7/29/2019 14979585

1/6

BASIC NUTRITIONAL INVESTIGATION

Dietary Fat (Virgin Olive Oil or Sunower Oil) andPhysical Training Interactions on Blood Lipids

in the RatJose L. Quiles, PhD, Jesus R. Huertas, PhD, Julio J. Ochoa, PhD, Maurizio Battino, PhD,

Jose Mataix, PhD, and Mariano Manas, PhDFrom the Institute of Nutrition and Food Technology, Department of Physiology, University of

Granada, Granada, Spain; and the Institute of Biochemistry, Faculty of Medicine,University of Ancona, Ancona, Italy

OBJECTIVE: We investigated whether the intake of virgin olive oil or sunower oil and performance of physical exercise (at different states) affect plasma levels of triacylglycerols, total cholesterol, and fattyacid prole in rats.METHODS: The study was carried out with six groups of male rats subjected for 8 wk to a diet based onvirgin olive oil (three groups) or sunower oil (three groups) as dietary fat. One group for each diet actedas sedentary control; the other two groups ran in a treadmill for 8 wk at 65% of the maximum oxygenconsumption. One group for each diet was killed 24 h after the last bout of exercise and the other waskilled immediately after the exercise performance. Triacylglycerols, total cholesterol, and fatty acidprole were analyzed in plasma. Analysis of variance was used to test differences among groups.RESULTS: Animals fed on virgin olive oil had lower triacylglycerol and cholesterol values. Physicalexercise reduced these parameters with both dietary treatments. Fatty acid prole showed highermonounsaturated fatty acid proportion in virgin olive fed oil animals and a higher -6 polyunsaturatedfatty acid proportion in sunower oil fed animals. Physical exercise reduced the levels of monounsatu-rated fatty acids with both diets and increased the proportions of -3 polyunsaturated fatty acids.CONCLUSIONS: Results from the present study supported the idea that physical exercise and the intake of virgin olive oil are very good ways of reducing plasma triacylglycerols and cholesterol, which is desirablein many pathologic situations. Concerning ndings on fatty acid prole, we had results similar to thoseof other investigators regarding the effect of different sources of dietary fat on plasma. The mostinteresting results came from the effect of physical exercise, with signicant increases in the levels of -3polyunsaturated fatty acids, which may contribute to the antithrombotic state and lower production of proinammatory prostanoids attributed to physical exercise. Nutrition 2003;19:363368. ElsevierScience Inc. 2003

KEY WORDS: olive oil, sunower oil, fatty acids, exercise, rats

INTRODUCTION

The intake of high amounts of fat is a major risk factor in theetiology of cardiovascular disease and cancer, two of the maincauses of death among populations in developed countries. How-ever, it is evident that the fatty acid composition of a particular fatis more important that its absolute concentration regarding thesediseases. 1 Indeed, there is an important number of studies support-ing the positive effects of the intake of sh oil (mainly rich in -3polyunsaturated fatty acids, or PUFAs) and virgin olive oil (mainlymonounsaturated fat) on these diseases. 1 In the same way, manyinvestigators have expressed their concern about the risk of anexcessive intake of saturated fat or edible oils rich in -6 PUFAs(mainly seed oils such as sunower or corn). 2

Physical exercise is widely recommended as a way of reducingweight gain and ameliorating the risk of hyperlipemias and other

deleterious effects of a high fat intake. 3 Thus, exercise trainingdecreases the plasma levels of total cholesterol and low-densitylipoprotein cholesterol, increases the levels of high-density li-poprotein cholesterol, reduces the concentration of triacylglycer-ols, reduces systolic and diastolic blood pressure, and is usuallyrecommended to prevent or ameliorate the effects of cardiovascu-lar diseases. 4 Moreover, considerable epidemiologic evidence hasbeen acquired linking increased physical activity with reducedoccurrence of breast and colon cancers. 5 In the same way, someexperimental studies performed with animals have suggested thatchronic exercise can retard, delay, or prevent the incidence, pro-gression, or metastasis of experimental tumours. 6 Some of thepossible mechanisms supporting these results include reducedbody fat, enhanced gut motility, stimulation of the immune system,and decreased time of exposure to estrogens and other hormones.

