Fellowships and Awards

ANDRE BENSADOUN Division of Nutritional Sciences Cornell University 321 Savage Hall Ithaca, New York 14853 Tel: 607-255-1927 Fax: 607-255-1033 E-mail: ab55@cornell.edu Education Universite de Toulouse, France Eng. of ENSAT engineering 1956 Universite de Toulouse, France Lic. Sc. biology 1956 Cornell University, Ithaca, NY Ph.D. nutrition 1960 Fellowships and Awards 1981-82 National Research Service Award Senior Fellow 1972-73 National Institutes of Health Special Fellowship 1971 Traveling Fellowship, Cornell University 1958-59 Henry Strong Denison Scholar, Cornell University 1974-03 NIH-NHLBI continuous funding of HL-14990-29 Professional Experience 2006-2011 Graduate School Professor, Cornell University 2006 Professor emeritus, Cornell University 1976-2006 Professor of Nutritional Biochemistry, Division of Nutritional Sciences, Cornell University 1981-82 Sabbatical leave, Research Fellow, Cell Biology, Baylor College of medicine, Houston 1974-1976 Associate Professor of Nutritional Biochemistry, Division of Nutritional Sciences, Cornell University 1972-1973 Sabbatical leave, Visiting Associate Professor of Medicine, Division of metabolic Diseases, University of California San Diego 1967-1974 Associate Professor of Physiology, Poultry Science Department and Joint appointee, Section of Physiology, Division of Biological Sciences, Cornell University 1965-1967 Assistant Professor, Department of Animal Sciences, University of Illinois, Champaign- Urbana 1960-1965 Research associate, Department of Animal Sciences, Cornell University

Bensadoun page 2 Memberships American Society for Biochemistry and Molecular Biology American Association for the Advancement of Science American Institute of Nutrition American Heart Association American Society for Cell Biology American Diabetes Society Teaching Responsibilities Courses taught until 2006 NS 602 (BS 619) Lipids -- graduate course on lipid metabolism. BS 731 -- Lipoprotein Receptors, short course taught periodically Research Interests * Isolation, cloning of heparan sulfate proteoglycans and studies of their functions by tissue specific gene knock out in mice. * Structure/function relationship of lipolytic enzymes. * Identification of major enhancers and silencers in lipoprotein lipase and hepatic lipase gene promoters. * Function of lipolytic enzymes and proteoglycans in lipoprotein receptor mediated endocytosis * Function of GPI anchored HDL-binding protein (GPI-HBP-1) in the hydrolysis of triglycerides-rich lipoproteins My laboratory is concerned with various aspects of the biochemistry of lipolytic enzymes and their functions in the genesis of circulating plasma lipoproteins. During the recent past, the laboratory has purified to homogeneity, lipoprotein lipase (LPL) and hepatic lipase (HL) in tissues of human and experimental animals. Panels of monoclonal antibodies against these two lipases have been isolated and cdna's and genomic sequences for several lipases have been isolated. A major thrust of our efforts is to conduct structural and functional studies of these lipases. Fine mapping of HL and LPL is being conducted by assigning monoclonal antibodies to specific epitopes on proteolytic, cyanogen bromide, chemically generated fragments and to peptides generated in vitro by expression vectors. Characterized monoclonal antibodies are being evaluated for their ability to inhibit functional properties of lipases: Binding of enzyme to lipid interfaces and heparan sulfate; catalytic activity; binding of the activator apocii to LPL and dimer formation from monomeric LPL. In identifying structural domains, where appropriate, chemical modifications, site directed mutagenesis, and reactivity of monoclonal antibodies to specific fusion products generated in vitro by expression vectors are employed. My laboratory has recently (2006) collaborated with the laboratory of Dr Stephen Young at UCLA in identifying the functions of GPI-HBP-1 (GPI anchored HDL-binding protein). The protein was first identified as an HDL-binding protein. We have discovered that it also binds LPL (lipoprotein lipase), VLDL, HDL and chylomicrons. Our current hypothesis is that GPI-HBP-1 functions as a lipolysis platform assembling a complex that would include LPL, substrate (VLDL, chylomicrons) and HDL. During lipolysis, the proximity of HDL would favor the channeling of surface remnants

Bensadoun page 3 (phospholipids, cholesterol, ApoCs ) from VLDL to HDL. In support of this hypothesis GPI-HBP-1 deficient adult mice and humans are grossly hypertriglyceredemic. Membership in Graduate Fields Nutrition Molecular and Integrative Physiology Biochemistry, Molecular and Cell Biology University Committees 1981-83 Research Policy Committee 1970-72 Standing Committee on Anatomy and Physiology, Chairman 1976 Honors Committee, Division of Nutritional Sciences 1976-79 Library Committee 1976-78 Animal Facility Committee 1975-82 Division of Biological Sciences Cell Biology Committee 1978-81 Seminar Committee, Field of Physiology 1978-81 Curriculum Committee, Section of Physiology 1978-81 Division of Nutritional Sciences Controlled Substances Committee 1979-81 Appointments and Tenure Committee 1984-87 Fellowship Board Committee of the Graduate School 1984-present University Radiation Safety Committee 1989-1993 Nutritional Biochemistry Faculty Search Committee 1989-91 Research Committee, Div. of Nutritional Sciences Curriculum Committee, Nutritional Sciences 1996-1998 Appointment and tenure Committee 1995-1998 Animal Welfare Committee 1996-1998 Search committee for Section of Biochemistry (two positions) Consultation with Public Agencies 1980-1981 Member of Committee B for Evaluation of SCOR's (Special Centers of Research) for the Heart, Lung and Blood Institute, NIH 1979-1982 Site visitor for Program Projects administered by the Heart, Lung and Blood Institute 1981-1986 Ad Hoc member of the Metabolism Study Section. National Institutes of Health. 1983-1984 Ad Hoc member of the Metabolism and Physiological Chemistry Study Section, National Institutes of Health 1986-1987 Member of Committee B for evaluation of SCOR's (Specialized Centers of Research on Atherosclerosis) 1991-1992 Tobacco-Related Disease Research Program. State of California: Study Section. April 2-3. 1991-1992 NIH Special Study Section of HLBI: RFA: Plasma Lipoprotein. 1994 NIH reverse site visit in Washington of UCLA lipase Program Project 1995-1997 Member of the American Heart Association Lipoproteins and Lipid Metabolism Research Study Section

Bensadoun page 4 1996 NIH Atherosclerosis subcommittee for Specialized Centers of Research in Molecular Medicine and Atherosclerosis. 1999 NIH study section: CNRU, Clinical Nutrition Research Centers 1999 NIH metabolism study section. 2005 NIH study section: Clinical Nutrition Research Units Ongoing Research Support 199-7418 Bensadoun (PI) 9/30/06-9/30/09 Hatch funds Percent time: 5% Direct costs: $17,000/year Regulation of hepatic lipase expression The major goal of this project is to study the transcriptional regulation of hepatic lipase RO1 HL087228 Young SG (PI) 2/01/07-1/31/12 Subcontract to Bensadoun Percent time: 10% Total costs: $77,000 for 2/01/08-1/31/09 Direct costs: $50,000 for 2/01/08-1/31/09 GPI-HBP1 and metabolism of triglyceride-rich lipoproteins The major goal is to study the function of GPIHBP1 1P01HL090553-01A1 Young SG (PI) 08/01/2008-06/30/2013 Subcontract to Bensadoun Percent time: 20% Total costs:$309,831/year Direct costs:$201,189/year New molecules in triglyceride metabolism and adipogenesis The major goal is to study the structure and molecular mechanism of GPIHBP1 Completed Research Support HL14990-29 Bensadoun (PI) 3/1/75-2/28/04 NIH Functional aspects of the lipoprotein-lipase-proteoglycan interaction The major goal of the project was to characterize heparan sulfate proteoglycans in adipose tissue and identify their function in lipoprotein lipase secretion. Conferences and Invited Talks Deuel Conference on Lipids. Invited talk. Effect of an anti-lipoprotein lipase on triglyceride removal, 1976. Gordon Conference on Lipid Metabolism. Invited talk: Regulation of lipoprotein lipase, 1976.

