, I. A. and Wiley, D. C. 1982, Changes in the conformation of influenza virus hemagglutinin at the pH optimum of virus-mediated membrane fusion. Proceedings of the National Academy of Sciences (USA), 79, 968–972.
   PubMed Web of Science ®Google Scholar
• Smith, C. L., Ahkong, Q. F., Fisher, D. and Lucy, J. A. 1982, Is purified poly(ethylene glycol) able to induce cell fusion? Biochimi- ca et Biophysica Acta, 692, 109–114.
   Google Scholar
• Spruce, A. E., Breckenridge, L. J., Lee, A. K. and Almers, W. 1990, Properties of the fusion pore that forms during exocytosis of a mast cell secretory vesicle. Neuron, 4, 643–654.
   PubMed Web of Science ®Google Scholar
• Spruce, A. E., Iwata, A. and Almers, W. 1991, The first milliseconds of the pore formed by a fusogenic viral envelope protein during membrane fusion. Proceedings of the National Academy of Sciences (USA), 88, 3623–3627.
   PubMed Web of Science ®Google Scholar
• Stegmann, T. 1993, Influenza hemagglutinin-mediated membrane fusion does not involve inverted phase lipid intermediates. Journal of Biological Chemistry, 268, 1716–1722.
   PubMed Web of Science ®Google Scholar
• Stegmann, T., White, J. M. and Helenius, A. 1990, Intermediates in influenza induced membrane fusion. Embo Journal, 9, 4231-4241.
   PubMed Web of Science ®Google Scholar
• Surewicz, W. K. 1983, Electron spin resonance study on the mechanism of polyethylene glycol-membrane interaction. FEBS Letters, 151, 228–232.
   PubMed Web of Science ®Google Scholar
• Sutton, R. B., Fasshauer, D., Jahn, R. and Brunger, A. T. 1998, Crystal structure of a SNARE complex involved in synaptic exo- cytosis at 2.4 A resolution [see comments]. Nature, 395, 347- 353.
   Google Scholar
• Suurkuusk, J., Lentz, B. R., Barenholz, Y., Biltonen, R. L. and Thompson, T. E. 1976, A calorimetric and fluorescent probe study of the gel-liquid crystalline phase transition in small, single- lamellar dipalmitoylphosphatidylcholine vesicles. Biochemistry, 15, 1393–1401.
   PubMed Web of Science ®Google Scholar
• Szoka, F. E., Magnusson, K. E., Wojcieszyn, J. W., Hou, Y., Derzko, Z. and Jacobson, K. A. 1981, Use of lectins and polyethylene glycol for fusion of glycolipid-containing liposomes with eukaryotic cells. Proceedings of the National Academy of Science (USA), 78. 1685–1689.
   Google Scholar
• Talbot, W. A., Zheng, L. X. and Lentz, B. R. 1997, Acyl chain unsaturation and vesicle curvature alter outer leaflet packing and promote poly(ethylene glycol)-mediated membrane fusion. Biochemistry, 36, 5827–5836.
   PubMedGoogle Scholar
• Tolar, L. A. and Pallanck, L. 1998, NSF function in neurotransmitter release involves rearrangement of the SNARE complex down- stream of synaptic vesicle docking. Journal of Neuroscience, 18, 10250–10256.
   PubMed Web of Science ®Google Scholar
• Tse, F. W., Iwata, A. and Almers, W. 1993, Membrane flux through the pore formed by a fusogenic viral envelope protein during cell fusion. Journal of Cell Biology, 121, 543–552.
   PubMed Web of Science ®Google Scholar
• Weber, T., Zemelman, B. V., McNew, J. A., Westermann, B., Gmachl, M., Parlati, F., Sollner, T. H. and Rothman, J. E. 1998, Snarepins —minimal machinery for membrane fusion. Cell, 92, 759–772.
   PubMed Web of Science ®Google Scholar
• Weissenhorn, W., Carfi, A., Lee, K. H., Skehel, J. J. and Wiley, D. C. 1998, Crystal structure of the ebola virus membrane fusion subunit, Gp2, from the envelope glycoprotein ectodomain. Molecular Cell, 2, 605–616.
   PubMed Web of Science ®Google Scholar
• Weissenhorn, W., Dessen, A., Harrison, S. C., Skehel, J. J. and Wiley, D. C. 1997, Atomic structure of the ectodomain from HIV-1 gp41 [see comments]. Nature, 387, 426–430.
   PubMed Web of Science ®Google Scholar
• Wharton, S. A., Calder, L. J., Ruigrok, R. W., Skehel, J. J., Steinhauer, D. A. and Wiley, D.C. 1995, Electron microscopy of antibody complexes of influenza virus haemagglutinin in the fusion pH conformation. Embo Journal, 14, 240–246.
   PubMed Web of Science ®Google Scholar
• White, J. M. 1990, Viral and cellular membrane fusion proteins. Annual Review of Physiology, 52, 675–697.
   PubMed Web of Science ®Google Scholar
• Wilson, I. A., Skehel, J. J. and Wiley, D. C. 1981, Structure of the haemagglutinin membrane glycoprotein of influenza virus at 3 A resolution. Nature, 289, 366–373.
   PubMed Web of Science ®Google Scholar
• Wu, H., Zheng, L. and Lentz, B. R. 1996, A slight asymmetry in the transbilayer distribution of lysophosphatidylcholine alters the surface properties and poly(ethylene glycol)-mediated fusion of dipalmitoylphosphatidylcholine large unilamellar vesicles. Bio- chemistry, 35, 12602–12611.
   Google Scholar
• Wu, J. R. and Lentz, B. R. 1991, Fourier transform infrared spectroscopic study of Ca2+ and membrane- induced secondary structural changes in bovine prothrombin and prothrombin fragment 1. Biophysics Journal, 60, 70–80.
   PubMedGoogle Scholar
• Wu, J. R. and Lentz, B. R. 1994, A method for quantitative interpretation of fluorescence detection of poly(ethylene glycol)- mediated 1-palmitoyl-2- [[[2-[4- (phenyl-trans-1.3.5-hexatrienyl)- phenyl] ethyl] oxyl] carboxyl] 3-sn -phosphatidylcholine (DPHpPC) transfer between phospholipid vesicles in the dehydrated state. Journal of Fluorescence, 4, 153–163.
   Google Scholar
• Yamazaki, M., Ohnishi, S. and Ito, T. 1989, Osmoelastic coupling in biological structures: decrease in membrane fluidity and osmo- phobic association of phospholipid vesicles in response to osmotic stress. Biochemistry, 28, 3710–3715.
   PubMed Web of Science ®Google Scholar
• Zimmerberg, J., Blumenthal, R., Sarkar, D. P., Curran, M. and Morris, S. J. 1994, Restricted movement of lipid and aqueous dyes through pores formed by influenza hemagglutinin during cell fusion. Journal of Cell Biology, 127, 1885-1894.
   PubMed Web of Science ®Google Scholar
• Zimmerberg, J., Curran, M. and Cohen, F. S. 1991, A lipid/protein complex hypothesis for exocytotic fusion pore formation. Annals of the New York Academy of Sciences, 635, 307–317.
   PubMed Web of Science ®Google Scholar
• Zimmerberg, J., Vogel, S. S. and Chernomordik, L. V. 1993, Mechanisms of membrane fusion. Annual Review of Biophysics and Biomolecular Structure, 22, 433–466.
   PubMedGoogle Scholar