Role of vesicle-mediated transport pathways in hepatocellular bile secretion

Seminars in Liver Disease

Volumn 16, Issue 2, 1996, Pages 169-189

  Crawford, James M ^a  

^a BRIGHAM AND WOMEN S HOSPITAL   (United States)

Author keywords

Bile canaliculus; Bile salt; Liver; Membrane; Protein; Trafficking

Indexed keywords

BILE ACID; CHOLESTEROL;

ANIMAL CELL; ARTICLE; BILE ACID SYNTHESIS; BILE FLOW; CELL MEMBRANE TRANSPORT; CHOLESTEROL TRANSPORT; ENDOCYTOSIS; INTRACELLULAR TRANSPORT; INTRAHEPATIC BILE DUCT; LIVER CELL; MEMBRANE VESICLE; NONHUMAN; PRIORITY JOURNAL; RAT; TRANSPORT KINETICS;

ANIMALS; BILE; BILE ACIDS AND SALTS; BILE CANALICULI; BIOLOGICAL TRANSPORT, ACTIVE; ENDOCYTOSIS; HUMANS; LIPID METABOLISM; LIVER; PROTEINS; RATS;

EID: 0030007590     PISSN: 02728087     EISSN: None     Source Type: Journal    
DOI: 10.1055/s-2007-1007230     Document Type: Article

References (252)

1. Phillips MJ, Poucell S, Petternson J, Valencia P: The liver: An atlas and text of ultrastructural pathology. New York, Raven Press, 1987. pp 1-35.
2. Mellman I, Yamamoto E, Whitney JA, et al: Molecular sorting in polarized and non-polarized cells: Common problems, common solutions. J Cell Sci 106(Suppl)17:1-7, 1993.
3. McIntyre JC, Sleight RG: Mechanisms of intracellular membrane lipid transport. Curr Topics Membr 40:453-81, 1994.
4. Saucan L, Palade GE. Membrane and secretory proteins are transported from the Golgi complex to the sinusoidal plasmalemma of hepatocytes by distinct vesicular carriers. J Cell Biol 125:733-41, 1994.
5. Gruenberg J, Maxfield FR: Membrane transport in the endocytic pathway. Curr Opin Cell Biol 7:552-63, 1995.
6. Lippincott-Schwartz J, Cole NB: Roles for microtubules and kinesin in membrane traffic between the endoplasmic reticulum and the Golgi copmlex. Biochem Soc Trans 23:544-8, 1995.
7. Oda H, Stockert RJ, Collins C, et al: Interaction of the microtubule cytoskeleton with endocytic vesicles and cytoplasmic dynein in cultured rat hepatocytes. J Biol Chem 270:15242-9, 1995.
8. Green RM, Crawford JM: Hepatocellular cholestasis: Pathobiology and histological outcome. Semin Liver Dis 15:372-89, 1995.
9. Meier PJ: The bile secretory polarity of hepatocytes. J Hepatol 9:124-9, 1989.
10. Suchy FJ: Hepatocellular transport of bile acids. Semin Liver Dis 13:235-47, 1993.
11. Radominska A, Treat S, Little J: Bile acid metabolism and the pathophysiology of cholestasis. Semin Liver Dis 13:219-34, 1993.
12. Nathanson MH, Boyer JL: Mechanisms and regulation of bile secretion. Hepatology 14:551-66, 1991.
13. Burwen SJ, Schmucker DL, Jones AL: Subcellular and molecular mechanisms of bile secretion. In: Jeon KW, Friedlander M (eds): International review of cytology: A survey of cell biology, Vol. 135. San Diego. Academic Press, Harcourt Brace Jovanovich, 1992, pp 269-313.
14. Reichen J, Simon FR, Cholestasis: In: Anas IM, Boyer JL, Fausto N, et al (eds): The liver: Biology and pathobiology. New York, Raven Press, 1994. pp 1291-326.
15. Verkade HJ, Vonk RJ, Kuipers F: Neu insights into the mechanism of bile acid-induced biliary lipid secretion. Hepatology 21:1174-89, 1995
16. Moseley RH: Cholestasis. In: Kaplowitz N (ed): Liver and biliary diseases Baltimore. Williams and Wilkins, 1992. pp 162-81.
17. Zaal KJM, Kok JW, Sormunen R, et al: Intracellular sites involved in the biogenesis of bile canaliculi in hepatic cells. Eur J Cell Biol 63:10-9, 1994.
18. Sormunen R, Eskelinen S, Lehto V-P: Bile canaliculus formation in cultured HEPG2 cells. Lab Invest 68:652-62, 1993.
19. Chiu J-H, Hu C, Lui W-Y, et al: The formation of bile canaliculi in human hepatoma cell lines. Hepatology 11:834-42, 1990.
20. Durand-Schneider A-M, Bouanga J-C, Feldmann G, Maurice M: Microtubule disruption interferes with the structural and functional integrity of the apical pole in primary cultures of rat hepatocytes. Eur J Cell Biol 56:260-8, 1991.
21. Crawford JM, Möckel G-M, Crawford AR, et al: Imaging biliary lipid secretion in the rat: Ultrastructural evidence for vesiculation of the hepatocyte canalicular membrane. J Lipid Res 36:2147-63, 1995.
22. Millington PF, Finean JB: Electron microscopic studies of the structure of the microvilli on principal epithelial cells of rat jejunum after treatment in hypo- and hypertonic saline. J Cell Biol 14:125-39, 1962.
23. Jones AL, Schmucker DL, Mooney JS, et al: Alterations in hepatic pericanalicular cytoplasm during enhanced bile secretory activity. Lab Invest 40:512-7, 1979.
24. Steer CJ, Klausner RD: Clathrin-coated pits and coated vesicles: Functional and structural studies. Hepatology 3:437-54, 1983.
25. Sakisaka S, Ng OC, Boyer JL: Tubulovesicular transcytotic pathway in isolated rat hepatocyte couplets in culture: Effect of colchicine and taurocholate. Gastroenterology 95:793-804, 1988.
26. Sakisaka S, Harada M, Gondo K, et al: Tubulovesicular transport of horseradish peroxidase in isolated rat hepatocyte couplets: Effects of low temperature, cytochalasin B and bile acids. Hepatology 20:1015-23, 1994.
27. Lake JR, Van Dyke RW, Scharschmidt BF: Acidic vesicles in cultured rat hepatocytes, Identification and characterization of their relationship to lysosomes and other storage vesicles. Gastroenterology 92:1251-61, 1987.