Bensadoun page 5 University of Paris, Dr. Polonovski's Laboratory. Invited talk: Lipoprotein lipase, 1976. Meetings of American Heart Association. Invited talk: Identification of a lipoprotein lipase in liver perfusate, 1976. Clinical Institute of Montreal. Invited talk: Studies on lipoprotein lipase, 1977. Department of Biochemistry, Medical College of Pennsylvania. Invited talk: Studies on lipoprotein lipase, 1977. Department of Medicine, Albert Einstein Medical College. Invited talk: Studies on lipoprotein lipase, 1977. Massachusetts Institute of Technology. Invited seminar speaker, 1978. Federation meetings. Symposium on lipid transport, 1979. Cooperative Extension, North Island, NY. Dietary lipids and cardiovascular disease, 1979. Department of Biology, Utica College of Syracuse University. Invited talk: Aspects of lipid transport in plasma of vertebrates, 1979. Department of Medicine, University of Washington. Invited talk: Synthesis and secretion of lipoprotein lipase, 1979. European Lipoprotein Club. Lead-off paper on synthesis of lipoprotein lipase, 1980. Louisiana State University Medical Center. Distinguished Lecturer, Trauma Training Program at the LSU Medical Center, 1980. Lipid Metabolism, Gordon Research Conference. Synthesis and secretion of lipoprotein lipase, 1981. Deuel Conference, California. Role of hepatic lipase, 1983. New Aspects in Lipoprotein Metabolism, International Symposium. Effect of camp on the synthesis and secretion of lipoprotein lipase by adipocytes in culture, 1983. Lipid Metabolism, Gordon Research Conference. Regulation of synthesis and secretion of lipoprotein lipase by adipocytes, 1984. Symposium, Lipids in Modern Nutrition, Vevey, Switzerland. Role of lipoprotein lipase and hepatic triglyceride lipase in lipoprotein genesis and catabolism, 1985. Gordon Research Conference, Lipid Metabolism. Molecular biology and structure of lipolytic enzymes, Chairman, 1986. Symposium de Creteil sur Lipids, Lipoproteines et Nutrition. Hepatic Lipase, 1988.

Bensadoun page 6 Baylor College of Medicine, Houston, TX. Regulation of lipoprotein lipase synthesis and degradation in cultured adipocytes. Binding of lipoprotein lipase to membrane heparan sulfate proteoglycans is necessary for degradation, 1988. Sandoz, New Jersey. Invited talk. Lipoprotein lipase synthesis, secretion and degradation. January 21-24, 1989. Biological Research Course (HPLC class) given by Waters. Boston, MA. April 30-May 3, 1989. Military Nutrition Research Committee. National Academy of Sciences. Washington, DC. May 22-23, 1989. Glaxo Company, Raleigh, N.C. Presented seminar. June 1-3, 1989. Second Symposium on the Genetic Engineering of Animals, Ithaca, NY. June 25-28, 1989. Planning Committee, Arteriosclerosis Section. 62nd Scientific Session of American Heart Association. Dallas, TX. July 10-11, 1989. American Society for Cell Biology, 1989. Workshop on Triglyceride Metabolism and Fatty Acid Utilization. Tenth International Symposium on Drugs Affecting Lipid Metabolism, Houston, TX. Cochair of the lipase round table. November 7-11, 1989. American Heart Association meeting, New Orleans, LA. Nov. 11-15, 1989. Committee on Military Nutrition Research, Washington, D.C. Committee Member. Washington, DC. February 6-8, 1990. Special Review Committee on Postprandial Lipoproteins and Atherosclerosis, Bethesda, MD. Washington, DC. March 4-5, 1990. Omega-3 Fatty Acid Meeting, Bethesda, MD. Organized by NIH for Principal Investigators. Washington, DC. March 18-19, 1990. Glycobiology Symposium. Bethesda, MD. April 29-May 1, 1990. Steenbach Symposium, Madison, WI. Invited talk: An "LPL-receptor" on adipocytes regulates LPL degradation. Madison, WI. June 3-5, 1990. Gordon Research Conference, Conference on Lipid Metabolism. Meriden, NH. June 18-22, 1990. Hepatic Cholesterol and Postprandial Lipoproteins Seminar, Aspen, CO. The function of the cell surface lipoprotein lipase receptor in lipase turnover. August, 1990. Committee on Military Nutrition Research, Washington, D.C. Committee Member. Washington, DC. November 27-29, 1990.

Bensadoun page 7 American Society for Cell Biology, San Diego, CA. December 8-13, 1990. Committee on Military Nutrition Research, Washington, D.C. Committee Member. Washington, DC. April 11-12, 1991. Journees Du Gerli: Cholesterol, Oxysterols et Acides Biliaires Conference, Paris, France. May 9-24, 1991. Parke-Davis, Ann Arbor, MI. Invited talk: Function of heparan sulfate proteoglycans in the turnover of lipoprotein lipase. June 14-15, 1991. Scientific Review Session of the Bristol-Myers Squibb/Mead Johnson Nutrition Grants Selection Committee, Washington, DC. Presented talk: "Function of heparan sulfate proteoglycans in the secretion of lipoprotein lipase". October 17, 1991. American Heart Association Scientific Sessions. Anaheim, CA. November 11-14, 1991. Glycomed, Inc., Alameda, CA. Invited talk: "Function of cellular heparan sulfate proteoglycans in the turnover of lipoprotein lipase in adipocytes". November 14-15, 1991. American Society for Cell Biology meeting, Boston, MA. November, 1991. Laboratorie de Biochimie, Orsay, France. Invited talk: "Function of heparan sulfate proteoglycans in the turnover of lipoprotein lipase". December 18, 1991. University of California, San Francisco, CA. Cardiovascular Disease Study Section on Tobacco- Related Disease Research Program. April 1-6, 1992. National Institutes of Health, Washington, DC. Study Section on Diet and Lipoproteins in Humans. June 23-24, 1992. Lipid, Gordon Research Conference, Meriden, NH. Secretion and transport of lipases, interaction with proteoglycans. Discussion leader, presented talk: "Function of cellular heparan sulfate proteoglycans in lipoprotein lipase turnover". June 15, 1992. Ottawa Cardiovascular Center, Ottawa, Canada. Presented talk: "Mechanisms of regulation of lipoprotein lipase". July 5-7, 1992. Medical Research Division, American Cyanamid Company, Pearl River, NY. Invited lecture: "Various aspects of the regulation of lipoprotein lipase". October 16, 1992. American Heart Association Meetings, New Orleans, LA. Chaired session "Lipolytic Enzymes". November 16-19, 1992. American Cyanamid Company, Pearl River, NY. Invited talk "Various aspects of the regulation of lipoprotein lipase", October 16, 1992.

