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	S:	Pharmocokinetics of cholesterol absorption and mechanism of cholesterol absorption inhibition by EPA-DHA rich phytosterol ester versus ALA rich phytosterol ester
Avery Sengupta* & Mahua Ghosh
Dept. of Chemical Technology
University of Calcutta
India

*Corresponding author :               Dr. Avery Sengupta
				      Dept. of Chemical Technology
				      University College of Science & Technology
				      University of Calcutta
				      92, A.P.C. Road, Kolkata 700009
				      India

Phone numbers 	:     +91 33 23510050/51/52, Ext. 276
Fax Number	:     +91 33 23519755
       e-mail		:  HYPERLINK "mailto:avery.sgupta@gmail.com" avery.sgupta@gmail.com; 
Abstract: Hypercholesterolemia is a major cause of coronary heart disease. The present study was designed with different objectives; firstly to examine and compare the efficacy of n-3 fatty acid (alpha linolenic acid and eicosapentanoic acid-docosahexanoic acid) esters of phytosterol on cholesterol absorption in hypercholesterolemia and secondly to evaluate the optimum dosage, action and time of treatment of phytosterol esters as a drug in hypercholesterolemia by performing cholesterol absorption kinetics to obtain best results. Sterol esters were prepared by packed bed bioreactor using Thermomyces lanuginosus. Sterol esters at two different dosages were fed to male Wistar rats for 32 days and plasma lipid profile was analyzed. Cholesterol absorption was measured at each week and absorption kinetics was performed. Results show that sterol esters reduced the hyperlipidemic effect of hypercholesterolemia both in plasma and liver. Cholesterol absorption decreased by the administration of sterol esters. Cholesterol absorption inhibition was brought about by increase in the expression of liver X receptor (LXR) as seen from Western Blot analysis. Higher dose (0.5mg/kg body weight) of EPA-DHA rich sterol ester provided the best result against the ill effects of hypercholesterolemia. The three week treatment of sterol ester was quite effective for hypercholesterolemia as indicated from the kinetic study. 
Keywords hypercholesterolemia; sterol esters; cholesterol absorption kinetics; liver X  receptor

