Amounts Of Sterols Research Articles

Screening of macro- and bioactive microconstituents of commercial finfish and sea urchin eggs

Crude and lipid composition, tocopherols, carotenoids, sterols and squalene content, together with fatty acids profiles of commercial marine egg products are reported, namely (a) canned eggs of lumpfish (Cyclopteris lumpus), Atlantic salmon (Salmo salar), trout (Oncorhynchus mykiss), smoked herring (Clupea harengus), and sea urchin (Paracentrotus lividus), (b) fresh sea urchin (P. lividus) eggs, and (c) beeswax covered, dried and salted mullet (Mugil cephalus) skeins -Greek “Avgotaracho”. The protein content of marine eggs ranged from 8.7 to 33.7 g/100 g on a fresh weight (fw) basis. Among neutral lipids triacylglycerols predominated, except for mullet roe where wax and steryl esters comprised 75% of neutral lipids. Phosphatidylcholine was the major phospholipid, followed by phosphatidylethanolamine. Marine eggs contained 0.18–2.39 g/100 g fw long chain ω3 polyunsaturated fatty acids (PUFA), mainly docosahexaenoic acid (DHA) in finfish eggs and eicosapentaenoic acid (EPA) in urchin eggs. They also contained 86.2–342.0 mg cholesterol/100 g fw, low amounts of squalene and other sterols and significant amounts of tocopherols (0.45–4.4 mg/100 g) and carotenoids (0.31–11.4 mg/100 g). The commercial finfish and urchin eggs are food of high nutritive value providing significant amounts of protein, ω3 PUFA, phosphatidylcholine, carotenoids, and tocopherols, with fatty acids profiles that exhibit low atherogenic and thrombogenic potential.

Wheat germ: not only a by-product

The wheat germ (embryonic axis and scutellum) represents about 2.5–3.8% of total seed weight and is an important by-product of the flour milling industry. The germ contains about 10–15% lipids, 26–35% proteins, 17% sugars, 1.5–4.5% fibre and 4% minerals, as well as significant quantities of bioactive compounds such as tocopherols [300–740 mg/kg dry matter (DM)], phytosterols (24–50 mg/kg), policosanols (10 mg/kg), carotenoids (4–38 mg/kg), thiamin (15–23 mg/kg) and riboflavin (6–10 mg/kg). Oil recovery is achieved by mechanical pressing or solvent extraction, which retrieve about 50% or 90% lipids, respectively; innovative approaches, such as supercritical carbon dioxide extraction, are also proposed. The oil is rich in triglycerides (57% of total lipids), mainly linoleic (18:2), palmitic (16:0) and oleic (18:1) acids, but relevant amounts of sterols, mono- and diglycerides, phospho- and glycolipids are present. The lypophilic antioxidants tocopherols and carotenoids are also abundant. The main by-product of oil extraction is defatted germ meal, which has high protein content (30–32%), is rich in albumin (34.5% of total protein) and globulin (15.6%), and thus presents a well-balanced amino acid profile. Its principal mineral constituents are potassium, magnesium, calcium, zinc and manganese, in decreasing order. Total flavonoid content is about 0.35 g rutin equivalent/100 g DM. The wheat germ is therefore a unique source of concentrated nutrients, highly valued as food supplement. While the oil is widely appreciated for its pharmaceutical and nutritional value, the defatted germ meal is a promising source of high-quality vegetable proteins. Better nutrient separation from the kernel and improved fractioning techniques could also provide high-purity molecules with positive health benefits.

An ultra performance liquid chromatographic method for determining phytosterol uptake by Caco-2 cells

A simple method for the determination of cellular uptake of phytosterols by Caco-2 cells has been developed by ultra performance liquid chromatography with ultraviolet detection (UPLC–UV). UPLC–UV was established using an ODS column, acetonitrile/H 2O (9:1, v/v) as a mobile phase, and a detection wavelength at 210 nm. As analytes, β-sitosterol, campesterol, stigmasterol, and brassicasterol were selected based on the abundance in foods and the similarity of their structures. A linear relation was observed between the peak area and the amount of sterol injected from 50 to 2000 pmol ( r > 0.999) with a relative standard deviation (RSD) of less than 2.5% ( n = 6). This method was applied to the determination of cellular uptake of phytosterols by Caco-2 cells. Recovery tests showed that phytosterols were extracted from the cell lysates by chloroform and determined by UPLC–UV with a recovery rate of more than 80.2% and an RSD of less than 11.3% ( n = 3). When Caco-2 cells were incubated with phytosterols at 37 °C, their uptake was increased with time in a concentration-dependent manner. This method will be useful for the simultaneous determination of cellular phytosterols in an in vitro intestine model.