Bensadoun page 8 Hercules, Wilmington, DE. Invited talk "Function of lipoprotein lipase in lipid transport". April 15, 1993. Cleveland Clinic Foundation, Cleveland, OH. Invited talk "Transcriptional and posttranscriptional regulation of lipoprotein lipase", April 16, 1993. American Society for Biochemistry and Molecular Biology and the Division of Biological Chemistry - ACS Joint Meeting, San Diego, CA. May 30-June 3, 1993. The Lipid Triad: Triglyceride, HDL, LDL and Cardiovascular Diseases Meeting, Milano, Italy. Invited talk: "Regulation of LPL synthesis by specific fatty acids". July 5-8, 1993. Bassett Research Institute, Cooperstown, NY. "Fatty acids as modulators of lipoprotein lipase activity", August 5, 1993. National Institutes of Health, Herndon, VA. Consultant on the Program Project Reverse Site Visit, October 14-16, 1993. American Heart Association's 66th Scientific Sessions, Atlanta, GA. Chaired session, on "Lipases". November 8-11, 1993. Keystone Symposia on Molecular and Cellular Biology, Park City, UT. January 14-21, 1994. The Creteil Symposium on "Nutrition, Lipides, Lipoproteines", Cedex, France. Invited talk "Molecular aspects of the regulation of lipoprotein lipase". March 25, 1994. FASEB Summer Research Conference, Santa Cruz, CA. July 1-10, 1994. American Heart Association Meetings, Dallas, TX. Nov. 12-16, 1994. Pennington Biomedical Research Center, Louisiana State University, Baton Rouge, Invited talk, Transcriptional and Post-transcriptional sites of Lipoprotein Lipase Regulation, May 18,1995. Columbia University, Ateriosclerosis SCOR seminar, Recent studies on lipoprotein lipase: Identification and significance of the heparin binding domains, April 6, 1998. Laval University, Ste-Foy, Laval University Medical Center, " Post-translational regulation of lipoprotein lipase" Spring 2000. Pennington biomedical research center, Baton-Rouge, LA, " Role of syndecan-4 in adipocyte cell proliferation. April 11, 2002. Columbia University, Ateriosclerosis SCOR seminar, Interaction of lipoprotein lipase with RAP, Spring 2004 Abstracts Accepted for Presentations at National Meetings Since 1976 Bensadoun, A., Koh, T.L. and Yang, M.W.W. Identification of an adipose-like lipoprotein lipase in liver perfusates. American Heart Association. Accepted for presentation at 1976 scientific meetings.

Bensadoun page 9 Brannon, P.M., Cheung, A.H. and Bensadoun, A. Synthesis of lipoprotein lipase in culture avian granulosa cells. Fed. Proc. 37: 348, 1978. Bensadoun, A., Cheung, A. and Chow, M.C. Direct solid phase radioimmunoassay of chicken lipoprotein lipase. Accepted for presentation at 5th International Symposium on Atherosclerosis, 1979. Jensen, G. and Bensadoun, A. Synthesis of lipolytic enzymes by cultured hepatocytes. American Heart Association. Accepted for presentation at November 12, 1979 scientific sessions. Baly, D.L. and Bensadoun, A. Effect of insulin on the clearance of portomicron remnants by primary cultures of avian hepatocytes. Accepted for presentation at FASEB meeting, 1980. Behr, S., Patsch, J., Forte, T. and Bensadoun, A. Plasma lipoprotein changes resulting from blocked lipolysis. American Heart Association. Accepted for presentation at November 17-20, 1980 scientific sessions. Cheng, C-F., Bensadoun, A. and Rosenberg, R. Binding of lipoprotein lipase to cultured endothelial cells. American Heart Association. Accepted for presentation at November 17-20, 1980 scientific sessions. Jensen, G.L. and Bensadoun, A. True alkaline triglyceride lipase activities in cultured hepatocytes by specific chlorpromazine inhibition of lysosomal lipase. American Heart Association. Accepted for presentation at November 17-20, 1980 scientific sessions. Cheng, C-F., Bersot, T. and Bensadoun, A. Monoclonal antibodies specific for human hepatic triglyceride lipase. Accepted for presentation at the 1984 American Heart Association Meeting, 1984. Bensadoun, A. and Marita, R. Dibutyryl cyclic AMP decreases adipocytes LPL synthesis and secretion rates. Fed. Proc. 44: 931, 1985. Daggy, B. and Bensadoun, A. Characterization of hyperlipoproteinemia following in vivo inhibition of rat hepatic triglyceride lipase. Fed. Proc. 44: 1763, 1985. Cornely-Moss, K.A. and Bensadoun, A. Lipolysis of triglyceride rich lipoproteins with hepatic lipase enhances the accessibility of apoe epitopes. J. Cell Biol. 107(6) Part 3, 1988. Sensel, M., Bensadoun, A. and Melford, K. Regulation of rat hepatic lipase. J. Cell Biol. 107(6) Part 3, 1988. Bensadoun, A. and Weinfeld, J. An LPL receptor on adipocytes regulates LPL degradation. Twentieth Steenbock Symposium "Molecular Biology of Atherosclerosis", Madison, Wisconsin, June 1990. Arteriosclerosis, pp. 71-77, 1990. Hoogewerf, A. and Bensadoun, A. Avian adipose lipoprotein lipase is a sulfonated glycoprotein. American Society for Biochemistry and Molecular Biology, New Orleans, LA, June 1990. FASEB Journal, #1385. April 26, 1990 Vol. 4, Number 7, page A1931.