Abbreviations
ALA alpha linolenic acid
EPA eicosapentanoic acid
DHA docosahexanoic acid
LXR liver X receptor
CAD Coronary artery disease
LDL low density lipoprotein
ABC ATP-binding cassette
TLIM Thermomyces lanuginosus Immobilized
CYP7A1 cholesterol 7�-hydroxylase
Introduction:       
Coronary artery disease (CAD) is the major cause of morbidity and mortality in industrialized countries. It is well established that plasma levels of cholesterol are strongly associated with the incidence of coronary atherosclerosis. Epidemiological studies have shown us the association between cholesterol and disease and the clinical studies have provided additional information regarding the benefits of lipid lowering therapies. The most commonly used cholesterol lowering agents contribute to the prevention of cardiovascular disease through their action on cholesterol metabolism, namely absorption and synthesis. 
        Plant sterols (phytosterols) have been proposed as one of the non-prescriptive methods for lowering plasma cholesterol levels to reduce CAD risk. The effectiveness of the phytosterols in lowering blood concentrations of cholesterol was recognized as early as 1950s. It is generally believed that the primary mechanism for lowering blood cholesterol levels is to reduce the absorption of cholesterol from the small intestine by competing for the limited space for cholesterol in bile salt micelles [ ADDIN EN.CITE <EndNote><Cite><Author>Ikeda</Author><Year>1989</Year><RecNum>79</RecNum><DisplayText>(Ikeda, Tanabe, &amp; Sugano, 1989)</DisplayText><record><rec-number>79</rec-number><foreign-keys><key app="EN" db-id="2vtasw5p4tf0w4ev2vyp2sagsxasdw25vvpe">79</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Ikeda, I.</author><author>Tanabe, Y.</author><author>Sugano, M.</author></authors></contributors><auth-address>Laboratory of Nutrition Chemistry, Kyushu University School of Agriculture, Fukuoka, Japan.</auth-address><titles><title>Effects of sitosterol and sitostanol on micellar solubility of cholesterol</title><secondary-title>J Nutr Sci Vitaminol (Tokyo)</secondary-title></titles><periodical><full-title>J Nutr Sci Vitaminol (Tokyo)</full-title></periodical><pages>361-9</pages><volume>35</volume><number>4</number><edition>1989/08/01</edition><keywords><keyword>Animals</keyword><keyword>Cholesterol/*metabolism</keyword><keyword>Intestinal Absorption/drug effects</keyword><keyword>Male</keyword><keyword>Micelles</keyword><keyword>Rats</keyword><keyword>Sitosterols/*pharmacology</keyword><keyword>Solubility</keyword><keyword>Spectrophotometry</keyword><keyword>Ultracentrifugation</keyword></keywords><dates><year>1989</year><pub-dates><date>Aug</date></pub-dates></dates><isbn>0301-4800 (Print)&#xD;0301-4800 (Linking)</isbn><accession-num>2585153</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/pubmed/2585153</url></related-urls></urls><language>eng</language></record></Cite></EndNote> HYPERLINK  \l "_ENREF_3" \o "Ikeda, 1989 #79" 1]. Phytosterols have also been described to interfere in cholesterol absorption by competing with cholesterol transporters. The inhibition of cholesterol absorption produces a state of relative cholesterol deficiency, followed by an up regulation of cholesterol biosynthesis and low-density lipoprotein (LDL) receptor activity [ ADDIN EN.CITE <EndNote><Cite><Author>Ling</Author><Year>1995</Year><RecNum>103</RecNum><DisplayText>(Ling &amp; Jones, 1995)</DisplayText><record><rec-number>103</rec-number><foreign-keys><key app="EN" db-id="2vtasw5p4tf0w4ev2vyp2sagsxasdw25vvpe">103</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Ling, W. H.</author><author>Jones, P. J.</author></authors></contributors><auth-address>School of Dietetics and Human Nutrition, McGill University at Macdonald Campus, Ste-Anee-de-Bellevue, PQ, Canada.</auth-address><titles><title>Dietary phytosterols: a review of metabolism, benefits and side effects</title><secondary-title>Life Sci</secondary-title></titles><periodical><full-title>Life Sci</full-title></periodical><pages>195-206</pages><volume>57</volume><number>3</number><edition>1995/01/01</edition><keywords><keyword>Animals</keyword><keyword>Antineoplastic Agents/pharmacology</keyword><keyword>Cholesterol/blood/metabolism</keyword><keyword>*Diet</keyword><keyword>Humans</keyword><keyword>Intestinal Absorption</keyword><keyword>Lipoproteins/biosynthesis/blood</keyword><keyword>Phytosterols/adverse effects/metabolism/*pharmacology</keyword><keyword>Sterol O-Acyltransferase/metabolism</keyword></keywords><dates><year>1995</year></dates><isbn>0024-3205 (Print)&#xD;0024-3205 (Linking)</isbn><accession-num>7596226</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/pubmed/7596226</url></related-urls></urls><electronic-resource-num>0024320595002636 [pii]</electronic-resource-num><language>eng</language></record></Cite></EndNote> HYPERLINK  \l "_ENREF_4" \o "Ling, 1995 #103" 2] After chronic phytosterol feeding, the ratio of cholesterol precursors to cholesterol rises to 46% [ ADDIN EN.CITE <EndNote><Cite><Author>Gylling</Author><Year>1999</Year><RecNum>122</RecNum><DisplayText>(Gylling, Puska, Vartiainen, &amp; Miettinen, 1999)</DisplayText><record><rec-number>122</rec-number><foreign-keys><key app="EN" db-id="2vtasw5p4tf0w4ev2vyp2sagsxasdw25vvpe">122</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Gylling, H.</author><author>Puska, P.</author><author>Vartiainen, E.</author><author>Miettinen, T. A.</author></authors></contributors><auth-address>Department of Medicine, University of Helsinki, P.O. Box 340, FIN-00029 HYKS, Helsinki, Finland.</auth-address><titles><title>Serum sterols during stanol ester feeding in a mildly hypercholesterolemic population</title><secondary-title>J Lipid Res</secondary-title></titles><periodical><full-title>J Lipid Res</full-title></periodical><pages>593-600</pages><volume>40</volume><number>4</number><edition>1999/04/07</edition><keywords><keyword>Anticholesteremic Agents/*administration &amp; dosage</keyword><keyword>Body Mass Index</keyword><keyword>Cholestanol/blood</keyword><keyword>Cholesterol/analogs &amp; derivatives/blood</keyword><keyword>Dietary Fats, Unsaturated/*administration &amp; dosage</keyword><keyword>Esters</keyword><keyword>*Fatty Acids, Monounsaturated</keyword><keyword>Humans</keyword><keyword>Hypercholesterolemia/*blood</keyword><keyword>Kinetics</keyword><keyword>Margarine</keyword><keyword>*Phytosterols</keyword><keyword>Sitosterols/*administration &amp; dosage/blood</keyword><keyword>Sterols/*blood</keyword></keywords><dates><year>1999</year><pub-dates><date>Apr</date></pub-dates></dates><isbn>0022-2275 (Print)&#xD;0022-2275 (Linking)</isbn><accession-num>10191283</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/pubmed/10191283</url></related-urls></urls><language>eng</language></record></Cite></EndNote> HYPERLINK  \l "_ENREF_2" \o "Gylling, 1999 #122" 3] and LDL receptor expression increases by 25% to 42%. Also, changes in LDL levels are correlated with LDL receptor expression in peripheral cells [ ADDIN EN.CITE <EndNote><Cite><Author>Plat</Author><Year>2002</Year><RecNum>128</RecNum><DisplayText>(Plat &amp; Mensink, 2002)</DisplayText><record><rec-number>128</rec-number><foreign-keys><key app="EN" db-id="2vtasw5p4tf0w4ev2vyp2sagsxasdw25vvpe">128</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Plat, J.</author><author>Mensink, R. P.