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
Universite de Toulouse, France
Cornell University, Ithaca, NY
Eng. of ENSAT
Lic. Sc.
Ph.D.
engineering
biology
nutrition
1956
1956
1960
Fellowships and Awards
1981-82
1972-73
1971
1958-59
1974-03
National Research Service Award Senior Fellow
National Institutes of Health Special Fellowship
Traveling Fellowship, Cornell University
Henry Strong Denison Scholar, Cornell University
NIH-NHLBI continuous funding of HL-14990-29
Professional Experience
2011-2016
Graduate School Professor, Cornell University
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, ChampaignUrbana
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
I am currently a Cornell Graduate School professor of nutritional biochemistry with a full time
commitment to research.
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.
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).
Aside from its function to bind LPL on the surface of endothelial cells we have shown that it also acts
as a unique transporter of LPL across endothelial cells. 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 (phospholipids, cholesterol, ApoCs ) from VLDL and chylomicrons to HDL. In
support of this hypothesis GPI-HBP-1 deficient adult mice and humans fail to clear triglycerides from
blood.
Membership in Graduate Fields
Nutrition
Molecular and Integrative Physiology
Biochemistry, Molecular and Cell Biology
University Committees
Bensadoun – page 3
1981-83
1970-72
1976
1976-79
1976-78
1975-82
1978-81
1978-81
1978-81
1979-81
1984-87
1984-present
1989-1993
1989-91
1996-1998
1995-1998
1996-1998
Research Policy Committee
Standing Committee on Anatomy and Physiology, Chairman
Honors Committee, Division of Nutritional Sciences
Library Committee
Animal Facility Committee
Division of Biological Sciences Cell Biology Committee
Seminar Committee, Field of Physiology
Curriculum Committee, Section of Physiology
Division of Nutritional Sciences Controlled Substances Committee
Appointments and Tenure Committee
Fellowship Board Committee of the Graduate School
University Radiation Safety Committee
Nutritional Biochemistry Faculty Search Committee
Research Committee, Div. of Nutritional Sciences
Curriculum Committee, Nutritional Sciences
Appointment and tenure Committee
Animal Welfare Committee
Search committee for Section of Biochemistry (two positions)
Consultation with Public Agencies
1980-1981
1979-1982
Member of Committee B for Evaluation of SCOR's (Special Centers of Research) for
the Heart, Lung and Blood Institute, NIH
Site visitor for Program Projects administered by the Heart, Lung and Blood Institute
1981-1986
1983-1984
Ad Hoc member of the Metabolism Study Section. National Institutes of Health.
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
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
1P01HL090553-01A1 Young SG (PI)
Subcontract to Bensadoun
Percent time: 20%
08/01/2008-06/30/2013
Bensadoun – page 4
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
RO1 HL 087228
Young SG (PI)
Subcontract to Bensadoun
Percent time: 10%
Total costs:$40,325/year
Direct costs:$25,000/year
12/01/2011-12/31/2016
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.
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.
Bensadoun – page 5
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.
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 2223, 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.
Bensadoun – page 6
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.
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 TobaccoRelated Disease Research Program. April 1-6, 1992.
Bensadoun – page 7
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.
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.
Bensadoun – page 8
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.
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 – page 9
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.
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
Bensadoun – page 10
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.
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.
Bensadoun – page 11
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.
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.
Bensadoun – page 12
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.
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.
Bensadoun – page 13
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.
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
Bensadoun – page 14
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.
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.
Bensadoun – page 15
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.
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.
Bensadoun – page 16
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.
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: 297302, 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.
Bensadoun – page 17
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.
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.
Bensadoun – page 18
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: 217229, 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.
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.
Bensadoun – page 19
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.
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: 11831192. 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.
Bensadoun – page 20
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.
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 PPAR 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-
Bensadoun – page 21
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
Takahashi, M., Hiyama, Y., Yokoyama, M., Yu, S., Hu, Y., Melford, K., Bensadoun, A., and Goldberg,
I.J. In Vivo Arterial Lipoprotein Lipase Expression Augments Inflammatory Responses and Impairs
Vascular Dilatation. Arterioscler. Thromb. Vasc. Biol. 28:455-462, 2008.
Weinstein, M.M., Yin, L., Beigneux, A.P., Davies, B.S., Gin, P., Estrada, K., Melford, K., Bishop, J.R.,
Esko, J.D., Dallinga-Thie, G.M., Fong, L.G., Bensadoun, A., and Young, S.G. Abnormal patterns of
lipoprotein lipase release into plasma in GPIHBP1-deficient mice. J.Biol.Chem. 283(50):34511-34518,
2008.
Gin, P., Yin, L., Davies, B.S., Weinstein, M.M., Ryan, R.O., Bensadoun, A., Fong, L.G., Young, S.G.,
and Beigneux, A.P. The acidic domain of GPIHBP1 is important for the binding of lipoprotein lipase
and chylomicrons. J. Biol. Chem. 283(43): 29554-29562, 2008.
Beigneux, A.P., Gin, P., Davies,B.S., Weinstein, M.M., Bensadoun, A., Ryan, R.O., Fong, L.G., and
Young, S.G. Glycosylation of Ans-76 in mouse GPIHBP1 is critical for its appearance on the cell
surface and the binding of chylomicrons and lipoprotein lipase. J.Lipid Res. 49(6): 1312-1321,2008.
Beigneux, A.P., Davies, B.S., Bensadoun, A., Fong, L.G., and Young, S.G. GPIHBP1, a GPIanchored protein required for lipolytic processing of triglyceride-rich lipoproteins. J. Lipid Res.
Suppl:S57-62. 2009.
Beigneux, A.P., Weinstein, M.M., Davies, B.S., Gin, P., Bensadoun, A., Fong, L.G., and Young, S.G.
GPIHBP1 and lipolysis: an update. Curr Opin. Lipidol. 20:211-216, 2009.
Beigneux, A.P., Gin, P., Davies, B.S., Weinstein, M.M., Bensadoun, A., Fong, L.G., and Young, S.G.
Highly conserved cysteines within the Ly6 domain of GPIHBP1 are crucial for the binding of
lipoprotein lipase. J. Biol. Chem. 284:30240-30247, 2009.
Bensadoun – page 22
Bensadoun, A., Beigneux, A.P., Fong, L.G., and Young, S.G. New developments in the lipolytic
processing of triglyceride-rich lipoproteins. In Adipose Tissue in Health and Disease. Edited by Leff,
T.A. and Granneman, J. Wiley-VCH Verlag GmbH & Co. KGA, Weinheim; p 97-108. 2009
Franssen, R., Young, S.G., Peelman, F., Hertecant, J., Sierts, J.A., Schimmel,A.W.M.,
Bensadoun, A., Kastelein, J.J.P., Fong, L.G., Dallinga-Thie, G.M., Beigneux, A.P.
Chylomicronemia with low postheparin lipoprotein lipase levels in the setting of GPIHBP1
defects. Circ. Cardiovasc. Genet. 3:169-178, 2010.
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