We have already demonstrated some of the links betweendietary fat and physical exercise in relation to health. In fact,previous studies have shown that a program of training for 8 wk leads to changes in the lipid prole of liver and skeletal musclemitochondrial membranes in rats. 7 These changes depended on thetype of fat consumed in affecting the functionality of such mem-branes 8 (contributing in some cases to the general benets

This work was supported by CICYT project ALI91-1113-C03-01.

Correspondence to: Jose L. Quiles, PhD, Instituto de Nutricion y Tecno-log a de Alimentos, Universidad de Granada, C/ Ramon y Cajal 4 (Edif.Fray Luis de Granada), 18071 Granada, Spain. E-mail: jlquiles@ugr.es

Nutrition 19:363368, 2003 0899-9007/03/$30.00Elsevier Science Inc., 2003. Printed in the United States. All rights reserved. PII S0899-9007(02)00949-8

7/29/2019 14979585

2/6

attributed to physical exercise) and modifying their susceptibilityto be oxidized by free radicals. 9

This evidence and our previous experience encouraged us toinvestigate whether the combination of physical exercise and theintake of monounsaturated fat (virgin olive oil) has any additionalpositive effect on blood parameters related to lipid metabolism.For the present study we used virgin olive oil and sun ower oil asdietary fats (the most commonly used vegetable oils in Spain) anda program of physical exercise involving a chronic model and achronic plus acute model. We estimated lipid pro le in plasma andthe levels of total cholesterol, serum triacylglycerols, and otherparameters related to physical exercise.

MATERIALS AND METHODS

Experimental Protocol

Male Wistar rats, initially weighing 80 to 90 g, were allocated ingroups of 10 per cage and maintained on a 12-h light/12-h dark-ness cycle, with free access to food and drinking water. The studylasted 9 wk (1 wk for animal selection followed by 8 wk for theexperiment). During the selection week, all rats were fed a non-puri ed diet and subjected to daily sessions on an exercise tread-mill at a speed of 15 m/min for 15 min. The rats were fedsemisynthetic and isoenergetic diets composed of (g/kg of diet):267 casein, 135.3 starch, 453 sucrose, 80 edible oil, 37 mineralsupplement, 10 vitamin supplement, 1.8 cellulose, 0.9 choline, and3 methionine. Three groups of eight rats each received sun oweroil as dietary fat and the other groups (one of eight rats and two of six rats each) r eceived virgin olive oil (the lipid pro le of both oilsare showed in Table I). For each diet one group was made up of sedentary rats (no exercise) and two groups followed a physicalexercise program (as described below). Thus, the groups were asfollow: rats fed virgin olive oil and sedentary (VS; n 8); rats fedvirgin olive oil and exercised (VT; n 6); rats fed virgin olive oiland exercised to exhaustion (VE; n 6); rats fed sun ower oil andsedentary (SS; n 8); rats fed sun ower oil and exercised (ST; n 8), and rats fed sun ower oil and exercised to exhaustion (SE;n 8).

The exercised animals (VT, VE, ST, and SE) underwent train-ing sessions on a horizontal treadmill throughout the 8 wk of theexperiment: for the rst 2 wk, the rats were exercised 5 d/wk, oncea day at a steadily increased rate, until they could run 40 min/d at35 m/min. These conditions, equivalent to 65% to 70% of maxi-mum oxygen consumption, 10 were maintained during the remain-ing 6 wk. The VT and ST groups were killed 24 h after the lastexercise bout. The VE and SE groups performed a nal exercisetest until exhaustion and immediately killed. Intakes for eachgroup were monitored daily.

The rats were killed at the same time of the day in all cases(between 12:00 and 1:00 PM) to avoid circadian uctuations. TheEthical Committee of the Interministerial Commission of Science

and Technology approved the protocols. Animals were handledaccording to the guidelines for care and use of laboratory animalsby the Spanish Society for Laboratory Animal Sciences.