</author></authors></contributors><auth-address>Department of Human Biology, Maastricht University, 6200 MD, Maastricht, The Netherlands. J.Plat@HB.UNIMAAS.NL</auth-address><titles><title>Effects of plant stanol esters on LDL receptor protein expression and on LDL receptor and HMG-CoA reductase mRNA expression in mononuclear blood cells of healthy men and women</title><secondary-title>FASEB J</secondary-title></titles><periodical><full-title>FASEB J</full-title></periodical><pages>258-60</pages><volume>16</volume><number>2</number><edition>2002/01/05</edition><keywords><keyword>Cholesterol/blood</keyword><keyword>Cholesterol, LDL/blood/drug effects</keyword><keyword>Esters/pharmacology</keyword><keyword>Female</keyword><keyword>Gene Expression Regulation/drug effects</keyword><keyword>Humans</keyword><keyword>Hydroxymethylglutaryl CoA Reductases/*genetics</keyword><keyword>Leukocytes, Mononuclear/cytology/*drug effects/metabolism</keyword><keyword>Male</keyword><keyword>Phytosterols/blood/chemistry/*pharmacology</keyword><keyword>RNA, Messenger/*drug effects/genetics/metabolism</keyword><keyword>Receptors, LDL/*drug effects/genetics/metabolism</keyword></keywords><dates><year>2002</year><pub-dates><date>Feb</date></pub-dates></dates><isbn>1530-6860 (Electronic)&#xD;0892-6638 (Linking)</isbn><accession-num>11772951</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/pubmed/11772951</url></related-urls></urls><electronic-resource-num>10.1096/fj.01-0653fje&#xD;01-0653fje [pii]</electronic-resource-num><language>eng</language></record></Cite></EndNote> HYPERLINK  \l "_ENREF_6" \o "Plat, 2002 #128" 4]. However the net result is a decrease in cholesterol concentration after phytosterol feeding. Recently studies have shown that, in general, phytosterols added to margarines reduce plasma total cholesterol by approximately 10% and low-density lipoprotein cholesterol (LDL-C) by up to 15%. But phytosterols have several disadvantages when administered as drugs. Esterification of phytosterols with fatty acids somewhat overcomes this drawback by increasing their lipophilicity and oil solubility, and so being started commercial applied by incorporating them into various products [5]. Food or drug components with high lipophilicity may compete with each other with similar lipophilicity for absorption [6].  
         Cholesterol homeostasis in liver is usually maintained by balancing intestinal cholesterol absorption and endogenous cholesterol synthesis. Reducing cholesterol biosynthesis and increasing cholesterol catabolism are both useful treatments for hypercholesterolemia. Currently, most clinical attention is focused on reducing biosynthesis using statin drugs; less attention has been paid to increasing cholesterol catabolism. Because cholesterol is principally excreted in the stool without undergoing tissue metabolism, the efficiency of cholesterol absorption is a critical determinant of cholesterol catabolism (i.e. excretion of unabsorbed dietary and biliary cholesterol is the largest component of reverse cholesterol transport) [7]. Cholesterol absorption seems well suited to play a regulatory role in whole-body cholesterol homeostasis by interacting with dietary and endogenous factors. There are three sources for intestinal cholesterol: the diet, the bile, and intestinal epithelial sloughing. Cholesterol absorption begins in the stomach when dietary constituents are mixed with lingual and gastric enzymes. Although the entire length of the small intestine has the capability to absorb cholesterol from the lumen, main sites of absorption are the duodenum and proximal jejunum. When the dietary conditions are controlled, biliary factors such as secretion rates of biliary lipids, cholesterol content of bile, as well as size, molecular composition, and hydrophilic-hydrophobic balance of the bile salt pool [ ADDIN EN.CITE  ADDIN EN.CITE.DATA  HYPERLINK  \l "_ENREF_14" \o "Wang, 1999 #394" 8] could together exert major influences on the efficiency of intestinal cholesterol absorption. Although a simple passive diffusion process is widely assumed for intestinal cholesterol absorption, several lines of evidence support the existence of an energy-independent, protein-facilitated mechanism for cholesterol uptake by the enterocyte [ ADDIN EN.CITE <EndNote><Cite><Author>Thurnhofer</Author><Year>1990</Year><RecNum>146</RecNum><DisplayText>(Thurnhofer and Hauser 1990)</DisplayText><record><rec-number>146</rec-number><foreign-keys><key app="EN" db-id="d5f02wfd6pwws1e9rvl5asxf9sew5ewt02s9">146</key></foreign-keys><ref-type name="Journal Article">17</ref-type><contributors><authors><author>Thurnhofer, H.</author><author>Hauser, H.</author></authors></contributors><auth-address>Laboratorium fur Biochemie, Eidgenossische Technische Hochschule Zurich, Switzerland.</auth-address><titles><title>Uptake of cholesterol by small intestinal brush border membrane is protein-mediated</title><secondary-title>Biochemistry</secondary-title></titles><periodical><full-title>Biochemistry</full-title></periodical><pages>2142-8</pages><volume>29</volume><number>8</number><edition>1990/02/27</edition><keywords><keyword>Animals</keyword><keyword>Cell Membrane/metabolism/ultrastructure</keyword><keyword>Centrifugation</keyword><keyword>Cholesterol/*metabolism</keyword><keyword>Cytosol/metabolism</keyword><keyword>Intestine, Small/*metabolism/ultrastructure</keyword><keyword>Kinetics</keyword><keyword>Microvilli/metabolism</keyword><keyword>Papain</keyword><keyword>Rabbits</keyword></keywords><dates><year>1990</year><pub-dates><date>Feb 27</date></pub-dates></dates><isbn>0006-2960 (Print)&#xD;0006-2960 (Linking)</isbn><accession-num>2328246</accession-num><urls><related-urls><url>http://www.ncbi.nlm.nih.gov/pubmed/2328246</url></related-urls></urls><language>eng</language></record></Cite></EndNote> HYPERLINK  \l "_ENREF_13" \o "Thurnhofer, 1990 #146" 9]. Intestinal cholesterol absorption can be specifically inhibited by cholesterol absorption inhibitors of different chemical structure. Literature showed that intestinal ATP-binding cassette (ABC) transporter ABCA1 serves to efflux cholesterol from the enterocyte back to the intestinal lumen, thereby modulating net cholesterol absorption efficiency [10], as well as cholesterol absorption and the transcriptional regulation of ABCA1 gene expression are mediated by the RXR/LXR heterodimer. Increasing the activity of ABCA1 transporter thus decreases cholesterol absorption also. There is enough literature regarding the role of phytosterols on cholesterol absorption, but very little information is provided regarding the role of phytosterol ester on cholesterol absorption. Moreover, there is also almost no information relating to the action of phytosterol ester as a drug on cholesterol absorption, the dosage of the drug and the amount of time the drug takes to perform its maximum action on preventing cholesterol absorption and thus treating hypercholesterolemia. 
	The present study was designed with multiple objectives; firstly to examine and compare the efficacy of n-3 fatty acid (alpha linolenic acid and eicosapentanoic acid- docosahexanoic acid) esters of phytosterol on cholesterol absorption in hypercholesterolemia; secondly to evaluate the optimum dosage, action and time of treatment of phytosterol esters as a drug in hypercholesterolemia by performing cholesterol absorption kinetics to obtain best results; and thirdly to determine the mechanism by which sterol esters reduce cholesterol absorption.