Bensadoun page 10 Montalto, M. and Bensadoun, A. Lipoprotein lipase synthesis and secretion: Effects of concentration and type of fatty acids in adipocyte cell culture. American Society for Cell Biology Meeting, San Diego, CA, December 1990. Kim, K.C. and Bensadoun, A. Isolation of Lipoprotein Lipase-Binding Heparan Sulfate Proteoglycan from the Cultured Avian Adipocytes. American Society for Cell Biology Meeting, Boston, MA, December 8-12, 1991. The Journal of Cell Biology Vol. 115, page 471a, 1991. Berryman, D. and Bensadoun, A.. Characterization of a Heparin-Binding Site in Lipoprotein Lipase. 32nd Annual American Society for Cell Biology Meeting, Denver, CO, November 15-19, Vol. 3, page 122a, 1992. Lauer, S.J., Ji, Z-S., Fazio, S., Bensadoun, A., Taylor, J.M. and Mahley, R.W. Human hepatic lipase secretion by transfected McA-RH7777 cells results in enhanced binding and uptake of remnant lipoproteins. American Heart Association Meeting, Atlanta, GA, November 6-9, 1993. Fan, J., Wang, J., Lauer, S.J., Dang, Q., Sanan, D.A., Bensadoun, A., Mahley, R.W., and Taylor, J.M. Correction of hepatic lipase deficiency in transgenic rabbits results in a dramatic reduction of high density lipoprotein levels in plasma. Clinical Research Meeting, Baltimore, MD, April 29-May 2, 1994. Klaus A.Dugi, Catherine L.Knapper, Deborah Applebaum-Bowden, Andre Bensadoun, Susan M. Meyn, Thong T.Le, Nobuyo Maeda, H.Bryan Jr. Brewer, Silvia Santamarina-Fojo. American Heart Association, Atlanta, New Orleans,LA, November 10-13,1996. Haudenschild,C.C., Dugi, K.A., Amar, M., Knapper, C.L., Bensadoun, A., Maeda, N., Brewer, B., and Santamarina-Fojo, S. Identification of a residue (433K) essential for hepatic lipase mediated lipoprotein metabolism independent of lipolysis. American Heart Association Meetings; Nov. 10/1997. Lambert, G.C., Remaley, A.T., Chase, M.B., Peterson, K.M., Bensadoun, A., Brewer, B., and Santamarina-Fojo, S. Role of hepatic lipase in the Cla-1 mediated selective uptake of HDL cholesterol esters. American Heart Association Meetings; Nov. 9/1998. Merkel, M., Kako, Y., Melford, K., Breslow, J.L., Bensadoun, A., and Goldberg, I.J. Transgenic mice expressing lipoprotein lipase(lpl) with a mutation in the carboxyl-terminal heparin binding region have large amounts of inactive LPL in preheparin plasma. American Heart Association Meetings; Nov 7-10, 1999. Van Vlijmen, B.,J., Rohlmann, A., Havekes, L., M., Van Berkel, T. J., Bensadoun, A., Herz, J. An extrahepatic LDL receptor and LRP-independent RAP-sensitive site is involved in the metabolism of chylomicron remnants. American Heart Association Meetings; Nov 7-10, 1999. Yagyu, H., Lutz,P., E., Kako,Y., Marks, S., Hu,Y., Bensadoun, A., and Goldberg, I.J. VLDL receptor(vldlr) deficiency leads to hypertriglyceridemia and abnormal regulation of lipoprotein lipase in lipoprotein lipase heterozygous knockout (LPL1) mice. Accepted for presentation, American Heart Association meetings, Nov 11-14, 2001.

Bensadoun page 11 Lutz, E.,P., Kako, Y., Marks, S., Melford, K., Bensadoun, A., Goldberg, I. Covalent dimeric heparinbinding defective lipoprotein lipase has increased enzymatic stability and prevents neonatal demise of lipoprotein lipase mice. Accepted for presentation, American Heart Association meetings, Nov 11-14, 2001. Conde,K., Bensadoun, A. and Shachter, N.S. Severe hypertriglyceridemia and remnant accumulation due to modest overexpression of human apolipoprotein (apo) C-1 in apoe-null mice: ApoC1-induced hyperlipidemia appears not to be mediated via effects on apo-e mediated clearance. American Heart Association Meetings. Nov 2002. Wernimont, S. and Bensadoun, A. Effect of heparan sulfate proteoglycans (HSPGs) on clpl turnover. American Society for Nutritional Sciences. April 2003. Mayumi Takahashi, Masayoshi Yokoyama, Toru Seo, André Bensadoun, Ira J. Goldberg; Endothelium specific overexpression of human Lipoprotein Lipase (LpL) leads to fatty liver in mice. Ateriosclerosis thrombosis Vascular Biology meeting. 2006 Chahil, T., Ngai, C., Bensadoun, A., Goldberg, I., Holleran, S., Ramakrishnan, R., Ginsburg, H. Effects of pioglitazone and rosiglitazone on lipid metabolism. American Diabetes Association, 67 Scientific Sessions, 2007. Franssen, R. Beigneux, A.P., Hertecant, J., Gin, P., Weinstein, M., Kastelein, J.P., Bensadoun, A., Fong, L.G., Young, S.G., Dallinga-Thie, G.M. Two mutations in GPIHBP1 cause severe chylomicronemia and provide evidence for a critical role of GPIHBP1 in human triglyceride metabolism. International Symposium on Atherosclerosis, 2009. Dallinga-Thie, G.M., Beigneux, A.P., Franssen, R., Hertecant, J., Gin, P., Weinstein, M., Kastelein, J.J.P., Bensadoun, A., Fong, L.G., Young, S.G. Two mutations in GPIHBP1 cause severe chylomicronemia and provide evidence for a critical role of GPIHBP1 in human triglyceride metabolism. Ateriosclerosis thrombosis Vascular Biology meeting, 2009. Theses Completed in Dr. Bensadoun's Laboratory at Cornell University) Chandrabose, K.A. Effect of hypophysectomy on some aspects of lipid metabolism in cockerels; Ph.D., 1970. Sadovsky, R.A. The nature of the thyroid iodohormones in the plasma of the rooster (Gallus domesticus) and their diurnal rhythm; Honors, 1970. Rothfeld, A. The absorption of micellar oleic acid from the intestine of the domestic fowl, Gallus domesticus; Honors, 1971. Clifford, C. Mechanism of plasma lipoprotein uptake by the avian follicles; Ph.D., 1971. Boyd, T.A. Aspects of lipoprotein lipase in the domestic fowl (Gallus domesticus); Honors, 1973.

Bensadoun page 12 Strauss, M.L. A study of fatty acid transport in the male chicken, Gallus domesticus; M.A., 1973. Bramer, M. Studies on serotonin levels in the pineal of the immature albino rat and the immature Japanese quail; Ph.D., 1973. Lomasky, S.J. Effect of triglyceride unsaturation on the catalytic activity of lipoprotein lipase; Honors, 1977. Spellun, J.A. Isolation and purification of the activator of adipose tissue lipoprotein lipase, apolipoprotein CII from hen's egg very low density lipoproteins. Cheung, A.H. Studies on avian lipoprotein lipase. I. Radioimmunoassay. II. Electron microscopic localization; Master's thesis, 1974. Baly, D.L. The role of insulin in the clearance of portomicron remnants by primary cultures of avian hepatocytes; Master's thesis, Cornell University, 1979. Brannon, P.M. Synthesis, turnover, and secretion of lipoprotein lipase in granulosa cells; Ph.D. thesis, Cornell University, 1979. Selling, B. Intermolecular and intramolecular specificity of lipoprotein lipase; Honors thesis, Biochemistry, Cornell University, 1979. Jensen, G. Hepatic triacylglycerol lipases. 1. Identification and characterization of lipolytic activities in liver perfusates and cultured hepatocytes. 2. Purification, stabilization and characterization of hepatic triglyceride lipase. Ph.D. thesis, Cornell University, January 1981. Behr, S.R. Lipoprotein metabolism in normolipemic and hypercholesterolemic roosters. Ph.D. thesis, Cornell University, August 1983. Cisar, L.A. An enzyme linked immunosorbent assay for rat hepatic triglyceride lipase. Master's thesis, Cornell University, August 1983. Daggy, B.P. Characterization of the hyperlipoproteinemia following in vivo inhibition of hepatic lipase in fed and fasting rats. Ph.D. thesis, Cornell University, August 1985. Raport, Carol J. Role of glycosylation in the synthesis and secretion of lipoprotein lipase in cultured chicken adipocytes. Master's thesis, Cornell University, January 1986. Cisar, L. Studies on the synthesis and secretion of hepatic lipase in Fu5AH rat hepatoma cells and primary cultures of rat hepatocytes. Ph.D. thesis, Cornell University, August 1986. Cornely-Moss, K. Immunological studies of triglyceride-rich lipoproteins from control and hepatic lipase-inhibited animals. Ph.D. thesis, Cornell University, August 1989. Sensel, M. Regulation of hepatic lipase: Isolation of cdna and genomic clones and in vivo effects of thyroid hormone and hypercholesterolemia. Ph.D. thesis, Cornell University, August 1989.