Sample Analysis

The rats were killed by decapitation, blood was collected in tubescoated with ethylene-diaminetetraacetic acid, and plasma was sep-

arated by centrifuge. Triacylglycerols, total cholesterol, glucose,and lactate concentrations were determined in plasma by enzy-matic methods using Boehringer-Mannheim kits (Munich,Germany).

Fatty acid pro le of plasma was measured by gas liquid chro-matography as described by Lepage and Roy. 11 A gas liquidchromatograph (Model HP-5890, Series II, Hewlett Packard, PaloAlto, CA, USA) equipped with a ame ionization detector wasused to analyze fatty acids as methyl esters. Chromatography wasperformed with a 60-m capillary column with 32 mm inner diam-eter and 20 mm thickness impregnated with Sp 2330 FS (SupelcoInc., Bellefonte, Palo Alto, CA, USA). The injector and detectorwere maintained at 250 C and 275 C, respectively; nitrogen wasused as the carrier gas, and the split ratio was 29:1. Temperatureprogramming (for 40 min) was as follows: initial temperature,160 C for 5 min, 6 C/min to 195 C, 4C/min to 220 C, 2C/minto 230 C, hold 12 min, 14 C/min to 160 C.

All chemical products and solvents, of the highest qualityavailable, were acquired from Sigma (St. Louis, MO, USA) andMerck (Darmstadt, Germany). The homologs of coenzyme Q weregifts from Eisai Co. (Tokyo, Japan). Virgin olive oil and sun oweroil were kindly provided by Coosur S.A. (Jaen, Spain).

Statistical Analysis

The results represent the mean and standard error of six (VT andVE) or eight (VS, SS, ST and SE) animals. A two-way analysis of variance was performed for effects of dietary fat and physicalactivity on each variable. Signi cant ( P 0.05) interaction termswere evaluated by Scheffe s F test. Previous to any statisticalanalysis, all variables were checked for normal and homogeneousvariance with Levene s test. When a variable was found not to benormal, it was log-transformed and reanalyzed.

RESULTS

Dietary intake did not vary signi cantly among groups (data notshown). The sedentary animals reached a hig her weig ht gain thandid exercised groups at the end of the study (Fig. 1A). Accordingto the two-way analysis of variance, physical exercise was the onlyfactor responsible for differences in rat weight gain. Sedentaryanimals fe d sunower oil reached the highest levels of triacyl-glycerols (Fig. 1B), and physical exercise (both models) led to alower concentrations of serum triacylglycer ols in th e animals fedwith both diets. Total cholesterol analysis (Fig. 1C) showed thatphysical exercise was responsible for lower levels of that moleculein both dietary groups. However, sedentary animals fed virginolive oil had a lower cholesterol amount than did the sedentaryanimals fed sun ower oil. Physical exercise and the interactionbetween exercise and dietary fat were responsible for the levels of total cholesterol, according to the two-way analysi s of vari ance.

For both dietary treatments, levels of lactate (Fig. 2A) weresimilar and both types of exercise led to higher values, with nodifferences between fats. No d ifferenc es were found between thesedentary groups for glucose (Fig. 2B), and physical exercise ledto higher levels in exhausted rats, with no differences between

diets.In relation to the plasma lipid pro le (Table II), for the satu-rated fatty acids, there were differences among groups only for

TABLE I.

FATTY ACID COMPOSITION OF DIETARY OILS (g/100 g)

Fatty acid Virgin olive oil Sun ower oil

16:0 11.32 7.1916:1( -9) 0.11 0.02

16:1( -7) 0.84 0.1918:0 4.34 4.5118:1( -9) 74.12 32.0818:2( -6) 7.64 54.2618:3( -3) 0.61 0.10

364 Quiles et al. Nutrition Volume 19, Number 4, 2003

7/29/2019 14979585

3/6

stearic acid (18:0), with all animals subjected to physical exercisereaching higher levels (there was no difference between diets). Forall the monounsaturated fatty acids (MUFAs), the highest levelswere found in animals fed virgin olive oil. In addition, physical

exercise led to lower levels of MUFA in both dietary groups.Dietary fat and physical exercise were responsible for thesechanges according to analysis of variance. In relation to the -6