Materials and methods: 
Preparation of phytosterol esters
A standard �-sitosterol sample was procured from Fluka Chemicals and analyzed at the laboratory by gas chromatography (GC). Fish oil (Mega-Shelcal capsules from Elder Pharmaceuticals, India) was used as the source of eicosapentanoic acid (EPA) and docosahexaenoic acid (DHA), and the GC analysis of the fish oil showed that the oil contained 32% EPA and 22% DHA. Refined and bleached linseed oil procured from V.K.V.K. Oil Limited, Kolkata, India, was used as the source of alpha linolenic acid (ALA), and the GC analysis of the oil showed the presence of 54% ALA in the oil. Thermomyces lanuginosus lipase (Lipozyme TLIM), used as biocatalyst, was a gift of Novozyme India, Ltd., Bangalore, India. Phytosterol esters were produced by enzymatic transesterification reaction with sterol and corresponding oil in a packed bed reactor and their fatty acid composition was analyzed by GC.
Animals and treatments
Adult male albino rats of Wistar strain were housed and given food and water ad libidum. The duration of the experimental period was 32 days. The animals were divided into six groups with six rats in each: Group I: vehicle treated control animals, Group II: rats were fed with a high cholesterol diet [rat stock diet supplemented with cholesterol (1% of dietary lipid) for 32 days, Group III: rats received EPA-DHA phytosterol ester (0.25g/kg body weight/day, oral gavage) for last 25days along with a high cholesterol diet for 32days, Group IV: : rats received EPA-DHA phytosterol ester (0.5g/kg body weight/day, oral gavage) for last 25days along with a high cholesterol diet for 32days, Group V: rats fed with high cholesterol diet for 32 days and ALA phytosterol ester (0.25g/kg body weight/day, oral gavage) for the last 25days, Group VI: rats fed with high cholesterol diet for 32 days and ALA phytosterol ester (0.5g/kg body weight/day, oral gavage) for the last 25days. Groundnut oil was used as the vehicle and given to all the groups. At the end of the experiment the feeding of rats was stopped and after 12h fasting, the rats were anesthetized by chloroform and 5ml of blood was taken from the heart. The plasma was obtained by centrifugation of the blood. The liver, brain and mesentery was removed, rinsed with ice-cold saline, blotted, weighed and stored at -20�C until analyzed. The experimental protocol was approved by the Animal Ethical Committee of Dept. of Chemical Technology, University of Calcutta. 
Plasma Lipid Profile
Blood samples for laboratory analyses were taken into EDTA-containing vacuum tubes after 12h fasting. Plasma was separated by centrifugation and stored at -70�C until analyzed. In the analysis of lipid profile plasma lipids, samples were taken into glass centrifuge tubes and homogenized with different proportions of chloroform/methanol (vol/vol) and then centrifuged. Redistilled water was added to the mixture and the chloroform layer was separated in a separatory funnel. The chloroform was then evaporated to obtain the plasma lipids. The lipid components such as total cholesterol, high density lipoprotein (HDL) cholesterol and triacylglycerol were analyzed using enzyme kits supplied by Merck.
Determination of cholesterol absorption
Cholesterol absorption was measured by the peroral double-isotope feeding method [11] each week for four successive weeks.
Pharmacokinetic Analysis of Cholesterol Absorption
The weight gain, food intake and cholesterol absorption data were analyzed by the four-parameter mathematical model for physiological responses. The equation for the model is as follows:
 EMBED Equation.3  