Bensadoun page 13 Hoogewerf, A. Sulfation of lipoprotein lipase and heparan sulfate proteoglycans in cultured chicken adipocytes, Ph.D. thesis, Cornell University, January 1991. Chen, David. Howard Hughes Fellow. Effect of bivalent and monovalent antibodies on the release of lipoprotein lipase in cultured avian adipocytes. Honors thesis, Cornell University, May 1991. Yamada, Reiko. Regulation of rat hepatic lipase by cholesterol, M.S. thesis, Cornell University, August 1991. Montalto, Michael. Lipoprotein synthesis and secretion: Effects of concentration and type of fatty acids in adipocyte cell culture. Ph.D. thesis, Cornell University, May 1992. Lu, Shao-Chun. Structure, sequence and transcriptional regulation of the avian lipoprotein lipase gene. Ph.D. thesis, Cornell University, January 1993. Berryman, Darlene. Interaction of avian lipoprotein lipase with heparan sulfate proteoglycans. Ph.D. thesis, Cornell University, May 1994. Wang, Ming-Eng. Transcriptional regulation of the rat hepatic lipase gene: Dissection of the 5' flanking region and characterization of two elements with similar sequences but distinct function. Ph.D. thesis, Cornell University, August 1994. A. Alex Hoffling. Characterization of the receptor-associated protein, a modulator of the low density lipoprotein receptor family. Honors thesis, May 1996. Soo Yeon Cho. Molecular cloning and characterization of C-K-glypican, glycosyl phosphatidyl inositol anchored heparan sulfate proteoglycan(gpi_hspg) with affinity for lipoprotein lipase in avian adipocytes. PhD thesis, Cornell University, Jan 1997. Yong Liu. Purification and cloning of a GPI-anchored heparan sulfate proteoglycan from the human liver carcinoma cell HepG2 cells. MS thesis, Cornell University, Aug 1997. Riggs Christine. The effect of fatty acids and insulin on lipoprotein lipase synthesis in adipocyte cell culture. PhD thesis, Cornell University, Aug 1997. Zhang Wei. Characterization of the cis-regulatory elements in the chicken lipoprotein lipase(lpl) gene and proteins binding to these elements, PhD thesis, Cornell University, Dec 1997. Rebecca Sendak. Structure/function studies of avian lipoprotein lipase: Identification of the primary heparin-binding domain, PhD thesis, Cornell University, May 1998. Reiko Yamada Landry. Characterization of syndecan-4 induction in mouse 3T3-F442A adipocytes. PhD thesis, Cornell University, January 2000. Shallee Page: High affinity binding between lipoprotein lipase and low density lipoprotein receptorrelated protein receptor-associated protein. PhD thesis, Cornell University, May 2000.

Bensadoun page 14 Guanguy Zhang: Glypican-4: Expression during pre-adipocyte differentiation. MS thesis, May 2001. Krysta D. Levac: Regulation of leptin expression by polyunsaturated fatty acids in 3T3-F442A adipocytes. PhD thesis, Cornell University, May 2001. Susan Wernimont: Role of the heparin sulfate proteoglycans (HSPG) syndecan-4 in regulating secretion, degradation, and transcription of lipoprotein lipase (LPL). MS thesis. 2004. Anne LeBourg: Sterol regulation of hepatic lipase. A novel role for sterol regulatory element binding protein in cholesterol homeostasis. PhD thesis, 2006. Past Doctoral Fellows: Veronica Hearn, 1971-74 John Benson, 1972-75 Putu Kompiang, 1974-76 Steve Behr, 1980-81 Chao Fu Cheng, 1978-85 Rosalind Marita, 1984-86 Laura Cisar, 1986-89 Philippe Legrand, 1989-90 Anne legrand, 1989-90 Paul Strielman, 1986-87 Mary Cupp, 1985-1987 Dawn Bresaemle, 1990-93 Dale Cooper, 1989-92 Kyung Kim, 1991-92 Karin Misra, 1992-94 Darlene Berryman, 1992-94 Julio Mulero, 1995-97 Carroll Brooks, 1999-2001 Vincent Rioux, 2001-02 Huaixing Li, 1999-2002 Publications (since 1974) Bensadoun, A., Ehnholm, C., Steinberg, D. and Brown, W.V. Purification and characterization of lipoprotein lipase from pig adipose tissue. J. Biol. Chem. 249: 2220, 1974. Clifford, C.K. and Bensadoun, A. In vivo uptake of triglyceride fatty acids and free fatty acids by ovarian follicles of hens (Gallus domesticus). Comp. Biochem. Physiol. 51: 133, 1975. Benson, J.D., Bensadoun, A. and Cohen, D. Lipoprotein lipase of ovarian follicles in the domestic chicken (Gallus domesticus). Proc. Soc. Exp. Biol. Med. 148: 347, 1975. Hearn, V. and Bensadoun, A. Plasma lipoproteins of the chicken, Gallus domesticus. Int. J. Biochem. 6: 295, 1975.

Bensadoun page 15 Benson, J.D., Hearn, V., Boyd, T. and Bensadoun, A. Triglyceride hydrolase of chicken and rat preand post-heparin plasma. Effects of fasting and comparison with adipose lipoprotein lipase. Int. J. Biochem. 6: 727, 1975. Bensadoun, A., Weinstein, D. and Steinberg, D. Assay of proteins in the presence of interfering material. Anal. Biochem. 70: 241, 1976. Shemesh, M., Bensadoun, A. and Hansel, W. Lipoprotein lipase activity in the bovine corpus leuteum during the estrous cycle and early pregnancy. Proc. Soc. Exp. Biol. Med. 151: 667, 1976. Kompiang, I.P., Bensadoun, A. and Yang, M.W.W. Effect of anti-lipoprotein lipase serum on plasma triglyceride removal. J. Lipid Res. 17: 498, 1976. Bensadoun, A., Koh, T.L., and Yang, M.W.W. Identification of an adipose-like lipoprotein lipase in liver perfusates. Circulation 54: 85, 1976. Benson, J.D. and Bensadoun, A. Response of adipose tissue lipoprotein lipase to fasting in the chicken and the rat -- A species difference. J. Nutr. 107: 990 1977. Ehnholm, C., Bensadoun, A., and Brown, W.V. Characterization of two triacylglycerol lipase activities in pig post heparin plasma. Biochem. J. 163: 347, 1977. Bensadoun, A. and Koh, T.L. Identification of an adipose tissue-like lipoprotein lipase in perfusates of chicken liver. J. Lipid Res. 18: 768, 1977. Brannon, P.M., Cheung, A.H., and Bensadoun, A. Synthesis of lipoprotein lipase in cultured avian granulosa cells. Biochim. Biophys. Acta 531: 96, 1978. Cheung, A.H., Bensadoun, A., and Cheng, C.-F. Direct solid phase radioimmunoassay for chicken lipoprotein lipase. Anal. Biochem. 94: 346, 1979. Bensadoun, A. and Kompiang, I.P. The role of lipoprotein lipase in plasma triglyceride removal. Fed. Proc. 38: 2622, 1979. Brannon, P.M. and Bensadoun, A. Determination of the half-life of lipoprotein lipase in cultured granulosa cells. Biochim. Biophys. Acta 618: 173, 1980. Jensen, G.L., Baly, D.L., Brannon, P.M., and Bensadoun, A. Synthesis and secretion of lipolytic enzymes by cultured chicken hepatocytes. J. Biol. Chem. 255: 11141, 1980. Behr, S.R., Patsch, J.R., Forte, T., and Bensadoun, A. Plasma lipoprotein changes resulting from immunologically blocked lipolysis. J. Lipid Res. 22: 443, 1981. Jensen, G.L. and Bensadoun, A. Purification, stabilization and characterization of rat hepatic triglyceride lipase. Anal. Biochem. 113: 246, 1981. Cheng, C.-F., Oosta, G.M., Bensadoun, A., and Rosenberg, R.D. Binding of lipoprotein lipase to endothelial cells in culture. J. Biol. Chem. 256: 12893, 1981.