28. Jackle S, Runquist EA, Miranda-Brady S, Havel RJ: Trafficking of the epidermal growth factor receptor and transferrin in three hepatocytic endosomal fractions. J Biol Chem 266:1396-402, 1991.
29. Knight A, Hughson E, Hopkins CR, Cutler DF: Membrane protein trafficking through the common apical endosome compartment of polarized Caco-2 cells. Mol Biol Cell 6:597-610, 1995.
30. Alvaro D, Benedetti A, Gigliozzi A, et al: Effect of Brefeldm A on transcytotic vesicular pathway and bile secretion: A study on the isolated perfused rat liver and isolated rat hepatocyte couplets. Heparology 21:450-9, 1995.
31. Small DM: Phase equilibria and structure of dry and hydrated egg lecithin. J Lipid Res 8:551-9, 1967.
32. Tiffany JM, Bloug HA: Estimation of the number of surface projections on myxo- and paramyxoviruses. Virology 41:392-4, 1970.
33. DePierre JW, Andersson G, Dallner G: Endoplasmic reticulum and Golgi complex. In: Arias IM, Jakoby WB, Popper H, et al (eds): The liver: Biology and pathobiology. 2nd ed. New York, Raven Press, 1988, pp 165-87
34. Pley U, Parham P: Clathrin: Its role in receptor-mediated vesicular transport and specialized functions in neurons. Crit Rev Biochem Mol Biol 28:431-64, 1993.
35. Dunn WA, Connolly AP, Hubbard AL: Receptor-mediated endocytosis of Epidermal Growth Factor by rat hepatocytes: Receptor pathway. J Cell Biol 102:24-36, 1986.
36. Weibel ER, Staubli W, Gnagi HR, Hess FA: Correlated morphometric and biochemical studies on the liver cell. I. Morphometric model, stereological methods, and normal morphometric data for rat liver. J Cell Biol 42:68-91, 1969.
37. Van Deurs B, Petersen OW, Olsnes S, Sandvig K: The ways of endocytosis. Int Rev Cytol 117:131-78, 1989.
38. Van Dyke RW, Belcher JD. Acidification of three types of liver endocytic vesicles: Similarities and differences. Am J Physiol 266:C81-94, 1994.
39. Scharschmidt BF, Lake JR, Renner EL, et al: Fluid phase endocytosis by cultured rat hepatocytes and perfused rat liver: Implications for plasma membrane turnover and vesicular trafficking of fluid phase markers. Proc Natl Acad Sci USA 83:9488-92, 1986.
40. Lake JR, Licko V, Van Dyke RW, Scharschmidt BF: Biliary secretion of fluid-phase markers by the isolated perfused rat liver Role of transcellular vesicular transport. J Clin Invest 76:676-89, 1985.
41. Moss AL, Ward WF: Multiple pathways for ligand internalization in rat hepatocytes II: Effect of hyperosmolarity and contribution of fluid-phase endocytosis. J Cell Physiol 149:319-23, 1991.
42. Schweizer FE, Betz H, Augustine GJ: From vesicle docking to endocytosis: Intermediate reactions of exocytosis. Neuron 14:689-96, 1995.
43. Roma MG, Marinelli RA, Rodríguez Garay EA: Biliary excretion of polyethylene glycol molecular weight 900. Evidence for a bile salt-stimulated vesicular transport mechanism. Biochem Pharmacol 42:1775-81, 1991.
44. Kuipers F, Havinga R, Bosschieier H, et al: Enterohepatic circulation in the rat. Gastroenterology 88:403-11, 1985.
45. Rigotti A, Núñez L, Amigo L, et al: Biliary lipid secretion: Immunolocalization and identification of a protein associated with lamellar cholesterol carriers in supersaturated rat and human bile. J Lipid Res 34:1883-94, 1993.
46. Offner GD, Gong D, Afdhal NH: Identification of a 130-kilodalton human biliary concanavalin A binding protein as aminopeptidase N. Gastroenterology 106:755-62, 1994.
47. Weimer HE, Roberts DM, Vallanuevo P, Porter HG: Genetic differences in electrophoretic patterns during the phlogistic response in the albino rat. Comp Biochem Physiol 43B:965-73, 1972.
48. Harada M, Sakisaka S, Yoshitake M, et al: Ultrastructure of the intracellular membranous system of rat hepatocytes in intrahepatic cholestasis induced by phalloidin. J Hepatol 21:560-6, 1994.
49. Layden TJ, Boyer JL: Influence of bile acids on bile canalicular membrane morphology and the lobular gradient in canalicular size. Lab Invest 39:110-9, 1978.
50. LaRusso NF: Proteins in bile: How they get there and what they do Am J Physiol 247:G199-205, 1984.
51. Reuben A: Biliary proteins. Hepatolgy 4:46S-50S, 1984
52. Mullock BM, Shaw LJ, Fitzharris B, et al: Sources of protein in bile. Gut 26:500-9, 1985.
53. Burwen SJ, Jones AL: Hepatocellular processing of endocytosed proteins. J Electron Microsc Tech 14:140-51, 1990.
54. LeSage GD, Robertson WE, Baumgart MA: Bile acid-dependent vesicular transport of lysosomal enzymes into bile in the rat. Gastroenterology 105:889-900, 1993.
55. Accatino L, Figueroa C, Pizarro M, Solís N: Enhanced biliary excretion of canalicular membrane enzymes in estrogen-induced and obstructive cholestasis, and effects of different bile acids in the isolated perfused rat liver. J Hepatol 22:658-70, 1995.
56. Saucan L, Palade GE: Differential colchicine effects on the transport of membrane and secretory proteins in rat hepatocytes in vivo: Bipolar secretion of albumin. Hepatology 15.734-21, 1992.
57. Graham R, Karnovsky M: The early stages of injected horseradish peroxidase in the proximal tubules of mouse kidney: Ultrastructural chemistry by a new technique. J Histochem Cytochem 14:291-302, 1966.
58. Lowe PJ, Kan KS, Barnwell SG, et al: Transcytosis and paracelluar movements of horseradish peroxidase in the isolated perfused rat liver. Biochem J 229:529-37, 1985.
59. Hayakawa T, Cheng O, Ma A, Boyer JL: Taurocholate stimulates transcytotic vesicular pathways labeled by horseradish peroxidase in the isolated perfused rat liver. Gastroenterology 99:216-28, 1990.