where r is the physiological response; I is the nutrient intake or concentration in the diet; b is the intercept on the r axis; Rmax is the maximum response; n is the apparent kinetic order; K0.5 is the intake for � (Rmax + b).
      Although several non-linear curve-fitting programs are available and satisfactory for fitting the model, we have used a Gauss-Newton method based on the BMDX 85 program of the UCLA computer center. This procedure gives rapid convergence when there are good starting estimates for the four parameters.
Western blot Analysis
Western blot analysis of Liver X Receptor was carried out with commercially available kits (Amersham) according to manufacturer's instructions. Briefly, 25 �g protein isolated from the liver cells were subjected to PAGE and transferred to a nitrocellulose membrane. Blots were incubated with mouse anti-LXR INCLUDEPICTURE "http://joe.endocrinology-journals.org/math/alpha.gif" \* MERGEFORMATINET  (1:1000) or rabbit anti-RXR INCLUDEPICTURE "http://joe.endocrinology-journals.org/math/alpha.gif" \* MERGEFORMATINET  (1:200) antibodies for 1 h, followed by 2� goat HRP-conjugated antibody (1:5000) for an additional hour and visualized by enhanced chemiluminescence. Anti-�-actin was used as a loading control. 
Statistical Analysis
Statistical analysis was performed with either 1-way ANOVA or 2-way ANOVA and Fisher s test, if appropriate. Statistical significance was designated as P<0.05. Values are expressed as Mean� SEM.