Bensadoun page 16 Jensen, G.L., Daggy, B., and Bensadoun, A. Triglyceride lipase, monoglyceride lipase and phospholipase activities of highly-purified rat hepatic lipase. Biochim. Biophys. Acta 710: 464, 1982. Bensadoun, A. and Behr, S.R. Use of specific antibodies in the study of the Function of Lipoprotein Lipase. Atherosclerosis VI, 1982. Cisar, L. and Bensadoun, A. Enzyme linked immunosorbent assay for rat hepatic triglyceride lipase. J. Lipid Res. 26: 380-386, 1985. Cheng, C.-F., Bensadoun, A., Bersot, T., Hsu, J.S.T., and Melford, K.H. Purification and characterization of human lipoprotein lipase and hepatic triglyceride lipase. Reactivity with monoclonal antibodies to hepatic triglyceride lipase. J. Biol. Chem. 260: 10720-10727, 1985. Gershenwald, J.E., Bensadoun, A., and Saluja, A. Monoclonal antibodies to avian lipoprotein lipase. Purification of the enzyme by immunoaffinity chromatography. Biochim. Biophys. Acta 836: 286-295, 1985. Daggy, B.P. and Bensadoun, A. Enrichment of apolipoprotein B-48 in the LDL density class following in vivo inhibition of hepatic lipase. Biochim. Biophys. Acta 877: 252-261, 1986. Bensadoun, A. and Marita, R.A. Dibutyryl cyclic AMP decreases the rate of lipoprotein lipase synthesis in cultured adipocytes. Biochim. Biophys. Acta 879: 253-263, 1986. Bensadoun, A. and Daggy, B.P. Role of lipoprotein lipase and hepatic lipase in the genesis of plasma lipoproteins. In: Lipids in Modern Nutrition 13. Raven Press, 17-27, 1987. Cisar, L. and Bensadoun, A. Characterization of the intracellular processing and secretion of hepatic lipase in Fu5AH rat hepatoma cells. Biochim. Biophys. Acta 927: 305-314, 1987. Strieleman, P.J. and Bensadoun, A. Cell free translation of avian adipose tissue lipoprotein lipase messenger RNA. Biochim. Biophys. Acta 908: 268-274, 1987. Cupp, M., Melford, K. and Bensadoun, A. Heparin decreases the degradation rate of lipoprotein lipase in adipocytes. J. Biol. Chem. 262: 6383-6388, 1987. Daggy, B., Arost, C. and Bensadoun, A. Dietary fish oil decreases VLDL production rates. Biochim. Biophys. Acta 920: 293-300, 1987. Bensadoun, A., Daggy, B.P. and Cornely-Moss, K. Action of hepatic lipase on VLDL and chylomicron remnant removal. Ann. Nutr. Metab. 32: 247, 1988. Cisar, L.A., Hoogewerf, A.J., Cupp, M., Rapport, C.A. and Bensadoun, A. Secretion and degradation of lipoprotein lipase in cultured adipocytes. Binding of lipoprotein lipase to membrane heparan sulfate proteoglycans is necessary for degradation. J. Biol. Chem. 264: 1767-1774, 1989.

Bensadoun page 17 Cooper, D.A., Stein, J.C., Strieleman, P.J. and Bensadoun, A. Avian adipose tissue lipoprotein lipase: cdna sequence and reciprocal regulation of mrna levels in adipose and heart. Biochim. Biophys. Acta. 1008: 92-101, 1989. Cisar, L.A., Melford, K.H., Sensel, M. and Bensadoun, A. Heparin decreases the degradation rate of hepatic lipase in Fu5AH rat hepatoma cells. A model for hepatic lipase efflux from hepatocytes. Biochim. Biophys. Acta. 1004: 196-204, 1989. Sensel, M.G., Legrand-Lorans, A., Wang, M-E. and Bensadoun, A. Isolation and characterization of clones for the rat hepatic lipase gene upstream regulatory region. Biochim. Biophys. Acta. 1048: 297-302, 1990. Bensadoun, A., Cisar, L.A., Hoogewerf, A.J. and Cupp, M. An "LPL-receptor" on adipocytes regulates LPL degradation. In: Molecular Biology of Atherosclerosis (Attie, A., ed). Elsevier: New York, pp. 71-77, 1990. Hoogewerf, A. and Bensadoun, A. Occurrence of sulfate in the asparagine-linked complex oligosaccharide of lipoprotein lipase. J. Biol. Chem. 266: 1048-1057, 1991. Bensadoun, A. Lipoprotein Lipase. Ann. Rev. Nutr. 11: 217-237, 1991. Hoogewerf, A.J., Cisar, L.A., Evans, D.C. and Bensadoun, A. Effect of chlorate on the sulfation of lipoprotein lipase and heparan sulfate proteoglycans: Sulfation of heparan sulfate proteoglycans affects lipoprotein lipase degradation. J. Biol. Chem. 266: 16564-16571, 1991. Legrand, P. and Bensadoun, A. Stearyl CoA desaturase activity in cultured rat hepatocytes. Biochim. Biophys. Acta 1086: 89-94, 1991. Cooper, D., Lu, S-C., Viswanath, R., Freiman, R. and Bensadoun, A. Structure and sequence of the avian lipoprotein lipase gene. Biochim. Biophys. Acta. 1129: 166-171, 1992. Montalto, M. and Bensadoun, A. Lipoprotein lipase synthesis and secretion: Effects of concentration and type of fatty acids in adipocyte cell culture. J. Lipid Res. 34: 397-407, 1993. Brasaemle, D., Cornely-Moss, K. and Bensadoun, A. Hepatic lipase treatment of chylomicron remnants increases exposure of apolipoprotein E. J. Lipid Res. 34: 455-465, 1993. Berryman, D. and Bensadoun, A.. Site-directed mutagenesis of a putative heparin binding domain in avian lipoprotein lipase. J. Biol. Chem. 268: 3272-3276, 1993. Lu, S-L. and Bensadoun, A. Identification of 5' regulatory elements of avian lipoprotein lipase (LPL) gene: Synergistic effects of multiple factors. Biochim. Biophys. Acta. 1216: 375-384, 1993. Shafi, S., Brady, S.E., Bensadoun, A. and Havel, R.J. Role of hepatic lipase in the uptake and processing of chylomicron remnants in rat liver. J. Lipid Res. 35: 709-720, 1994.