Bioactive compounds in unsaponifiable fraction of oils from unconventional sources

AbstractThe aim of the research was to characterize bioactive components of unsaponifiable fraction of selected unconventional oils. Nine oils were analyzed as far as the content of tocopherols, squalene, phenolic compounds, and sterols were concerned. Tocopherols and squalene were analyzed by HPLC coupled with diode array detector and fluorescent detector (HPLC‐DAD‐FLD). The content of sterols in oils was determined by GC coupled with MS (GC‐MS). The total amount of phenolic compounds in oils was determined by the colorimetric methods using Folin–Ciocalteau phenol reagent. The examined oils were characterized by differentiated amount of particular forms of tocopherols. The oil obtained from the seeds of amaranth was the richest source of squalene (over 52 mg/g oil). The presence of 22 different compounds of sterols were identified, whereas β‐sitosterol was found in the largest amount. Total amount of sterols in the oils ranged from 90 (walnut) to 850 mg/100 g (evening primrose). Significant differentiation of total amount of phenolic compounds was observed in the examined oils. Evening primrose oil showed the highest amount of phenolic compounds (679 mg/kg). The presented results prove that plant oils obtained from nonconventional sources are a potential source of bioactive compounds.

CHEMICAL EVALUATION OF FLOWER BUD AND OILS OF TUMBLEWEED (GUNDELIA TOURNEFORTI L.) AS A NEW POTENTIAL NUTRITION SOURCES

ABSTRACT The chemical composition and mineral contents of flower bud of tumbleweed (Gundelia tourneforti L.) wild growing in Turkey were studied. The tumble weed flower buds were rich in crude oil, crude protein and crude fiber. At the same time, it proved to be a good source for minerals K, Ca, P, Na, Fe, Mg, Zn. The oil of flower buds were rich in linoleic (57.8%), followed by oleic (28.5%) and palmitic (8.1%). Stearic, vacsenic and arachidic were also found. The total content of sterol of the oil was established as 3,766.60 mg/kg, with β-sitosterol as the predominant sterol that accounted for more than 51.76% of the total amount of sterols. Other sterols were established as 18.52% stigmasterol, 9.82% 5-avenasterol, 6.02% campesterol, 3.68% 7-stigmastenol and 2.63% 7-avenasterol. The total content of vitamin E active compounds in the oil was determined as 51.9 mg/100 g. The predominant isomers were α-tocopherol (48.9 mg/100 g) and γ-tocopherol (1.0 mg/100 g). PRACTICAL APPLICATION The head part of Gundelia tournefortii is composed of several groups of flowers. The gundelia is an edible spiny, thistle-like flowering plant. The edible parts are the leaves, stems, roots, and undeveloped flower buds. The soft white part of the young leaves is used in soup. The flowers, leaves, seeds and stems of Tumbleweed are consumed as food sources. At the same time, it is an important food source and a well-known medicinal plant in Eastern Anatolia.

STEROL PROFILES IN 18 MACROALGAE OF THE PORTUGUESE COAST(1).

The sterol profiles of dominant macroalgae occurring in the western Portuguese coast were evaluated. An analytical procedure, involving alkaline hydrolysis and extraction followed by separation by reversed-phase HPLC-diode array detection (HPLC-DAD), was optimized for the study of their sterols composition. The validated methodology is short in analysis time (as the compounds are determined in <20 min), sensitive, reproducible, and accurate. It was then successfully applied to the determination of campesterol, cholesterol, desmosterol, ergosterol, fucosterol, stigmasterol, and β-sitosterol in 18 species (three Chlorophyta, five Rhodophyta, and 10 Phaeophyta). The profiles obtained for the several macroalgal species were considerably different. C29 sterols were predominant in Phaeophyta and Chlorophyta (71%-95% of total sterol content), while in Rhodophyta cholesterol content is significantly higher (34%-87%). Among the studied species, Asparagopsis armata Harv. contained the lowest sterol amount (555 mg · kg(-1) dry weight), and Cystoseira tamariscifolia (Huds.) Papenf. the highest one (6,502 mg · kg(-1) dry weight). Data obtained may be helpful in identifying suitable marine sources of sterols, with potential applications in the food and pharmaceutical industries.