60. ++ mechanisms defective in cholestasis. J Clin Invest 92:2984-93, 1993.
61. Hoshino M, Hirano A, Hayakawa T, et al: Comparative studies on bile flow and biliary lipid excretion after bile-acid loading in normal and partially hepatectomized rats. Biochem J 305:367-71, 1995.
62. Reynier MO, Hasfaieh IA, Crotte C, et al: Monensin action on the Golgi complex in perfused rat liver: Evidence against bile salt vesicular transport. Gastroenterology 102:2024-32, 1992.
63. Araki N, Ohno J, Lee T, et al: Nematolysosomes (elongate lysosomes) in rat hepatocytes: Their distribution, microtubule dependence, and role in endocytic transport pathway. Exp Cell Res 204:181-91, 1993.
64. Knapp PE, Swanson JA: Plasticity of the tubular lysosomal compartment in macrophages. J Cell Sci 95:433-9, 1990.
65. Smythe E, Warren G: The mechanism of receptor-mediated endocytosis. Eur J Biochem 202:689-99, 1991.
66. Quintart J, Baudhuin P, Courtoy PJ: Marker enzymes in rat liver vesicles involved in transcellular transport. Eur J Biochem 184:567-74, 1989.
67. Barroso M, Sztul ES: Basolateral to apical transcytosis in polarized cells is indirect and involves BFA and trimeric G protein sensitive passage through the apical endosome. J Cell Biol 124:83-100, 1994.
68. Jones AL, Schmucker DL, Renston RH, Murakami T: The architecture of bile secretion: A morphological perspective of physiology. Dig Dis Sci 25:609-29, 1980.
69. Nuñez M, Gaete V, Watkias JA, Glass J: Mobilization of iron from endocytic vesicles: The effects of acidification and reduction. J Biol Chem 265:6688-92, 1990.
70. Goltz JS, Wolkoff AW, Novikoff PM, Stocken RJ, Satir P: A role for microtubules in sorting endocytic vesicles in rat hepatocytes. Proc Natl Acad Sci USA 89:7026-30, 1992.
71. Schiff JM, Fisher MM, Underdown BJ: Receptor-mediated biliary transport of immunoglobulin A and asialoglycoprotein: Sorting and missorting of ligands revealed by two radiolabeling methods. J Cell Biol 98:79-89, 1984.
72. Regoeczi E, Chindemi PA: Translocation of different forms of transferrin from blood to bile in the rat. Hepatology 21:1063-9, 1995.
73. Kim DC, Sugiyama Y, Sato H, et al: Kinetic analysis of in vivo receptor-dependent binding of human epidermal growth factor by rat tissues. J Pharm Sci 77:200-7, 1988.
74. Coffey RJ Jr. Kost LJ, Lyons RM, et al: Hepatic processing of transforming growth factor b in the rat: Uptake, metabolism, and biliary excretion. J Clin Invest 80:750-7, 1987.
75. 125I]insulin Gastroenterology 78:1373-88, 1980.
76. Nakano A, Marks DL, Tietz PS, et al. Quantitative importance of biliary excretion to the turnover of hepatic lysosomal enzymes. Hepatology 22:262-6, 1995.
77. LaRusso NF, Fowler S: Coordinate secretion of acid hydrolases in rat bile: Hepatocyte exocytosis of lysosomal protein? J Clin Invest 64:948-54, 1979.
78. LeSage GD, Robertson WE, Baumgart MA: Demonstration of vesicular-dependent bile flow in the sucrose-loaded rat. Gastroenterology 99:478-87, 1990.
79. Sewell RB, Barham SS, Zinsmeister AR, LaRusso NF: Microtubule modulation of biliary excretion of endogenous and exogenous hepatic lysosomal constituents. Am J Physiol 246:G8-15, 1984.
80. Sewell RB, Barham SS, LaRusso NF: Effect of chloroquine on the form and function of hepatic lysosomes. Morphologic modifications and physiologic alterations related to the biliary excretion of lipids and proteins. Gastroenterology 85:1146-53, 1983.
81. Hardison WGM, Weiner RG, Hatoff DE, Miyai K: Similarities and differences between models of extrahepatic biliary obstruction and complete biliary retention without obstruction in the rat. Hepatology 3:383-90, 1983
82. Spies M Hiraziker W, Lodish HG, Sememza G: Molecular cell biology of brush border hydrolases: Sucrose-isomaltase and gamma-glutamyl transpeptidase. In: Kenny J (ed): Mammalian ectoenzymes. Amsterdam, Elsevier, 1987. pp 87-110.
83. Scott LJ, Hubbard AL: Dynamics of four rat liver plasma membrane proteins and polymeric IgA receptor. Rates of synthesis and selective loss into the bile. J Biol Chem 267:6099-106, 1992.
84. Low MG: Biochemistry of the glycosyl-phosphatidylinositol membrane protein anchors. Biochem J 244:1-13, 1987
85. Holdsworth G, Coleman R: Enzyme profiles of mammalian bile. Biochim Biophys Acta 389:47-50, 1975.
86. Hatoff DE, Hardison WG: Bile acid-dependent secretion of alkaline phosphatase in rat bile. Hepatology 2:433-9, 1982.
87. Barnwell SG, Lowe PJ, Coleman R: Effect of taurochenodeoxycholate or tauroursodeoxycholate upon biliary output of phospholipids and plasma-membrane enzymes, and the extent of cell damage, in isolated perfused rat livers. Biochem J 216:107-11, 1983.
88. Mullock BM, Dobrota M, Hinton RH: Sources of proteins of rat bile. Biochim Biophys Acta 543:497-507, 1978.
89. Evans WH, Kremmer T, Culvenor JG: Role of membranes in bile formation Comparison of the composition of bile and a liver-bile canalicular plasma-membrane subfraction. Biochem J 154:589-95, 1976.
90. Barnwell SG, Godfrey PP, Lowe PJ, Coleman R: Biliary protein output by isolated perfused rat livers. Biochem J 210:549-57, 1983.
91. Núñez L, Amigo L, Mingrone G, et al: Biliary aminopeptidase-N and the cholesterol crystallization defect in cholelithiasis. Gut 37:422-6, 1995.
92. Schachter D: The hepatocyte plasma membrane: Organization and differentiation. In: Arias IM, Boyer JL, Fausto N, et al (eds): The liver: Biology and pathobiology. 3rd ed. New York, Raven Press, 1994. p 145-54.