PUFAs, animals fed sun ower oil showed the highest levels.Physical exercise reached higher levels compared with their con-trols for 20:4( -6) in both diets and for total -6 PUFA in animalsfed virgin olive oil. Concerning -3 PUFA, animals from bothdietary groups had similar levels of 22:6( -3), and physical exer-cise led to higher proportions than in sedentary animals. Physicalexercise affected total -3 PUFA in animals fed olive oil. The totalPUFA index showed higher levels in all the animals fed sun oweroil and those subjected to physical exercise (for bot h diets).

The highest 20:4( -6)/22:6( -3) ratio (Fig. 3A) belonged toanimals fed sun ower oil. This ratio was lower i n trained a nimalsthan in controls but not in the exhausted groups. Figure 3B showsthe ratio -6 PUFA to -3 PUFA, with results similar to thosedescribed for the ratio of arachidonic to docosahexaenoic acid, butin this case physical exercise decreased this ratio in exhaustedanimals fed sun ower oil.

DISCUSSION

There is contrasting evidence about the bene ts of speci c dietary

fats and physical exercise on pathologic states such as cardiovas-cular disease and certain types of cancer. The goal of the presentwork was to investigate whether the combination of both factors

FIG. 1. Effects of physical exercise and dietary fat on weight gain, plasmatriacylglycerols, and plasma total cholesterol of rats. Values are means standard error of the mean. For each chart, columns not sharing superscriptletters are signi cantly different ( P 0.05). A two-way analysis of variance was performed to test the effects of fat, exercise, and the inter-action between fat and exercise. Effects were considered signi cant at P 0.05. The P values for this two-way analysis were as follows. For weightgain: fat, 0.454; exercise, 0.000; fat exercise, 0.341. For triacylglycerols:fat, 0.690; exercise, 0.000; fat exercise, 0.015. For total cholesterol: fat,0.741; exercise, 0.000; fat exercise, 0.016. SE, sun ower oil fed animalsand exercised; SS, sun ower oil fed animals and sedentary; ST, virginolive oil fed animals and training exercised; VE, virgin olive oil fedanimals and exercised; VS, virgin olive oil fed animals and sedentary; VT,virgin olive oil fed animals and training exercised.

FIG. 2. Effects of physical exercise and dietary fat on plasma lactate andglucose of rats. Values are means standard error of the mean. For eachchart, columns not sharing superscript letters are signi cantly different ( P 0.05). A two-way analysis of variance was performed to test the effectsof fat, exercise, and the interaction between fat and exercise. Effects areconsidered signi cant at P 0.05. The P values for this two-way analysis

were as follows. For lactate: fat, 0.559; exercise, 0.001; fat exercise,0.359. For glucose: fat, 0.290; exercise, 0.000; fat exercise, 1.000. SE,sun ower oil fed animals and exercised; SS, sun ower oil fed animals andsedentary; ST, virgin olive oil fed animals and training exercised; VE,virgin olive oil fed animals and exercised; VS, virgin olive oil fed animalsand sedentary; VT, virgin olive oil fed animals and training exercised.

Nutrition Volume 19, Number 4, 2003 365Dietary Fat and Exercise Effect on Blood Lipids

7/29/2019 14979585

4/6

(fat and exercise) affects blood lipid metabolism, thus modifyingthe predisposition to these diseases.