THE INFLUENCE OF CONTROLLED ATMOSPHERE STORAGE ON PHYTOSTEROLS IN BROCCOLI HEADS

The objective of this study was to analyze changes in sterols content during storage of broccoli. Broccoli heads of “Marathon” and “Cezar” cultivars were stored for 6 weeks in a cold store, under normal (0% CO2 + 21% O2) and controlled atmosphere conditions (CA, 5% CO2 + 2% O2) at the temperature of 0–1C. Sterols, in the forms of free sterols and glucosides, were determined by high-performance liquid chromatography. Color parameters of broccoli were determined spectrophotometrically using CIE L*a*b* system. Amount of free sterols in the freshly harvested heads reached 54.3 mg/100 g dw and was the highest in the heads of “Marathon.” The dominant free sterol was b-sitosterol. During the storage of broccoli, sterol content decreased but was higher in the heads stored in CA. The broccoli showed smaller changes of color when stored in CA than in normal atmosphere.

Quantitative role of LAL, NPC2, and NPC1 in lysosomal cholesterol processing defined by genetic and pharmacological manipulations

Lipoprotein cholesterol taken up by cells is processed in the endosomal/lysosomal (E/L) compartment by the sequential action of lysosomal acid lipase (LAL), Niemann-Pick C2 (NPC2), and Niemann-Pick C1 (NPC1). Inactivation of NPC2 in mouse caused sequestration of unesterified cholesterol (UC) and expanded the whole animal sterol pool from 2,305 to 4,337 mg/kg. However, this pool increased to 5,408 and 9,480 mg/kg, respectively, when NPC1 or LAL function was absent. The transport defect in mutants lacking NPC2 or NPC1, but not in those lacking LAL, was reversed by cyclodextrin (CD), and the ED₅₀ values for this reversal varied from ~40 mg/kg in kidney to >20,000 mg/kg in brain in both groups. This reversal occurred only with a CD that could interact with UC. Further, a CD that could interact with, but not solubilize, UC still overcame the transport defect. These studies showed that processing and export of sterol from the late E/L compartment was quantitatively different in mice lacking LAL, NPC2, or NPC1 function. In both npc2(-/-) and npc1(-/-) mice, the transport defect was reversed by a CD that interacted with UC, likely at the membrane/bulk-water interface, allowing sterol to move rapidly to the export site of the E/L compartment.

Fatty acids, tocopherol, and sterol contents of some Nigella species seed oil

The lipid compositions of the seed oils of some Nigella species were investigated. The total oil content of the seeds ranged from 28.0 to 36.4%. GC-MS fatty acid compositional analysis of the Nigella seed oils revealed the content of linoleic acid to be the highest (40.3&amp;ndash;58.9%). Other prominent fatty acids were as follows: oleic (18.7&amp;ndash;28.1%), palmitic (10.1&amp;ndash;12.5%), 22:1 D11 (3.2&amp;ndash;3.8%) and stearic (2.6&amp;ndash;3.1%) acids. All the Nigella seed oils analysed exhibited differences in their tocopherol contents and the differences were estimated. The oils extracted from the seeds contained between 1.70&amp;ndash;4.12 mg/100 g &amp;alpha;-T, 0.97&amp;ndash;4.51 mg/100 g &amp;gamma;-T, and 4.90&amp;ndash;17.91 mg/100 g &amp;beta;-T3. The total tocopherol content in seeds varied between 9.15 mg/100 g to 24.65 mg/100 g. The compositions of the sterol fractions were determined by gas liquid chromatography. The total amounts of sterols ranged between 1993.07 mg/kg to 2182.17 mg/kg. The main component was &amp;beta;-sitosterol (48.35&amp;ndash;51.92%), followed by 5-avenasterol, campesterol, and stigmasterol.