93. Sztul E, Colombo M Stahl P, Samanta R: Control of protein traffic between distinct plasma membrane domains: Requirement for a novel 108,000 protein in the fusion of transcytotic vesicles with the apical plasma membrane. J Biol Chem 268:1876-85, 1993.
94. Bartles JR, Feracci HM, Stieger B, Hubbard AL: Biogenesis of the rat hepatocyte plasma membrane in vivo: Comparison of the pathways taken by apical and basolateral proteins using subcellular fractionation. J Cell Biol 105:1241-51, 1987.
95. Bartles JR, Hubbard AL. Biogenesis of the rat hepatocyte plasma membrane. Methods Enzymol 191:825-40, 1990.
96. Maurice M, Schell MJ, Lardeux B, Hubbard AL: Biosynthesis and intracellular transport of a bile canalicular plasma membrane protein: Studies in vivo and in the perfused rat liver. Hepatology 19:648-55, 1994.
97. Barr VA, Hubbard AL: Newly synthesized hepatocyte plasma membrane proteins are transported in transcytotic vesicles in the bile duct-ligaied rat. Gastroenterology 105-554-71, 1993.
98. Hubbard AL, Barr VA, Scott LJ: Hepatocyte surface polarity. In: Arias IM, Boyer JL, Fausto N, et al (eds): The liver: Biology and pathobiology. New York, Raven Press, 1994. pp 189-213.
99. Ihrke G, Neufeld EB, Meads T, et al: WIF-B cells: An in vitro model for studies of hepatocyte polarity. J Cell Biol 123:1761-75, 1993.
100. Bruck R, Haddad P, Graf J, Boyer JL: Regulatory volume decrease stimulates bile flow, bile acid excretion, and exocytosis in isolated perfused rat liver Am J Physiol 262:G806-812, 1992.
101. Häussinger D, Saha N, Hallbrucker C, et al. Involvement of microtubules in the swelling-induced stimulation of transcellular taurocholate transport in perfused rat liver, Biochem J 291:355-60, 1993.
102. Brack R, Benedetti A, Strazzabosco M, Boyer JL: Intracellular alkalinization stimulates bile flow and vesicular-mediated exocytosis in IPRL. Am J Physiol 265:G347-53, 1993.
103. + +], increase in pancreatic acinar cells and its relation to amylase secretion. Am J Physiol 259:G792-801, 1990.
104. Graf J, Rupnik M, Zupancic G, Zorec R: Osmotic swelling of hepatocytes increases membrane conductance but not membrane capacitance. Biophys J 68:1359-63, 1995.
105. Landmann L, Meier PJ, Bianchi L: Bile duct ligation-induced redistribution of canalicular antigen in rat hepatocyte plasma membranes demonstrated by immunogold quantitation. Histochemistry 94:373-9, 1990.
106. Fricker G, Landmann L, Meier PJ: Extrahepatic obstructive cholestasis reverses the bile salt secretory polarity of rat hepatocytes. J Clin Invest 84:876-85, 1989.
107. Stieger B, Meier PJ, Landmann L: Effect of obstructive cholestasis on membrane traffic and domain-specific expression of plasma membrane proteins in rat liver parenchymal cells. Hepatology 20.201-12, 1994.
108. Matter A, Orci L, Rouiller C: A study on the permeability barriers between Disse's space and the bile canaliculi. J Ultrastruct Res 11(Suppl):1-71, 1969.
109. Jones AL, Hradek GT, Schmucker DL, Underdown BJ: The rate of polymeric and secretory immunoglobulin A after retrograde infusion into the common bile duct in rats. Hepatology 4:1173-83, 1984.
110. Hoekstra D, Kok JW: Trafficking of glycosphingohpids in eukaryotic cells: Sorting and recycling of lipids. Biochim Biophys Acta 1113:277-94, 1992.
111. van Generen IL, Van Meer G: Phospholipid translocation, asymmetry and membrane fusion. Biochem Soc Trans 21:235-9, 1993.
112. Rosenwald AG, Pagano RE: Intracellular transport of ceramide and its metabolites at the Golgi complex: Insights from short-chain analogs. Adv Lipid Res 26:101-16, 1993.
113. Young WW Jr: Molecular approaches to studying the intracellular trafficking of glycosphingolipids. Adv Lipid Res 26:161-79, 1993.
114. Trotter PJ, Voelker DR: Lipid transport processes in eukaryotic cells. Biochim Biophys Acta 1213:241-62, 1994.
115. Kok JW, Hoekstra D: Glycosphingolipid trafficking in the endocytic pathway. Curr Topics Membr 40:503-37, 1994.
116. Van't Hof W, Van Meer G: Lipid polarity and sorting in epithehal cells. Curr Topics Membr 40:539-63, 1994.
117. Schnitzer E, Lichtenberg D: Re-evaluation of the structure of low density lipoproteins. Chem Phys Lipids 70:63-74, 1994.
118. Van Antwerpen R, Gilkey JC: Cryo-electron microscopy reveals human low density lipoprotein substructure. J Lipid Res 35:2223-31, 1995.
119. Carey MC, LaMont JT: Cholesterol gallstone formation. 1. Physical-chemistry of bile and biliary lipid secretion. Prog Liv Dis 10:139-63, 1992.
120. Robins SJ, Fasulo JM, Robins VF, Patton GM: Utilization of different fatty acids for hepatic and biliary phosphatidylcholine formation and the effect of changes in phosphatidylcholine molecular species on biliary lipid secretion. J Lipid Res 32:985-92, 1991.
121. Patton GM, Fasulo JM, Robins SJ: Hepatic phosphatidylcholines: Evidence for synthesis in the rat by extensive reutilization of endogenous acylglycerides. J Lipid Res 35:1211-21, 1994.
122. Patton GM, Fasulo JM, Robins SJ: Origin of bile lecithins: Evidence for extensive remodeling and a novel pathway of synthesis. Gastroenterology 104:A970, 1993.
123. Chanussot F, Lafont H, Hauton J, Tuchweber B, Yousef I. Studies on the origin of biliary phospholipid. Effect of dehydrocholic acid and cholic acid infusions on hepatic and biliary phospholipids. Biochem J 270:691-5, 1990.
124. Schersten T, Nilsson S, Cahlin E: Relationship between the biliary excretion of bile acids and the excretion of water, lecithin, and cholesterol in man. Eur J Clin Invest 1.242-7, 1971.