Results:
Fatty Acid Composition of Phytosterol Esters
The transesterification reaction was carried out in the packed bed reactor.  In the first set of reaction fish oil was used as a source of EPA & DHA fatty acids and in the second set, linseed oil was used as a source of ALA fatty acid. Approximately 32% EPA & 22% DHA was present in fish oil and 54% of ALA in linseed oil. Analysis of fatty acid composition of sterol-esters by GC showed that almost all the fatty acids present in the different oils (in TAG form) were incorporated in the corresponding  esters. Table 1 shows the fatty acid profile of the sterol esters produced in the packed bed reactor.
Changes in Plasma Lipid Profile
The plasma lipid profile of rats of different groups is shown in Table 2. The type of fat consumed altered the different lipid concentration in plasma. Rats fed with control Groundnut Oil had plasma total cholesterol of 54.37 mg/dL which was significantly increased to 246.83 mg/dL by feeding them high cholesterol diet. Both the doses of phytosterol ester brought about a decrease in total cholesterol which was much more in case of EPA-DHA rich sterol ester. The higher doses produced better hypocholesterolemic effect than the lower dose. Similar results were found in case of estimation of non-HDL cholesterol levels and triacylglycerol levels. Thus the phytosterol esters lowered the levels of non-HDL cholesterol and triacylglycerol significantly (p<0.05). The level of HDL cholesterol, which was known as good cholesterol, increased significantly (p<0.05) by treating the rats with phytosterol esters. The increase in the HDL level was much more in case of the rats treated with EPA-DHA rich sterol ester.
Kinetic Analysis
Weight gain and food intake responses
The observed body weight and food intakes are shown in Figure 1.A. and 1.B. The curves are sigmoidal in shape and demonstrate the usual threshold phase, sensitive or rapidly changing phase and the plateau or saturation phase. After 21 days the values of all the groups become saturated and attain a plateau stage. 
Cholesterol absorption kinetics
The cholesterol absorption depended on the type and dose of treatment of sterol esters. The functional relationship between cholesterol absorption and sterol esters was quite varied. Many of the responses as well as some of the pools and ratios can be described by the four-parameter mathematical model. The cholesterol absorption decreased with time and after some days it became stable. Sterol esters brought about the decrease in cholesterol absorption. EPA-DHA rich sterol ester was more effective in reducing cholesterol absorption in comparison with ALA rich sterol ester. Similarly the higher doses of both the esters showed better results (Figure 2).
Western blot results
We evaluated the LXR expression by western blot analysis to investigate the cellular mechanism of cholesterol absorption. LXR expression was detected in liver samples and the effect of sterol esters on LXR expression to attenuate hypercholesterolemia was investigated. It is seen from Figure 3 that LXR expression was enhanced by administration of both the sterol esters to hypercholesterolemic subjects. LXR expression was maximum in case of higher dose of EPA-DHA rich sterol ester and minimum in case of lower dose of ALA rich sterol ester. 
Discussion:
       This study shows that under controlled dietary conditions, both sterol esters possess improved efficiency in reducing plasma total cholesterol (TC), LDL-cholesterol (LDL-C) and triacylglycerol (TG) concentrations at both the dietary doses. The effect of EPA-DHA rich sterol ester was much improved in comparison to ALA rich sterol ester. At higher doses the hypolipidemic effect was much enhanced compared to the lower dose. Similar results were also seen in the case of hepatic lipid profile. This reduction occurred as a result of the ability of sterol esters to depress cholesterol absorption. The fact that the final circulatory TC level was positively associated with the cholesterol absorption, particularly in the group showing the most pronounced cholesterol lowering, indicates inhibition of absorption as a chief mechanism in the cholesterol-modulating effects of these dietary agents. In addition to sterols in sterol esters, dietary fatty acid composition may have also been partially responsible for the changes in lipids observed.  Differences between study design including sterol ester type and dosage, or dietary composition may be responsible for the variable results observed in the present study. 
         Major sub-objectives of the present research were to assess mechanisms of action through which these sterol esters act. To meet these objectives, we utilized a novel combination of technique for measurement of cholesterol absorption, kinetics and mechanism of cholesterol absorption. 
       The dual isotope absorption measurement method uses intravenously administered labeled cholesterol as a tracer to monitor of exit rate of both body and dietary cholesterol from the rapid pool relative to orally administered cholesterol labeled with a second tracer to monitor the intestinal absorption. Presently from the data it could be seen that cholesterol absorption decreased by about 45% with the lower dose of sterol esters and to about 75% with the higher dose. Thus it could be conferred that sterol esters successfully reduced plasma total cholesterol by reducing cholesterol absorption. 
           Living organisms are considered to be thermodynamically open systems in a condition of dynamic, steady-state equilibrium. The organism has available to it homeostatic mechanisms by which it regulate its interaction with the environment so that it can maintain equilibrium. These dynamic processes preserve the constancy of physiological variables in time, i.e. a steady state. During nutrient deprivation, new equilibrium states may be established and steady-state values may be adjusted. In many instances these values can be shown to be functions of the environmental configuration, but within limitations imposed by the genetic makeup of the organism. In this experiment, the environmental configuration was manipulated by changing the type of a dietary nutrient (lipid), the physiological variables measured were weight gain, food intake and cholesterol absorption kinetics, and the functional relationship was expressed by the four-parameter model. The four-parameter model was based on the concept of saturation kinetics. In a series of reactions there is a rate-limiting saturable step, and the observed physiological response reflects the kinetics of that step. Figure 1.A. and 1.B. shows weight gain and food intake as functions of time. Both responses tend to �saturate�, i.e. plateau out. Since the diets are isoenergetic, differences in food intake are probably caused by specific effects of lipids or their metabolites. Subsequent metabolic activities cause alterations in the level of plasma and liver cholesterol.  Figure 3 shows the cholesterol absorption as function of time. It also shows the rate of absorption of cholesterol by treatment with different doses and types of sterol esters. Kinetic data shows that the absorption of cholesterol decreases with time and becomes constant after 3 weeks. After 3 weeks there is no further decrease in cholesterol absorption. Figure 3 also shows that cholesterol absorption decreases much more in higher doses of sterol esters than the lower one. There was however a significant negative correlation between cholesterol absorption with time in rats fed with cholesterol in the diet. The negativity increased with the increase in the dose. 
            Nuclear receptors are transcription factors that act as intracellular receptors for small biologically active molecules such as lipids and steroid hormones. In contrast to extracellular receptors that bind peptide ligands and activate a cascade of cytoplasmic kinases, nuclear receptors are intracellular, have lipophilic ligands, and directly regulate the expression of target genes by binding to specific DNA sequences in the promoter of target genes. Liver X receptors (LXR) belong to the family of ligand-activated transcription factors with a key role in the control of lipid metabolism and inflammation. In the liver, LXR activation promotes cholesterol elimination in two different ways: first, by induction of the expression of cholesterol 7�-hydroxylase (CYP7A1), the rate-limiting enzyme in the classical pathway of bile acid biosynthesis [12]; second, by promoting the transcription of genes for ATP-binding cassette transporters, ABCG5 and ABCG8 that transport cholesterol from the hepatocytes to the bile canaliculi [13]. LXR also offers extrahepatic tissue protection from cholesterol accumulation. Through the control of reverse cholesterol transport in macrophages, LXR ligands induce the expression of ABCA1, ABCG1, and ABCG4 transporters that promote the efflux of cholesterol to HDL [14,15]. In our study sterol ester treatment resulted in increased expression of LXR in comparison to hypercholesterolemic control. The higher dose of sterol esters showed increased concentration of LXR in liver tissue in comparison to the lower one. This shows the actual mechanism of cholesterol lowering which is due to the increased presence of LXR in the liver tissues. 
                In conclusion, the present study demonstrates that in esterified forms, phytosterol efficacy in total, LDL-cholesterol and triacylglycerol lowering both in plasma and liver may be influenced by both phytosterol and the fatty acids present in the esters. Moreover, a mechanism contributing to the action of plant sterols in lowering the above lipid parameters is through reduction of absorption of dietary cholesterol. Kinetic data showed that body weight of the rats increased up to 3 weeks of treatment as food intake became constant after 3 weeks. Cholesterol absorption kinetics also proved that inhibition of cholesterol absorption was maximum at 3 weeks or 21 days and after that there was no further decrease in the cholesterol absorption. Therefore to conclude that to combat against hypercholesterolemia 0.5g of EPA-DHA rich sterol ester/day for 21 days is advisable as a drug. 