Metabolic engineering of monoterpene synthesis in yeast

Terpenoids are one of the largest and most diverse families of natural compounds. They are heavily used in industry, and the trend is toward engineering modified microorganisms that produce high levels of specific terpenoids. Most studies have focused on creating specific heterologous pathways for sesquiterpenes in Escherichia coli or yeast. We subjected the Saccharomyces cerevisiae ERG20 gene (encoding farnesyl diphosphate synthase) to a set of amino acid mutations in the catalytic site at position K197. Mutated strains have been shown to exhibit various growth rate, sterol amount, and monoterpenol-producing capacities. These results are discussed in the context of the potential use of these mutated strains for heterologous expression of monoterpenoid synthases, which was investigated using Ocimum basilicum geraniol synthase. The results obtained with up to 5 mg/L geraniol suggest a major improvement compared with previous available expression systems like Escherichia coli or yeast strains with an unmodified ERG20 gene that respectively delivered amounts in the 10 and 500 µg/L range or even a previously characterized K197E mutation that delivered amounts in the 1 mg/L range.

Affinity of PIP-aquaporins to sterol-enriched domains in plasma membrane of the cells of etiolated pea seedlings

The hypothesis that sterol-enriched domains represent sites of preferred localization of PIP-aquaporins was tested in experiments on plasma membranes isolated from cells of etiolated pea (Pisum sativum L.) seedlings. Plasma membranes were isolated from microsomes by the partition in the aqueous two-phase polymer system and separated into vesicle fractions of different buoyant density by flotation in discontinuous OptiPrep gradient. Two types of plasma membrane preparations were used: one was treated with cold 1% Triton X-100 and the other was not. In untreated preparations, three populations of plasma membrane vesicles were obtained, while in the case of treated preparations, fractions of detergent-resistant membranes (DRM) and solubilized membrane proteins were obtained. In all membrane fractions collected after OptiPrep flotation, the amounts of proteins, sterols, and PIP-aquaporins were determined. The highest sterol content was detected in the membrane fraction with buoyant density 1.098 g/cm3 and in the DRM fraction (1.146 g/cm3). These fractions contained much more PIP-aquaporins than the other ones. Phase state of the lipid bilayer was determined by measuring generalized polarization excitation of fluorescence (GPEX) of laurdan incorporated into the membranes of different fractions. It was revealed that the lipid bilayer of the membranes with density of 1.098 g/cm3 had a higher extent of ordering than that of the fractions with density of ∼1.146 g/cm3. The results indicated that uppermost local concentrations of PIP-aquaporins were associated with tightly packed sterol-enriched domains. Moreover, upon solubilization of plasma membrane with Triton X-100, PIP-aquaporins mainly resided in DRM, thus exhibiting a high affinity to sterols.

Effect of chemical refining steps on the minor and major components of cottonseed oil

During chemical refining steps (neutralization, bleaching and deodorization) of crude cottonseed oil (CSO), minor and major components, namely, tocopherols, sterols and fatty acids were evaluated by HPLC and GLC. HPLC analysis of total tocopherols (T) revealed that loss in these components were 7.12, 12.28 and 18.47 % in neutralized, bleached and deodorized oil respectively. On the other side, quantitative GLC analysis of sterols (as trimethylsilyl derivative, TMS), showed that the losses were 5.6, 8.7 and 16.4 % at the different refining steps respectively. In case of the fatty acids (as methyl esters, ME), as determined by GLC analysis, it was noticed that essential fatty acids (EFA), namely, linoleic and linolenic were nearly kept constant. The total amounts of sterols and tocopherols lost in the deodorized distillates (D D) as well as their patterns were also determined. It was found that total sterols content was 28500 ppm, whereas tocopherols content was 2640 ppm. Sterenes (reference samples) were also used to follow the formation of sterenes during the chemical refining steps of CSO.

Comparison of Supercritical Fluid and Hexane Extraction Methods in Extracting Kenaf (Hibiscus cannabinus) Seed Oil Lipids

AbstractThe objective of this study was to investigate and compare fatty acids, tocopherols and sterols of kenaf seed oil extracted by supercritical carbon dioxide and traditional solvent methods. Fatty acids, tocopherols and sterols were determined in the extracted oils as functions of the pressure (400 bar, 600 bar), temperature (40 °C, 80 °C) and CO2 flow rate (25 g/min) using a 1‐L extraction vessel. Gas chromatography was used to characterize fatty acids and sterols of the obtained oils while tocopherols were quantified by HPLC. No differences were found in the fatty acid compositions of the various oil extracts and the main components were found to be linoleic (38%), oleic (35%), palmitic (20%) and stearic acid (3%). Extraction of tocopherols using high pressure (600 bar/40 °C, 600 bar/80 °C) gave higher total tocopherols (88.20 and 85.57 mg/100 g oil, respectively) when compared with hexane extraction which gave yield of 62.38 mg/100 g oil. Extraction of kenaf seed oil using supercritical fluid extraction at high temperature (80 °C) gave higher amounts of sterols when compared with hexane extraction.