Bensadoun page 18 Misra, K., Kim, K.C., Cho, S-Y., Low, M.G. and Bensadoun, A. Purification and characterization of heparan sulfate proteoglycans with affinity for lipoprotein lipase in adipocytes. J. Biol. Chem. 269: 23838-23844, 1994. Fan, J., Wang, J., Bensadoun, A., Lauer, S., Qi, D., Mahley, R.W. and Taylor, J.M. Overexpression of hepatic lipase in transgenic rabbits leads to a marked reduction of plasma high density lipoproteins and intermediate density lipoproteins. Proc. Nat. Acad. Sci. USA. 91:8724-8728, 1994. Ji, Z.S., Lauer, S.J., Fazio, S., Bensadoun, A., Taylor, J.M., and Mahley, R.W. Enhanced binding and uptake of remnant lipoproteins by hepatic lipase-secreting hepatoma cells in culture. J. Biol. Chem. 269:13429-13436. 1994. Berryman,D.E. and Bensadoun,A. Heparan sulfate proteoglycans are primarily responsible for the maintenance of enzyme activity, binding, and degradation of lipoprotein lipase in Chinese hamster ovary cells. J. Biol. Chem. 270:24525-24531,1995. Havel, R.J., Herz, J., Shafi, S., Mokuno, H., Bensadoun, A., Brady, S., Kotite, L., and desilva, H. 1995. Mechanism of hepatic uptake and endocytosis of chylomicron remnants. Atherosclerosis X. 168-170. Bensadoun, A. Sandwich-immunoassay for the measurement of human hepatic lipase. Methods in Enzymol. 263:333-338, 1996. Knudsen,P., Antikainen M., Ehnholm S., Uusioukari, M., Tenkanen, H., Lahdenpera, S., Kahri, J., Tilly-Kiesi, M., Bensadoun A., Taskinen, M.R., and Ehnhlom, C. A compound heteozygote for hepatic lipase gene mutations Leu334 Phe and Thr383 Met: correlation between HL activity and phenotypic expression. J. Lipid Res. 37:825-834, 1996. Bensadoun, A. and Berryman,D.E. Genetics and Molecular Biology of hepatic lipase. Curr. Opin. Lipidology, 7:77-81, 1996. Sanan, D.A., Fan, J., Bensadoun, A., and Taylor, J.M. Hepatic lipase is abundant on both hepatocyte and endothelial cell surfaces in the liver, J. Lipid Research, 38: 1002-1013, 1997. Knudsen, P., Antikainen, M., Uusi-Oukari, M., Ehnholm S., Lahdenpera, S., Bensadoun A., Funke, H., Wiebusch, H., Assmann, G., Taskinen, M.R., EhnHolm, C. Heterozygous hepatic lipase deficiency, due to two misense mutations R186H and L334F, in the HL gene. Atherosclerosis,128: 165-174, 1997. Haudenschild, C.C, Dugi, K.A., Amar, M.J., Knapper, C.L., Bensadoun, A., Maeda, N., et al. Identification of a residue (433K) essential for hepatic lipase mediated remnant lipoprotein metabolism independent of lipolysis. Circulation, 96: 1-38, 1997. Berryman, D.E., Mulero, J.J., Hughes, L.B., Brasaemle, D.L. and Bensadoun, A. Oligomeric structure of hepatic lipase: Evidence from a novel epitope tag technique. Biochim. Biophys. Acta, 1382: 217-229, 1998. Sendak, R.A., Melford, K., Kao, A., and Bensadoun, A. Identification of the epitope of a monoclonal antibody that inhibits heparin binding of lipoprotein lipase: new evidence for a carboxyl-terminal heparin-binding domain. J. of Lipid Res.39: 633-646, 1998.

Bensadoun page 19 Sendak, R.A., and Bensadoun, A. Identification of a heparin-binding domain in the distal carboxylterminal region of lipoprotein lipase by site directed mutagenesis. J. Lipid Res. 39: 1310-1315, 1998. Qiu, S., Bergeron, N., Kotite, L., Krauss, R.M., Bensadoun,A. and Havel, R. Concentration, composition and metabolism of lipoproteins containing apolipoprotein B in hepatic lipase-deficient mice. J. Lipid Res. 39: 1661-1668, 1998 Bensadoun,A., Hsu, J. and Hughes, B. Large scale lipoprotein lipase purification from adipose tissue. Methods in Molecular Biology, 109: 145-150, 1998. Bensadoun, A., Hughes, L.B., Melford, K., Hsu, J. and Braesaemle, L. Purification of rat hepatic lipase essentially free of apolipoprotein E and apolipoprotein B. Methods in Molecular Biology, 109 : 151-156, 1998. Bergeron, N., Kotite, L., Verges, M., Blanche, P., Hamilton, R.L., Krauss, R.M., Bensadoun, A. and Havel, R. Lamellar lipoproteins uniquely contribute to hyperlipidemia in mice doubly deficient in apolipoprotein E and hepatic lipase. Proc. Nat.Acad.Sci., USA, 95: 15647-15652, 1998. Zhang, W. and Bensadoun, A. Identification of a silencing element in the chicken lipoprotein lipase gene promoter: Characterization of the silencer-binding protein and delineation of its target nucleotide sequence. Biochim. Biophys. Acta, 1436 : 390-404. 1999. Lambert, G., Chase, M.B., Dugi, K., Bensadoun, A., Brewer, H.B., and Santamarina-Fojo, S. Hepatic lipase promotes the selective uptake of high density lipoprotein-cholesteryl esters via the scavenger receptor B1. J. Lipid Res. 40:1294-1303. 1999. Vlijmen, B.J.M., Rohlmann, A., Bensadoun, A., Page, S., van Berkel T. J.C., Havekes, L.M. and Herz, J. An extrahepatic LDL receptor and LRP-independent RAP-Sensitive site is involved in the metabolism of chylomicron remnants. J. Biol. Chem. 274:35219-35226. 1999. Sendak, R.A., Berryman, D.E., Gellman, G., Melford, K., and Bensadoun, A. Binding of hepatic lipase to heparin: Identification of specific heparin-binding residues in two distinct positive charge clusters. J. Lipid Res. 41:260-268, 2000. Dugi, K.A., Amar, M.J.A., Haudenschild, C.C., Shamburek, R.D., Bensadoun, A., Hoyt, Jr, R.F., Fruchart-Najib, J., Madj,Z., Brewer, Jr, H.B., and Santamarina-Fojo, S. In vivo evidence for both lipolytic and nonlipolytic function of hepatic lipase in the metabolism of HDL. Arterioscler. Thromb. Vas. Biol. 20:793-800. 2000. Zambon, A., Deeb, S.S., Bensadoun, A., Foster, K.E., and Brunzell, J.D. Evidence for a role of hepatic lipase in human apo B lipoprotein metabolism, independent of its lipolytic activity. J. Lipid Res. 41: 2094-2099. 2000. Concannon, P., Levac, K., Rawson, R., Tennant, B., and Bensadoun, A. Seasonal changes in serum leptin, food intake and body weight in photoentrained woodchucks. Endocrinogy.Am. J. Physiol. 281:R951-R959. 2001.