125. Cohen DE, Angelico M, Carey MC: Structural alterations in lecithin-cholesterol vesicles following interactions with monomeric and micellar bile salts: Physical-chemical basis for subselection of biliary lecithin species and aggregative states of biliary lipids during bile formation. J Lipid Res 31:55-70, 1990.
126. Cohen DE, Leonard MR, Carey MC: In vitro evidence that phospholipid secretion into bile may be coordinated intracellularly by the combined actions of bile salts and the specific phosphatidylcholine transfer protein of liver. Biochemistry 33:9975-80, 1994.
127. Leonard AN, Cohen DE: In vitro evidence that sterol carrier protein 2 may provide a scavenger pathway for phosphatidylcholine secretion into bile. Hepatology 22:295A, 1995.
128. Crawford JM, Berken CA, Gollan JL: Role of the hepatocyte microtubular system in the excretion of bile salts and biliary lipid: Implications for intracellular vesicular transport. J Lipid Res 29:144-56, 1988
129. Coleman R, Rahman K: Lipid flow in bile formation. Biochim Biophys Acta 1125:113-33, 1992.
130. Bilhartz LE, Dietschy JM: Bile salt hydrophobiticy influences cholesterol recruitment from rat liver in vivo when cholesterol synthesis and uptake are constant. Gastroenterology 95:771-9, 1988.
131. Scheibner J, Fuchs M, Hörmann E, et al: Biliary cholesterol secretion and bile acid formation in the hamster: The role of newly synthesized cholesterol. J Lipid Res 35:690-7, 1994.
132. Robins SJ, Fasulo JM, Lessard PD, Patton GM: Hepatic cholesterol synthesis and the secretion of newly synthesized cholesterol in bile. Biochem J 289:41-4, 1993.
133. Van Meer G, Simons K: The function of tight junctions in maintaining differences in lipid composition between the apical and the basolateral cell surface domains of MDCK cells. EMBO J 5:1455-64, 1986.
134. Robins SJ, Fasulo JM, Collins MA, Patton GM: Evidence for separate pathways of transport of newly synthesized and preformed cholesterol into bile. J Biol Chem 260:6511-3, 1985.
135. Robins SJ, Fasulo JM, Salen G, Patton GM: Preferential sterol transport from HDL to bile. Hepatology 22:240A, 1995.
136. Acton S, Rigotti A, Landschulz KT, et al: Identification of scavenger receptor SR-BI as a High Density Lipoprotein receptor. Science 271:518-20, 1996.
137. Sleight RG: Intracellular lipid transport in eukaryotes. Annu Rev Physiol 49:193-208, 1987.
138. Puglielli L, Rigotti A, Greco AV, et al: Sterol carrier protein-2 is involved in cholesterol transfer from the endoplasmic reticulum to the plasma membrane in human fibroblasts. J Biol Chem 270:18723-6, 1995.
139. Puglielli L, Rigotti A, Amigo L, et al: Evidence for the involvement of sterol carrier protein (SCP-2) in a rapid, microtubule-, and Golgi-independent transport of newly-synthesized cholesterol to bile. Gastroenterology 108:A1150, 1995.
140. Vanier MT, Pentchev P, Rodriguez-Lafrasse C, Rousson R: Niemann-Pick disease type C: An update. J Inherited Metab Dis 14:580-95, 1991.
141. Liscum L, Underwood KW: Intracellular cholesterol transport and compartmentation. J Biol Chem 270:15443-6, 1995.
142. Lange Y, Steck TL: Cholesterol-rich intracellular membranes: A precursor to the plasma membrane. J Biol Chem 260:15592-7, 1985.
143. Allan D, Kallen K: Transport of lipids to the plasma membrane in animal cells. Prog Lipid Res 32:195-219, 1993.
144. Roman ID, Thewles A, Coleman R: Fractionation of livers following diosgenin treatment to elevate biliary cholesterol. Biochim Biophys Acta Lipids Lipid Metab 1255:77-81, 1995.
145. Koval M, Pagano RE: Intracellular transport and metabolism of sphingomyelin. Biochim Biophys Acta 1082:113-25, 1991.
146. Jeckel D, Karrenbauer A, Birk R, et al: Sphingomyelin is synthesized in the cis Golgi. FEBS Lett 261:155-7, 1990.
147. Futerman AH, Stieger B, Hubbard AL, Pagano RE: Sphingomyelin synthesis in rat liver occurs predominantly at the cis and medial cisternae of the Golgi apparatus. J Biol Chem 265:8650-7, 1990.
148. Koval M, Pagano RE: Lipid recycling between the plasma membrane and intracellular compartments: Transport and metabolism of fluorescent sphingomyelin analogues in cultured fibroblasts. J Cell Biol 108:2169-81, 1989.
149. Lipsky NG, Pagano RE: Intracelluar translocation of fluorescent sphingolipids in cultured fibroblasts: Endogenously synthesized sphingomyelin and glucocerebroside analogues pass through the Golgi apparatus en route to the plasma membrane. J Cell Biol 100:27-34, 1985.
150. Van Meer G, Van't Hof W: Epithelial sphingolipid sorting is insensitive to reorganization of the Golgi by nocodazole, but is abolished by monensin in MDCK cells and by brefeldin A in Caco-2 cells. J Cell Sci 104:833-42, 1993.
151. Shiao Y-J, Vance JE: Sphingomyelin transport to the cell surface occurs independently of protein secretion in rat hepatocytes. J Biol Chem 268:26085-92, 1993.
152. Boyer JL, Soroka CJ: Vesicle targeting to the apical domain regulates bile excretory function in isolated rat hepatocyte couplets. Gastroenterology 109:1600-11, 1995.
153. Gautam A, Ng O, Strazzabosco M, Boyer JL: Quantitative assessment of canalicular bile formation in isolated hepatocyte couplets using microscopic optical planimetry. J Clin Invest 83:565-73, 1989.
154. Hay DW, Carey MC: Chemical species of lipids in bile. Hepatology 12(Suppl)6S-16S, 1990.
155. Oude Elfermk RPJ, Bakker CTM, Roelofsen H, et al: Accumulation of organic anion in intracellular vesicles of cultured rat hepatocytes is mediated by the canalicular multispecific organic anion transporter. Hepatology 17:434-44, 1993.
156. 2. Int J Biochem 22:1453-5, 1990.