Sterol composition from inflorescences of Hieracium pilosella L.

The fraction of sterol acetates from the inflorescences of &lt;em&gt;Hieracium pilosella&lt;/em&gt; has been isolated in the typical way from petroleum ether extract. By means of the weight method the total amount of sterols was determined (0.2659%). The mixtures of sterol acetates and free sterols were investigated using GC-MS techniques. The occurrence of about 18 sterols has been observed. Cholesterol, cholest-8(14)-en-3b-ol, cholesta-5.7-dien-3b-ol, cholest-7-en-3b-ol, ergosta-5.24-dien-3b-ol, campesterol, stigmasterol, b-sitosterol, fucosterol, 5a-stigmast-7-en-3a-ol were identified. The probable structures of lophenol, isofucosterol, 5a-stigmasta-7.24-dien-3b-ol, lanosta-9(11).24-dien-3b-ol and 24-ethylidene lophenol were stated on the basis of literature data. The last 4 sterols occur in a vestigial quantity, which made its identification impossible. Sitos erol and cholesterol are remarkably dominating sterols in the fraction.

Bioactive lipids and fatty acids profile of Cistanche phelypaea

Oil extracted from the wild plant of Cistanche phelypaea was analyzed for its fatty acid, sterol, hydrocarbon and tocopherol contents. Total lipids (TL) content was 10 g/kg (on dry weight basis). The majority of fatty acids were of the unsaturated type (50.4 % of total fatty acids), while the saturated (mainly palmitic acid) were about 43.2 % of the total fatty acids. Oleic acid was the dominating fatty acid followed by palmitic and linoleic acids. High amounts of sterols were found in the oil with the main component β-sitosterol. Other phytosterols (e.g. stigmasterol, ∆7-avenasterol and ∆5-avenasterol) were present at approximately equal amounts (6–9 % of total sterols). The main identified hydrocarbon compounds were C21, C26 and C32 constituting about 61.2 % of total hydrocarbons. Small amounts of C12, C18 and C22, however, were also detected. Tocopherol levels were high in the oil (3.36 g/kg oil), wherein β-tocopherol was the main component followed by α-isomer. Both tocopherol components comprised more than 87 % of total vitamin E content in the oil. Furthermore, γ- and δ-tocopherols were detected in small amounts in the oil accounting for 14–16 % of the total vitamin E content. Information provided by the present work will be of importance for food applications and chemotaxonomy of Cistanche phelypaea.

Clinical observations, molecular genetic analysis, and treatment of sitosterolemia in infants and children

The clinical observation and treatment of young children with sitosterolemia has rarely been reported. We report clinical, biochemical, and molecular genetic observations and treatment outcomes for five Chinese children from four separate families presenting with sitosterolemia in whom we identified two new (Y329X, G269R) and three known (R446X, N437K, R389H) mutations in the ABCG5 gene. The R389H mutation was found in 50% of alleles. Three of these five patients received cholestyramine therapy with a very good response. However, all patients discontinued this therapy because of poor compliance. Finally, all patients were on ezetimibe therapy and had satisfactory total serum cholesterol levels, though their plant sterol levels were still higher than normal. Another noteworthy finding is that a female infant had a serum cholesterol level of 654 mg/dl at 7 months of age, despite being breast fed (with very tiny amounts of plant sterols) since birth and undergoing 4 months of ezetimibe administration. Although she failed to respond to ezetimibe during this period, she did show improvement when the therapy was started again at 2 years of age. It is possible that another 23-month-old female patient also responded more slowly to ezetimibe treatment than older patients.