Animals adapted to diet, as showed by plasma fatty acid pro-les (animals fed virgin olive oil had a higher proportion of MUFAand the highest -6 PUFA was found in animals fed sun ower oil).Animals also adapted to the training program, as show by thesigni cantly higher plasma triacylglycerols and total cholesterollevels in sedentary animals and by the levels of lactate and glucose.This was also con rmed by the lesser weight gain produced byexercise, which has been well documented in humans andanimals. 3,12

Regarding triacylglycerols, both models of exercise reducedserum levels of this molecule. Others previously reported the sameeffect in rats and humans. 13,14 The hypotriglyceridemic effect maybe secondary to reduced hepatic triacylglycerol secretion and/orenhanced removal of triacylglycerol-rich lipoproteins by the ex-trahepatic tissues, according to Tan et al. 15 Altered activity of lipoprotein lipase, which is found in several tissues includingskeletal muscle, heart, and adipose tissue, also may be involvedbecause it is responsible for the clearance of postprandial chylo-microns and endogenous very low-density lipoprotein. 16 Withrespect to the effect of dietary fat on serum triacylglycerols, wefound that sedentary animals fed sun ower oil had higher levelsthan those fed virgin olive oil. After physical exercise the valuesreached with both diets were the same, but in terms of reduction,sun ower oil reduced triacylglycerols more than virgin olive oilafter exercise. The reduction of serum triacylglycerols after intakeof speci c dietary fats such as sh oil have been described, 17 butthere are no reports describing differences between olive oil andsun ower oil. Our study supported the idea that physical exerciseand the intake of unsaturated oils as the fat source reduce triacyl-glycerols, which is desirable in many pathologic situations. Be-cause sun ower oil produces a higher proportion of triacylglycer-ols in sedentary animals than does virgin olive oil, the practice of physical exercise could be recommended more often if the con-sumed diet is rich in -6 PUFAs.

Many studies,18

but not all,15

have shown that training lowersserum cholesterol in rats, and our data supported this hypothesis.In humans, the type of exercise performed appears to affect the

hypocholesterolemic effect of training. 18 However, in the presentwork both types of exercise led to a reduction in the level of totalcholesterol to the same extent. This means that exhaustive exercisein the VE and SE groups not compound the hypocholesterolemiceffect in comparison with that obtained with the 8-wk training inthe VT and ST groups. Regarding the effects of dietary fat,controversy exists about the effect of olive oil and sun ower oil ontotal cholesterol. Our results showed that, even if a diet enrichedwith olive oil leads to lower levels of serum cholesterol than withsun ower oil, the concentration of cholesterol in both diets may beconsidered normal and the intake of any one of the two diets led tosimilar levels of cholesterol after physical exercise.

Plasma lipid pro le of the rats showed that the proportion of saturated fatty acids (SFA) is barely in uenced by the ingestion of any of the studied fat sources. SFA appears to be a relatively stablefraction within the general composition of plasma which is inaccordance with the ndings of others. 19 In relation to physicalexercise, Rocquelin and Juaneda 20 investigated SFA and foundthat 16:0 but not 18:0 fatty acids decrease in the epididymal fat of trained rats. Our results showed that 18:0 increased in both dietarygroups after physical exercise, as did the total SFA index for thesun ower oil group. Many studies have reported that plasmapositively adapts itself to the consumption of a MUFA-rich diet,demonstrating in its composition the type of diet ingested. 21 Wecon rmed those results in rats fed the diet rich in olive oil. Physicalactivity led to a decrease in all MUFAs in all animals. With regardto the -6 PUFA, the highest levels were obtained in animals fedthe polyunsaturated fat source, as reported by other. 22 Physicalexercise increased arachidonic (20:4 -6) acid in both dietarygroups, as described by Vapaatalo et al. 23 in the plasma of youngsubjects after short-term heavy exercise. In relation to -3 PUFA,both dietary treatments led to similar values, and physical exerciseincreased the amount of these fatty acids.

In the present study, the effects of dietary fat on plasma fattyacid pro les were in agreement with those found by others, 21 butthe most interesting results concern the effect of physical exercise.

It is very dif cult to know whether the changes found among thedifferent fractions of fatty acids after physical exercise are relatedto changes in a particular metabolic function, a higher or lower

TABLE II.