Acknowledgement:  
Financial support obtained from University of Calcutta is gratefully acknowledged by the authors. Authors are sincerely thankful to Dr. S. Ghosh, Dept. of Chemical Technology for his help and encouragement. 

 ADDIN EN.REFLIST References
1.Ikeda I, Tanabe Y, Sugano M (1989) Effects of sitosterol and sitostanol on micellar solubility of cholesterol. J Nutr Sci Vitaminol 35:361-369.
2. Ling WH, Jones PJ (1995) Dietary phytosterols: a review of metabolism, benefits and side effects. Life Sci 57:195-206. 
3. Gylling H, Puska P, Vartiainen E, et al. (1999) Serum sterols during stanol ester feeding in a mildly hypercholesterolemic population. J Lipid Res 40:593-600.
4. Plat J, Mensink RP (2002) Effects of plant stanol esters on LDL receptor protein expression and on LDL receptor and HMG-CoA reductase mRNA expression in mononuclear blood cells of healthy men and women. FASEB J 16:258-260.
5. Weststrate JA, Meijer GW (1998) Plant sterol-enriched margarines and reduction of plasma total- and LDL-cholesterol concentrations in normocholesterolaemic and mildly hypercholesterolaemic subjects. Eur J Clin Nutr 52:334-343.
6. Bailey C (1996) New guidelines for cardiac rehabilitation. Colo Nurse 96:9-10.
7. Ostlund RE (2004) Phytosterols and cholesterol metabolism. Curr Opin Lipidol 15:37-41. 
8. Wang DQ, Lammert F, Cohen DE, et al. (1999) Cholic acid aids absorption, biliary secretion, and phase transitions of cholesterol in murine cholelithogenesis. Am J Physiol 276:G751-760.
9. Thurnhofer H, Hauser H (1990) Uptake of cholesterol by small intestinal brush border membrane is protein-mediated. Biochem 29:2142-2148.
10. Repa JJ, Turley SD, Lobaccaro JA, et al. (2000) Regulation of absorption and ABC1-mediated efflux of cholesterol by RXR heterodimers. Science 289:1524-1529. 
11. Crouse JR, Grundy SM (1978) Evaluation of a continuous isotope feeding method for measurement of cholesterol absorption in man. J Lipid Res 19:967-971.
12. Peet DJ, Turley SD, Ma W, et al. (1998) Cholesterol and bile acid metabolism are impaired in mice lacking the nuclear oxysterol receptor LXR alpha. Cell 93:693-704. 
13. Berge KE, von Bergmann K, Lutjohann D, et al. (2002) Heritability of plasma noncholesterol sterols and relationship to DNA sequence polymorphism in ABCG5 and ABCG8. J Lipid Res 43:486-494.
14. Costet P, Luo Y, Wang N, et al. (2000) Sterol-dependent transactivation of the ABC1 promoter by the liver X receptor/retinoid X receptor. J Biol Chem 275:28240-28245. 
15. Engel T, Lorkowski S, Lueken A, et al. (2001) The human ABCG4 gene is regulated by oxysterols and retinoids in monocyte-derived macrophages. Biochem Biophys Res Commun 288:483-488. 