157. Koval M, Pagano RE: Sorting of an internalized plasma membrane lipid between recycling and degradative pathways in normal and Niemann-Pick, type A fibroblasts. J Cell Biol 111:429-42, 1990.
158. Carey MC, Duane WC: Enterohepatic circulation. In: Anas IM, Boyer JL, Fausto N, et al (eds): The liver: Biology and pathobiology. New York, Raven Press, 1994, pp 719-67.
159. Stolz A, Hammond L, Lou H, et al: cDNA cloning and expression of the human hepatic bile acid-binding protein. A member of the monomeric reductase gene family. J Biol Chem 268:10448-57, 1993.
160. Takikawa H, Stolz A, Kaplowitz N: Cyclical oxidation-reduction of the C3 position on bile acids catalyzed by rat hepatic 3α-hydroxysteroid dehydrogenase. I. Studies with the purified enzyme, isolated rat hepatocytes, and inhibition by indomethacin. J Clin Invest 80:852-60, 1987.
161. Takikawa H, Fernandez-Checa JC, Kuhlenkamp J, et al: Effect of indomethacin on the uptake, metabolism and excretion of 3-oxocholic acid: Studies in isolated hepatocytes and perfused rat liver. Biochim Biophys Acta 1084:247-50, 1991.
162. Dubin M, Maurice M, Feldmann G, Erlinger S: Influence of colchicine and phalloidin on bile secretion and hepatic ultrastructure in the rat: Possible interaction between microtubules and microfilaments Gastroenterology 79:646-54, 1980.
163. Gregory DH, Vlahcevic ZR, Prugh MF, Swell L: Mechanism of secretion of biliary lipids. Role of a microtubular system in hepatocellular transport of biliary lipids in the rat. Gastroenterology 74:93-100, 1978.
164. Barnwell SG, Lowe PJ, Coleman R: The effects of colchicine on secretion into bile of bile salts, phospholipids, cholesterol and plasma membrane enzymes: Bile salts are secreted unaccompanied by phospholipids and cholesterol. Biochem J 220:723-31, 1984.
165. Crawford JM, Crawford AR, Strahs DCJ: Microtubule-dependent transport of bile salts through hepatocytes: Cholic vs. taurocholic acid. Hepatology 18:903-11, 1993.
166. Erlmger S: Role of intracellular organelles in the hepatic transport of bile acids. Biomed Pharmacother 44:409-16, 1990.
167. Crawford JM, Strahs DCJ, Crawford AR, Barnes S: The role of bile salt hydrophobicity in hepatic microtubule-dependent bile salt secretion. J Lipid Res 35:1738-48, 1994.
168. Crawford JM, Gollan JL: Transcellular transport of organic anions in hepatocytes: Still a long way to go (Editorial). Hepatology 14:192-7, 1991.
169. Suchy FJ, Balistreri WF, Hung J, et al: Intracellular bile acid transport in rat liver as visualized by electron microscope autoradiography using a bile acid analogue. Am J Physiol 245:G681-9, 1983.
170. Goldsmith MA, Huling S, Jones AJL: Hepatic handling of bile salts and protein in the rat during intrahepatic cholestasis. Gastroenterology 84 978-86, 1983.
171. Lamri Y, Roda A, Dumont M, et al: Immunoperoxidase localization of bile salts in rat liver cells: Evidence for a role of the Golgi apparatus in bile salt transport. J Clin Invest 82:1173-82, 1988.
172. Crawford JM, Barnes S, Stearns RC, et al: Ultrastructural localization of a fluorinated bile salt in hepatocytes. Lab Invest 71:42-51, 1994.
173. Lipsky NG, Pagano RE: A vital stain for the Golgi apparatus. Science 228:745-7, 1985.
174. Crawford JM, Vinter DW, Gollan JL: Taurocholate induces pericanalicular localization of C6-NBD-ceramide in isolated hepatocytes couplets. Am J Physiol 260:G119-32, 1991.
175. Kitamura T, Gatmaitan Z, Arias IM: Serial quantitative image analysis and confocal microscopy of hepatic uptake, intracellular distribution and biliary secretion of a fluorescent bile acid analog in rat hepatocyte couplets. Hepatology 12:1358-64, 1990.
176. Simion FA, Fleischer B, Fleischer S. Subcellular distribution of bile acids, bile salts and taurocholate binding sites in rat liver. Biochemistry 23:6459-66, 1984.
177. Simion FA, Fleischer B, Fleischer S: Two distinct mechanisms for taurocholate uptake in subcellular fractions from rat liver J Biol Chem 250:10814-22, 1984.
178. Verkade JJ, Wolbers MJ, Havinga R, et al: The uncoupling of biliary lipid from bile acid secretion by organic anions in the rat. Gastroenterology 99:1485-92, 1990.
179. Apstem MD: Inhibition of biliary phospholipid and cholesterol secretion by bilirubin in the Sprague-Dawley and Gunn rat. Gastroenterology 87:634-8, 1984.
180. Apstein MD, Russo AR. Ampicillin inhibits biliary cholesterol secretion. Dig Dis, Sci 30:253-6, 1985.
181. Monte MJ, Parslow RA, Coleman R: Inhibitory action of cyclobutyrol on the secretion of biliary cholesterol and phospholipids. Biochem J 266:165-71, 1990.
182. Bellringer ME, Steele NJ, Rahman K, Coleman R: Ampicillin inhibits the movement of biliary secretory vesicls in rat hepatocytes. Biochim Biophys Am 941:71-5, 1988.
183. Yamashita G, Tazuma S, Horikawa K, et al: Partial characterization of mechanism(s) by which sulphobromophthalein reduces biliary lipid secretion. Biochem J 291:173-7, 1993.
184. Cava F, Gonzalez J, Gonzalez-Buitrago JM, Muriel C, Jimenez R: Inhibition of biliary cholesterol and phospholipid secretion by cefmetazole. The role of vesicular transport and of canalicular events. Biochem J 275:591-5, 1991.
185. Yamashita G, Tazuma S, Kajiyama G: Effects of organic anions on biliary
186. Deroubaix X, Coche T, Depiereux E, Feytmans E: Saturation of hepatic transport of taurocholate in rats in vivo. Am J Physiol 260:G189-96, 1991.
187. Weinman SA, Maglova LM: Free concentrations of intracellular fluorescent anions determined by cytoplasmic dialysis of isolated hepatocytes. Am J Physiol 267:G922-31, 1994.