PLASMA FATTY ACID COMPOSITION (g/100 g) OF RATS SUBJECTED TO PHYSICAL EXERCISE AND DIFFERENT DIETARY FATS*

Fatty acid or index

Experimental groups Two-way ANOVA

VS (n 8) VT ( n 6) VE ( n 6) SS (n 8) ST (n 8) SE (n 8) Fat Exercise Fat exercise

16:0 17.4 0.7a

17.1 0.7a

17.3 0.7a

17.1 0.4a

17.1 0.4a

17.5 0.3a

0.132 0.312 0.10318:0 8.7 1.1 a 10.2 0.8 b 11.5 0.9 b 7.1 0.3 a 10.1 0.6 b 9.2 1.3 b 0.132 0.045 0.353Saturated 36.0 5.2 b 30.3 0.5 b 31.7 0.6 b 26.1 0.2 a 29.4 0.5 b 28.9 1.4 b 0.133 0.898 0.029

18:1( -7) 4.5 0.2 c 3.6 0.3 b 3.1 0.1 b 3.4 0.2 b 2.1 0.1 a 2.27 0.18 a 0.000 0.000 0.16918:1( -9) 35.4 1.8 c 32.3 0.3 b 31.8 2.1 b 18.6 0.4 b 15.3 1.1 a 14.6 1.2 a 0.000 0.015 0.421

MUFA 43.8 2.6 d 38.1 1.4 c 35.2 2.2 c 25.7 0.8 b 19.1 1.3 a 20.9 1.6 a 0.000 0.012 0.42318:2( -6) 6.6 0.7 a 7.9 0.4 a 8.8 1.4 a 28.7 1.6 b 29.2 1.1 b 25.1 2.1 b 0.000 0.185 0.30020:4( -6) 10.5 1.2 a 14.1 0.7 b 18.5 1.5 bc 15.3 1.1 b 19.1 1.6 c 20.8 1.7 c 0.011 0.001 0.762

PUFA( -6) 18.6 1.5 a 23.4 1.2 b 28.2 1.7 b 46.9 0.7 c 49.4 1.1 c 48.4 1.1 c 0.000 0.103 0.03622:6( -3) 0.9 0.1 a 1.6 0.3 b 1.5 0.3 b 0.6 0.2 a 1.3 0.4 b 1.3 0.3 b 0.102 0.012 0.617

PUFA( -3) 1.2 0.1 a 1.9 0.4 b 1.9 0.3 b 0.9 0.1 a 1.6 0.2 b 1.6 0.2 b 0.109 0.010 0.123PUFA 19.1 2.8 a 27.2 2.2 b 31.1 2.1 b 48.2 0.9 c 51.5 1.1 d 51.1 1.1 d 0.000 0.002 0.071

* Results are presented as means standard error of the mean. For each fatty acid or index, values in a row not sharing superscript letters are signi -cantly different ( P 0.05). P values for the two-way ANOVA for the effects of fat, exercise, and the interaction between fat and exercise (signi cant at P 0.05).ANOVA, analysis of variance; MUFA, monounsaturated fatty acid; PUFA, polyunsaturated fatty acid; SE, sun ower oil fed animals and exercised; SS,sun ower oil fed animals and sedentary; ST, virgin olive oil fed animals and training exercised; VE, virgin olive oil fed animals and exercised; VS,virgin olive oil fed animals and sedentary; VT, virgin olive oil fed animals and training exercised.

366 Quiles et al. Nutrition Volume 19, Number 4, 2003

7/29/2019 14979585

5/6

disposability of the fatty acids, or if this is due to a higherregulation mechanism. It has been documented that 18:1 -9 andother MUFAs have pr efe rential mobilization and oxidation ratesthan other fatty acids. 21 Leyton et al. 24 demonstrated that ratsoxidize fatty acids in the following order: oleic (18:1 -9)

linoleic (18:2 -6) palmitic (16:0) and stearic (18:0). Similardifferences exist in humans. 25 This could justify the decrease of MUFA in our model after physical exercise because of the higherenergetic requirements it caused. It could be mediated, accordingto Helge et al., 26 by an increase in the solubility of lipids incirculating lipoproteins and, hence, an enhanced enzyme-substratecontact, due to the unsaturation of the lipids. Another potentialmechanism could be related to changes in lipoprotein lipase ac-tivity in heart and skeletal muscle mediated by adaptations todifferent diets, as suggested by Shimomura et al. 27 If this hypoth-esis is true, the increase in some SFAs and PUFAs together witha decrease in MUFAs as a consequence of physical exercise couldrepresent a change in the relative proportions of fatty acids inplasma. That is easy to see if we keep in mind that lipid pro le

results are expressed as relative proportions of the different fattyacids, and a decrease in any of the fractions could lead to a relativeincrease in all the others.