Bensadoun page 20 Lutz, E.P., Merkel, M., Kako, K., Melford K., Radner, H., Breslow, J.L., Bensadoun, A., and Goldberg, I.J. Heparin-binding defective lipoprotein lipase (LPL): evidence that the carboxyl terminal region affects LPL binding to cells, its clearance, and physiological regulation. J. Clin. Invest. 107: 1183-1192. 2001. Landry R., Rioux, V., and Bensadoun, A. Characterization of syndecan-4 expression in 3T3-F442-A mouse adipocytes: Link between syndecan-4 induction and cell proliferation. Cell Growth and Diff.12: 497-504. 2001. Rioux, V., Landry, R.Y., and Bensadoun, A. Sandwich immunoassay for the measurement of murine syndecan-4. J. Lipid Res. 43: 167-173. 2002. Yagyu,H., Lutz, E.P., Kako,Y., Marks,S., Hu, Y., Choi,S., Bensadoun,A., and Goldberg,I.J. VLDL receptor deficient mice have hypertriglyceridemia and abnormal regulation of lipoprotein lipase. J. Biol. Chem. 277: 10037-10043. 2002. Gonzalez-Navarro, H., Nong, Z., Freeman, L., Bensadoun, A., Peterson, K., Kruth, H., and Santamarina-Fojo, S. Identification of mouse and human macrophages as a site of synthesis of hepatic lipase. J. Lipid Res. 43:671-675. 2002. Conde-Knape K., Bensadoun, A., Sobel, J.L., Cohn, J.S., and Shachter, N.S. Overexpression of apolipoprotein C-1 in Apolipoprotein E-null Mice: Severe hypertriglyceredemia due to inhibition of hepatic lipase. J. Lipid Res. 43: 2136-2145. 2002. Yagyu, H., Chen,G., Yokoyama, M., Hirata, K., Augustus, A., Kako, Y., Seo T., Hu, Y., Lutz,P., merkel,m., Bensadoun, A., Homma, S. and Goldberg, I.J. Lipoprotein lipase (LPL) on the surface of cardiomyocytes increased lipid uptake and produced a cardiomyopathy. J. Clin. Inv. 111:419-426. 2003. Verges, M., Bensadoun, A., Herz, J., Belcher, J., and Havel, R.J. Endocytosis of hepatic lipase and lipoprotein lipase into rat hepatocytes in vivo is mediated by the low density lipoprotein receptorrelated protein. J. Biol. Chem. 279: 9030-9036, 2004 Lutz, E.P., Kako, Y.,Yagyu, H., Heeren, J.,Marks, S., Wright,T., Melford, K., Ben-Zeev, O., Radner,H., Merkel, M., Bensadoun, A., Wong, H., Goldberg, I.J. Mice expressing only covalent dimeric heparin-binding deficient lipoprotein lipase: Muscles inefficiently secrete dimeric enzyme. J.Biol.Chem. 279: 238-244. 2004. Espirito Santo, S.M.S., Pires, N.M.M., Gerritsen, G., Bovenschen, N., van Dijk, K.W., Princen, H.M.G., Bensadoun, A., Herz, J., Havekes, L.M., and van Vlijmen, B.J.M. Hepatic Low-Density Lipoprotein Receptor-Related Protein Deficiency in Mice Increases Atherosclerosis Independent of Plasma Cholesterol. Blood. 103: 3777-3782, 2004. Augustus, A.S., Yagyu, H., Haemmerle, G., Bensadoun, A., Vikramadithyan, R.K. Park, S.Y., Kim,J.K., Zekner, R., Goldberg, I.J. Cardiac-specific knockout of lipoprotein lipase alters plasma lipoprotein triglyceride metabolism and cardiac gene expression. J Biol Chem. 279: 25050-25057, 2004.

Bensadoun page 21 Li, H., Melford, K., Judson, A., and Bensadoun, A.. Murine glypican-4 gene structure and expression. Sp1 and Sp3 play a major role in glypican-4 expression in 3T3-F442A cells. Bioch. Biophys. Acta. 1679: 141-155, 2004. Ruel, I.L., Couture, P., Cohn, J.S., Bensadoun, A., Marcil, M., and B. Lamarche. Evidence that hepatic lipase deficiency in humans is not associated with proatherogenic changes in HDL composition and metabolism. J.Lipid Res. 45: 1528-1537, 2004. Nagashima, K., Lopez, C., Donovan, D., Ngai, C., Fontanez, N., Bensadoun, A., Fruchart-Najib, J., Holleran, S., Cohn, J.S., Ramakrishnan, R., Ginsberg, H.N. Effects of the PPARγ agonist, pioglitazone, on lipoprotein metabolism in patients with type 2 diabetes mellitus. J. Clin. Invest. 115:1323-1332, 2005. Espirito Santo SM, Rensen PC, Goudriaan JR, Bensadoun A, Bovenschen N, Voshol PJ, Havekes LM, Van Vlijmen BJ. Triglyceride-rich lipoprotein metabolism in unique VLDL receptor, LDL receptor, and LRP triple-deficient mice. J. Lipid Res. 46:1097-1102, 2005. Page, S. Judson, A., Melford, K., and Bensadoun, A. Interaction of lipoprotein lipase and receptor associated protein. J. Biol. Chem. 281:13931-13938, 2006. MacArthur, J.M., R. Bishop, J.R., Wang, L., Stanford, K.I., Bensadoun, A., Witztum, J.L., and Esko, J.D. Hepatic clearance of triglyceride-rich lipoproteins depends on heparan sulfate J. Clin. Invet. 117:153-164, 2007. Beigneux, A.P., Davies, B.S.J., Gin, P., Weinstein, M.M., Farber, E., Qiao, X., Peale, F., Bunting, S., Walzem, R.L., Wong, J.S., Blaner, W.S., Ding, Z.M., Melford, K., Wongsiriroj, N., Shu, X., de Sauvage, F., Ryan, R.O., Fong, L.G., Bensadoun, A., and Young, S.G. Glycosylphosphatidylinositol- Anchored High-density Lipoprotein Binding Protein 1 plays a critical role in the lipolytic processing of chylomicrons. Cell, metabolism. 5: 293-291, 2007. Young S.G., Davies, B.S., Fong, L.G., Gin, P., Weinstein, M.M., Bensadoun, A., and Beigneux, A.P. GPIHBP1: an endothelial cell molecule important for the lipolytic processing of chylomicrons. Curr Opin Lipidol. 18:389-96, 2007. Véniant, M.M., Beigneux, A.P., Bensadoun, A., Fong, L.G., and Young, S.G. 2007.Lipoprotein Size and Susceptibility to Atherosclerosis Insights from Genetically Modified Mouse Models. In Murine Atherosclerosis. G. Getz, editor: Betham Science Publishers. Bensadoun, A., Beigneux, A.P., Fong, L.G., and Young, S.G. 2007. New Developments in the Lipolytic Processing of Triglyceride-rich Lipoproteins. In Adipose Tissue in Health and Disease. T. Leff, editor: Wylie. Gin,P., Beigneux, A.P., Davies, B., Young, M.F., Ryan, R.O., Bensadoun, A., Fong, L.G., and Young, S.G. Normal binding of lipoprotein lipase, chylomicrons, and apo-av to GPIHBP1 containing a G56R amino acid substitution. Biochim. Biophys. Acta. 1771:1464-1468, 2007. Mayumi Takahashi, Yaeko Hiyama, Masayoshi Yokoyama, Shuiqing Yu, Yunying