188. Coche T, Deroubaix X, Depiereux E, Feytmans E: Compartmental analysis of steady-state taurocholate transport kinetics by isolated rat hepatocytes. Hepatology 13:1203-14, 1991.
189. Hayakawa T, Katagiri K, Hoshino M, et al: Papaverine inhibits transcytotic vesicle transport and lipid excretion into bile in isolated perfused rat liver. Hepatology 16:1036-42, 1992.
190. Schubert R, Beyer K, Wolburg H, Schmidt K: Structural changes in membranes of large unilamellar vesicles after binding of sodium cholate. Biochemistry 25:5263-9, 1986.
191. Lowe PJ, Barnwell SG, Coleman R: Rapid kinetic analysis of the bile-salt-dependent secretion of phospholipid, cholesterol and a plasma-membrane enzyme into bile. Biochem J 222:631-7, 1984.
192. Kumai T, Hoshino M, Hayakawa T, Higashi K: Papaverine inhibits bile acid excretion in isolated perfused rat liver. Hepatology 20:692-9, 1994.
193. Nakai T, Katagiri K, Hoshino M, et al: Microtubule-independent choleresis and anti-cholestatic action of tauroursodeoxycholate in colchicine-treated rat liver. Biochem J 288:613-7, 1992.
194. Tanaka A, Katagiri K, Hoshino M, et al: Endothelin-1 stimulates bile acid secretion and vesicular transport in the isolated perfused rat liver. Am J Physiol 266:G324-9, 1994
195. LeSage GD, Schteingart CD, Hofmann AF: Effect of bile acid hydrophobicity on biliary transit time and intracellular mobility: A comparison of seven fluorescent bile acid analogues. Hepatology 20:256A, 1994.
196. Alves C, Von Dippe P, Amoui M, Levy D: Bile acid transport into hepatocyte smooth endoplasmic reticulum vesicles is mediated by microsomal epoxide hydrolase, a membrane protein exhibiting two distinct topological orientations. J Biol Chem 268:20148-55, 1993.
197. Reuben A, Allen RM. Taurocholate transport by rat liver golgi vesicles (Abstract). Gastroenterology 98:A624, 1990.
198. Berg C, Gollan JL: Taurocholate transport by hepatocyte plasma and intracellular membranes: Evaluation by "ultra-rapid" filtration. Hepatology 14:146A, 1991.
199. Marks DL, LaRusso MF, McNiven MA: Isolation of the microtubule-vesicle motor kinesin from rat liver: Selective inhibition by cholestatic bile acids Gastroenterology 108:824-33, 1995.
200. Higgins JV, Paul JM, Dumaswala R, Heubi JE: Downregulation of taurocholate transport by ileal BBM and liver BLM in biliary-diverted rats. Am J Physiol 267:G501-7, 1994.
201. +/bile acid cotransporter in extrahepatic cholestasis. Gastroenterology 110:199-209, 1996.
202. Hoshino M, Ohiwa T, Hayakawa T, et al: Effects of dibutyryl cyclic AMP and papaverine on intrahepatocytic bile acid transport. Scand J Gastroenterol 28:833-8, 1993.
203. Takikawa H, Sano N, Yamazaki R, Yamanaka M: Colchicine inhibits lithocholate-3-O-glucuronide-induced cholestasis in rats. J Hepatol 22:88-93, 1995.
204. Ohiwa T, Katagin K, Hoshini M, et al: Tauroursodeoxycholate and tauro-β-muricholate exert cytoprotection by reducing intrahepatocyte taurochenodexycholate content Hepatology 17:470-6, 1993.
205. Accatino L, Hono J, Koenig C, et al: Adaptive changes of hepatic bile salt transport in a model of reversible interruption of the enterohepatic circulation in the rat. J Hepatol 19:95-104, 1993.
206. Accatino L, Hono J, Maldonado M, et al: Adaptive regulation of hepatic bile salt transport. Effect of prolonged bile salt depletion in the rat. J Hepatol 7:215-23, 1988.
207. Crawford JM, Crawford AR: Down-regulation of hepatocyte canalicular bile salt transport by biliary diversion. Gastroenterology 108:A1053, 1995.
208. Gatmaitan ZC, Nies AT, Arias IM: Selectively increased translocation of apical membrane proteins by taurocholate and cAMP. Hepatology 22:322A, 1995.
209. Bouscarel B, Gettys TW, Fromm H, Dubner H: Ursodeoxycholic acid inhibits glucagon-induced cAMP formation in hamster hepatocytes: A role for PKC. Am J Physiol 268:G300-10, 1995.
210. Crawford JM, Gollan JL: Hepatocyte cotransport of taurocholate and bilirubin glucuronides: Role of microtubules. Am J Physiol 255:G121-31, 1988.
211. Moseley RH, Van Dyke RW: Organic cation transport by rat liver lysosomes. Am J Physiol Gastrointest Liver Physiol 268:G480-6, 1995.
212. Van Dyke RW, Faber ED, Meijer DKF: Sequestration of organic cations by acidified hepatic endocytic vesicles and implications for biliary excretion. J Pharmacol Exp Ther 261:1-11, 1992.
213. Berenson MM, El-Mir MY, Zhang L-K: Mechanism of bile acid facilitation of biliary protoporphyrin excretion in rat liver. Am J Physiol 268:G754-63, 1995.
214. Oude Elferink RPJ, Meijer DKF, Kuipers F, et al: Hepatobiliary secretion of organic compounds: Molecular mechanisms of membrane transport. Biochim Biophys Acta Rev Biomembr 1241:215-68, 1995.
215. Dijkstra M, Kuipers F, Smit EP, et al: Biliary secretion of trace elements and minerals in the rat. Effects of bile flow variation and diurnal rhythms. J Hepatol 13:112-9, 1991.
216. Brissot P, Zanninelli G, Guyader D, et al: Biliary excretion of plasma non-transferrin-bound iron in rats: Pathogenetic importance in iron-overload disorders. Am J Physiol 267:G135-42, 1994.
217. Hershko C, Graham G, Bates GW, Rachmilewitz EA: Non-specific serum iron in thalassaemia: An abnormal serum iron fraction of potential toxicity. Br J Haematol 40:255-63, 1978.
218. Wright TL, Lake JR: Mechanisms of transport of nontransferrin-bound iron in basolateral and canalicular rat liver plasma membrane vesicles. Hepatology 12:498-504, 1990.