Considerable controversy exists concerning the relative impor-tance of -3 and -6 PUFAs in the prevention of cardiovasculardiseases and cancer. Increase in -3 fatty acids has been related toa decrease in serum triacylglycerols and an increase in high-density lipoprotein cholesterol, whereas increases in linoleic acidand -linoleic acid mainly decrease total cholesterol and low-density lipoprotein cholesterol. 28 In a similar way, high levels of

-3 PUFA may decrease cancer progression and metastasis, andthe opposite is true for -6 PUFA. 29 Eicosapentaenoic acid (20:5 -3) inhibits platelet aggregation induced by arachidonic acidand docosahexaenoic acid (22:6 -3) reduces platelet aggregatingprostanoids (prostaglandins G 2 and H 2 , thromboxane A 2 ).30 Thus,low levels of 20:4 -6/22:6 -3 or, by extension, -6 PUFA/ -3PUFA in plasma re ect a state of antithrombogenesis. In fact, ahigh dietary intake of eicosapentaenoic and docosahexaenoic acidin volunteers reported by decreased plasma triacylglycerols and/orplasma total cholesterol, prolonging bleeding time, and also led toan antithrombotic state. 30 In our study the lower levels of 20:4 -6/22:6 -3 and -6 PUFA/ -3 PUFA were present in animals fedvirgin olive oil. Moreover, regular training reduced these levelswith both dietary fats. Thus, we could say that virgin olive oil isless prothrombotic than sun ower oil and that physical exercisepromotes an antithrombotic state. In addition, an increase in the

ratio of -3 PUFA to -6 PUFA may contribute to a lowerproduction of proin ammatory prostanoids and a higher produc-tion of the anti-in ammatory ones.

In conclusion, under our experimental conditions, physicalexercise and intake of virgin olive oil were very effective inreducing plasma triacylglycerols and cholesterol. Concerning nd-ings on the fatty acid pro le, the most interesting results comefrom the effect of physical exercise, with signi cant increases inthe levels of -3 PUFAs. These effects are very desirable in manypathologic situations.

ACKNOWLEDGMENTS

Special thanks to Ms. Monica Glebocki for elaboration of themanuscript.

REFERENCES

1. Willett WC, Sacks F, Trichopoulou A, et al. Mediterranean diet pyramid: acultural model for healthy eating. Am J Nutr 1995;61:1321S

2. Ramirez-Tortosa MC, Urbano G, Lopez-Jurado M, et al. Extra-virgin more thanrened olive oil increases the resistance of LDL to oxidation in free-living menwith peripheral vascular disease. J Nutr 1999;129:2177

3. Bell RR, Spencer MJ, Sherriff JL. Voluntary exercise and monounsaturatedcanola oil reduce fat gain in mice fed diets high in fat. J Nutr 1997;127:2006

4. Brooks GA, Fahey TD, White TP, Baldwin KM. Exercise physiology. Humanbioenergetics and its applications, 3th ed. Mountain View, California: May eldPublishing Company, 1999

5. Thune I, Furberg AS. Physical activity and cancer risk: dose-response and cancer,all sites and site-speci c. Med Sci Sports Exerc 2001;33:S530

6. Woods JA. Exercise and resistance to neoplasia. Can J Physiol Pharmacol1998;76:581

7. Quiles JL, Huertas JR, Man as M, Battino M, Mataix J. Physical exercise affectsthe lipid pro le of mitochondrial membranes in rats fed with virgin olive oil orsun ower oil. Br J Nutr 1999;81:21