Supercritical CO 2 extraction of oil, carotenoids, squalene and sterols from lotus ( Nelumbo nucifera Gaertn) bee pollen

The high-quality oil, abundant in carotenoids, squalene and sterols (mainly consisting of campesterol, stigmasterol, β-sitosterol and β-amyrin), was extracted by supercritical CO 2 from lotus bee pollen for its potential nutraceutical use. The effects of extraction pressure and temperature on the yields and the compositions of the extracts were investigated by using a two-factor central composite rotatable design experiment. ANOVA for response surface model demonstrated that the data were adequately fitted into four polynomial models. The yields of the oil, carotenoids, squalene and sterols were significantly influenced by the experimental variables. It was predicted that maximum oil yield obtained at the extraction pressure of 38.2 MPa and temperature of 49.7 °C contained the maximum amount of carotenoids, squalene and sterols. GC–FID analysis of the fatty acid composition of lotus bee pollen oil showed that polyunsaturated fatty acids accounted for approximately 22% of the total fatty acids.

Estudio de la Desodorización de Aceite de Soja por Simulación

This work presents simulation results of a vegetable oil deodorization process. For simulation purposes, the soybean oil was modelled as a five component mixture: triglyceride, fatty acid, sterol, tocopherol and squalene. Two types of flow, cross and countercurrent, were studied. The process temperature, pressure and contact time were changed, the amount of tocopherol, sterol and squalene in the distillate and the presence of trans compounds in the vegetable oil were analyzed. The results were in agreement with experiment and indicate that countercurrent flow is more convenient from the point of view of quality of the distillate and of the oil, as well as of steam consumption.

SOME FACTORS INFLUENCING FREE STEROLS CONTENT IN BROCCOLI (BRASSICA OLERACEA L. VAR. BOTRYTIS ITALICA PLENCK.)

The aim of this study was to determine free sterol accumulation in edible parts of broccoli cultivars of different vegetation periods, in relation to a part of the plant and growing term. Broccoli cvs. “Chevalier” F1, “Marathon” F1 and “Cezar” were cultivated as a spring, summer or autumn crop. Sterol compounds were determined by high-performance liquid chromatography in the top part (florets) and in the lower part (stems) of broccoli. The florets contained higher amount of sterols than the stems. The dominant sterol, both in florets and stems, were β-sitosterol and brassicasterol, but sterol content differed between the cultivars. Total amount of free sterols in florets reached 230 mg/100 g dw. “Chevalier” was the cultivar of the highest content of sterols in florets and “Cezar,” of the lowest. The concentration of sterols differed for the terms of growing and was the lowest for the spring term. Sterol content changed during florets development and was the lowest at the nonmature stage of the broccoli. Cooking of broccoli increased free sterol content in fresh and dry weights of the edible part. PRACTICAL APPLICATIONS Many authors point out that consumption of broccoli positively influences the health condition of humans. There are data showing high phytosterol content in this vegetable. Phytosterols are very valuable components of the human diet due to high biological activity of these phytochemicals. In this study, the differentiation in free phytosterol accumulation between some standard cultivars of broccoli is shown. The differentiation in free phytosterol concentration in various sections of the edible part of broccoli, as well as the differentiation resulting from the plants' development, was also found as important to a potential consumer.

Influence of class B scavenger receptors on cholesterol flux across the brush border membrane and intestinal absorption

To learn more about how the step of cholesterol uptake into the brush border membrane (BBM) of enterocytes influences overall cholesterol absorption, we measured cholesterol absorption 4 and 24 h after administration of an intragastric bolus of radioactive cholesterol in mice with scavenger receptor class B, type 1 (SR-BI) and/or cluster determinant 36 (CD36) deleted. The cholesterol absorption efficiency is unaltered by deletion of either one or both of the receptors. In vitro determinations of the cholesterol uptake specific activity of the BBM from the mice reveal that the scavenger receptors facilitate cholesterol uptake into the proximal BBM. It follows that cholesterol uptake into the BBM is not normally rate-limiting for the cholesterol absorption process in vivo; a subsequent step, such as NPC1L1-mediated transfer from the BBM into the interior of the enterocyte, is rate-limiting. The absorption of dietary cholesterol after 4 h in mice lacking SR-BI and/or CD36 and fed a high-fat/high-cholesterol diet is delayed to more distal regions of the small intestine. This effect probably arises because ATP binding cassette half transporters G5 and G8-mediated back flux of cholesterol from the BBM to the lumen of the small intestine limits absorption and causes the local cholesterol uptake facilitated by SR-BI and CD36 to become rate-limiting under this dietary condition.