219. LeSage GD, Kost LJ, Barham SS, LaRusso NF: Biliary excretion of iron from hepatocyte lysosomes in the rat: A major excretory pathway in experimental iron overload. J Clin Invest 77:90-7, 1986.
220. Gross JB, Myers BM, Kos LJ, et al: Biliary copper excretion by hepatocyte lysosomes in the rat: Major excretory pathway in experimental copper overload. J Clin Invest 83:30-9, 1989.
221. Dijkstra M, In't Veld G, Van den Berg GJ, et al: Adenosine triphosphate-dependent copper transport in isolated rat liver plasma membranes. J Clin Invest 95.412-6, 1995.
222. Houwen R, Dijskrra M, Kuipers F, et al: Two pathways for biliary copper excretion in the rat. The role of glutathione. Biochem Pharmacol 39:1039-44, 1990.
223. Verkade HJ, Wolters H, Gerding A, et al: Mechanism of biliary lipid secretion in the rat: A role for bile acid-independent bile flow. Hepatology 17:1074-80, 1993.
224. Berr F, Meier PJ, Stieger B: Evidence for the presence of a phosphatidylcholine translocator in isolated rat liver canalicular plasma membrane vesicles. J Biol Chem 268:3976-9, 1993.
225. Smit JJM, Schinkel AH, Oude Elferink RPJ, et al: Homozygous disruption of the murine mdr2 P-glycoprotein gene leads to a complete absence of phospholipid from bile and to liver disease. Cell 75:451-62, 1993.
226. Ruetz S, Gros P: Phosphatidylcholine translocase: A physiological role for the mdr2 gene. Cell 77:1071-81, 1994.
227. Oude Elferink RPJ, Ottenhoff R, van Wijland M, et al: Regulation of biliary lipid secretion by mdr2 P-glycoprotein in the mouse. J Clin Invest 95:31-8, 1995.
228. Möckel G, Gorti S, Tandon RK, et al: Microscope laser light scattering spectroscopy of vesicles within canaliculi of rat hepatocyte couplets. Am J Physiol 269:G73-84, 1995.
229. Cohen DE, Leighton LS, Carey MC: Bile salt hydrophobicity controls vesicle secretion rates and transformations in native bile. Am J Physiol 263:G386-95, 1992.
230. Sömjen GJ, Merikovsky Y, Lelkes P, Gilat T: Cholesterol-phospholipid vesicles in human bile: An ultrastructural study. Biochim Biophys Acta 879:14-21, 1986.
231. Ulloa N, Garrido J, Nervi F: Ultracentrifugal isolation of vesicular carriers of biliary cholesterol in native human and rat bile. Hepatology 7:235-44, 1987.
232. Wolf DE: Microheterogeneity in biological membranes. Curr Topics Membr 40:143-65, 1994.
233. Bittman R, Kasireddy CR, Mattjus P, Slotte JP: Interaction of cholesterol with sphingomyelin in monolayers and vesicles. Biochemistry 33:11776-81, 1994.
234. Sankaram MB, Thompson TE: Cholesterol-induced fluid-phase immiscibility in membranes. Proc Natl Acad Sci USA 88:8686-90, 1991.
235. Cabrai DJ, Small DM, Lilly HS, Hamilton JA: Transbilayer movement of bile acids in model membranes. Biochemistry 26:1801-4, 1987.
236. Kamp F, Hamilton JA: Movement of fatty acids, fatty acid analogues, and bile acids across phospholipid bilayers. Biochemistry 32:11074-86, 1993.
237. Carey MC: Formation and growth of cholesterol gallstones: The new synthesis. In. Fromm H, Leuschner U (eds): Bile acids - Cholestasis - Gallstones. Falk Symposium No. 84. Dordrecht, Kluwer, 1996, pp 147-175.
238. Almeida PFF, Vaz WLC, Thompson TE: Lateral diffusion and percolation in two-phase, two-component lipid bilayers. Topology of the solid-phase domains in-plane and across the lipid bilayer. Biochemistry 31:7198-210, 1992.
239. Evans E, Yeung A: Hidden dynamics in rapid changes of bilayer shape. Chem Phys Lipids 73:39-56, 1994.
240. Nichols JW: Low concentrations of bile salts increase the rate of spontaneous phospholipid transfer between vesicles. Biochemistry 25:4596-601, 1986.
241. Tall AR, Small DM: Body cholesterol removal: Role of plasma high-density lipoproteins. Adv Lipid Res 17:1-51, 1980.
242. Handa T, Saito H, Miyajima K: Phospholipid monolayers at the triolein-saline interface: Production of microemulsion particles and conversion of monolayers to bilayers. Biochemistry 29:2884-90, 1990.
243. Ostapchuk P, Hearing P, Ganem D: A dramatic shift in the transmembrane topology of a viral envelope glycoprotein accompanies hepatitis B viral morphogenesis. EMBO J 13:1048-57, 1994.
244. Yeagle PL: Cholesterol and the cell membrane. Biochim Biophys Acta 822:267-87, 1985.
245. Ipsen JH, Mouritsen OG, Bloom M: Relationships between lipid membrane area, hydrophobic thickness, and acyl-chain orientational order. The effects of cholesterol. Biophys J 57:405-12, 1990.
246. Walde P, Sunamoto J, O'Connor CJ: The mechanism of liposomal damage by taurocholate. Biochim Biophys Acta 905:30-8, 1987.
247. O'Connor CJ, Wallace RG, Iwamoto K, et al: Bile salt damage of egg phosphatidylcholine liposomes. Biochim Biophys Acta 817:95-102, 1985.
248. Carruthers A, Melchior DL: Studies of the relationship between bilayer water permeability and bilayer physical state. Biochemistry 22:5797-807, 1983.
249. Small DM: The formation of gallstones. Adv Intern Med 16:243-64, 1970.
250. Donovan JM, Carey MC: Separation and quantitation of cholesterol "carriers" in bile. Hepatology 12(Suppl)94S-105S, 1990.
251. Steck TL, Kezdy FJ, Lange Y: An activation-collision mechanism for cholesterol transfer between membranes. J Biol Chem 263:13023-31, 1988.
252. Vlahcevic ZR, Gurley EC, Heuman DM, Hylemon PB: Bile salts in submicellar concentrations promote bidirectional cholesterol transfer (exchange) as a function of their hydrophobicity. J Lipid Res 31:1063-71, 1990.