Figure Legends

Fig. 1.A  Body weight as a function of time. Each value is the mean of 6 rats. (p<0.05). 

Fig. 1.B Mean daily food intake as a function of time. Each value is the mean of 6 rats. 
              (p<0.05).   

Fig. 2 Cholesterol absorption as a function of time. Values are Mean S.E.M. (p<0.05)
           A. Changes in cholesterol absorption in case of EPA-DHA rich Ester 
           B. Changes in cholesterol absorption in case of ALA rich Ester

Fig. 3 LXR� Expression in Different Groups of Rats



  




 

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��d��^��gd!&terol Esters used as Nutraceutical
       Fatty     
          acid
Sample
                    Fatty Acid (% w/w)       C14:0C16:0C18:0C18:1C18:2C18:3C20:0C20:1C20:5C22:0C22:1C24:0C22:6DHA-EPA rich Ester 0.10
�
0.002.44
�
0.041.23
�
0.021.0
�
0.151.90
�
0.273.05
�
0.342.75
�
0.645.12
�
0.8339.20
�
1.105.65
�
0.372.19
�
0.083.84
�
0.1931.98
�
1.63ALA rich Ester
-12.48 
�
0.601.89 
�
0.1728.28 
�
1.0912.83 
�
1.0144.52 
�
1.76-------Values are expressed as mean � S.E.M

Table 2. Plasma Lipid Profiles (mg/dL) of Different Rats fed with Different Dietary Lipids
ParametersGroup I
Group II
Group III
Group IV
Group V
Group VI
Total Cholesterol54.37�1.20246.83�11.32a87.07�2.30b63.27�2.45b,c96.76�1.98b,e65.75�0.40b,dHDL Cholesterol23.98�1.2212.23�1.09a20.80�0.98b39.19�0.38b,c23.36�0.44b,e34.23�1.76b,d,fNon-HDL Cholesterol20.39�0.77213.44�3.40a54.21�0.99b14.52�0.78b,c58.66�1.56b,e20.63�0.56b,d,fTriglycerides49.98�0.57105.79�4.50a60.32�0.59b47.82�0.33b,c73.69�1.89e54.47�2.90b,d,fValues are expressed as mean � S.E.M. for six animals. The superscript letters also represent statistical significance at p<0.05.
a Comparisons are made between Groups I and II
b Comparisons are made between Groups II and III-VI
c Comparisons are made between Groups III and IV
d Comparisons are made between Groups V and VI
e Comparisons are made between Groups III and V
f Comparisons are made between Groups IV and VI


 EMBED Origin50.Graph  

 EMBED Origin50.Graph  

Fig. 1

 EMBED Origin50.Graph  
A
 EMBED Origin50.Graph  
B
Fig. 2




                                              
Fig. 3











 PAGE   \* MERGEFORMAT 23



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