8. Quiles JL, Huertas JR, Man as M, et al. Dietary fat type and regular exercise affectmitochondrial composition and function depending on speci c tissue in rat.J Bioenerg Biomembr 2001;33:127

9. Mataix J, Quiles JL, Huertas JR, Battino M, Man as M. Tissue speci c interac-tions of exercise, dietary fatty acids, and vitamin E in lipid peroxidation. FreeRadic Biol Med 1998;24:511

10. Amstrong RB, Laughlin MH, Rome L, Taylor CL. Metabolism of rats running upand down an incline. J Appl Physiol 1983;55:518

11. Lepage G, Roy CC. Direct transesteri cation of all classes of lipids in one-step

reaction. J Lipid Res 1986;27:11412. Blair SN. Evidence for success of exercise in weight loss and control. Ann Intern

Med 1993;119:702

FIG. 3. Effects of physical exercise and dietary fat on 20:4( -6)/22:6( -3)and PUFA ( -6)/PUFA( -3) indices in plasma of rats. Values are means standard error of the mean. For each chart, columns not sharing super-script letters are signi cantly different ( P 0.05). A two-way analysis of variance was performed to test the effects of fat, exercise, and the inter-action between fat and exercise. Effects were considered signi cant for P

0.05. The P values for this two-way analysis were as follows. For20:4( -6)/22:6( -3): fat, 0.000; exercise, 0.005; fat exercise, 0.349. ForPUFA( -6)/PUFA( -3): fat, 0.000; exercise, 0.015; fat exercise, 0.175.PUFA, polyunsaturated fatty acid; SE, sun ower oil fed animals andexercised; SS, sun ower oil fed animals and sedentary; ST, virgin olive oilfed animals and training exercised; VE, virgin olive oil fed animals andexercised; VS, virgin olive oil fed animals and sedentary; VT, virgin oliveoil fed animals and training exercised.

Nutrition Volume 19, Number 4, 2003 367Dietary Fat and Exercise Effect on Blood Lipids

7/29/2019 14979585

6/6

13. Narayan AK, McMullen JJ, Butler DP, Wake eld T, Calhoun WK. Effect of exercise on tissue lipids and serum lipoproteins of rats fed two levels of fat. J Nutr1975;195:581

14. Costill DL, Cleary P, Fink WJ, et al. Training adaptations in skeletal muscle of juvenile diabetics. Diabetes 1979;28:818

15. Tan MH, Bonen A, Garner JB, Belcastro AN. Physical training in diabetic rats:effect on glucose tolerance and serum lipids. J Appl Physiol 1982;52:1514

16. Deshaies Y, Vallerand AL, Bukowiecki LJ. Serum lipids and cholesterol distri-bution in lipoproteins of exercise-trained female rats fed sucrose. Life Sci

1983;33:7517. Durrington PN, Bhatnagar D, Mackness MI, et al. An omega-3 polyunsaturatedfatty acid concentrate administered for one year decreased triacylglycerols insimvastatin treated patients with coronary heart disease and persisting hypertri-glyceridaemia. Heart 2001;85:544

18. Bjo rtorp P, Berchtold P, Grimby G. Effects of physical training on glucosetolerance plasma insulin and lipids on body composition in men after myocardialinfarction. Acta Med Scand 1972;192:439

19. Rueda P, Lopez-Frias M, Llopis J, Mataix J, Urbano G. In uence of dietary faton plasma fatty acid composition in rats. Nutr Res 1992;12:757

20. Rocquelin G, Juaneda P. In uence of prolonged physical training on the com-position of fatty acids of epididymal adipose tissue and of the carcass in theyoung rat on a dietary regime of sun ower, rapeseed or primor oil. Rep Nutr Dev1981;21:1015

21. Seiquer I, Man as M, Martinez-Victoria E, Ballesta MC, Mataix FJ. Long-terminuence of dietary fat (sun ower oil, olive oil, lard and sh oil) in the serumfatty acid composition and in the different lipidic fractions, in miniature swine.Int J Vitam Nutr Res 1996;66:171

22. Jacotot B, Lasserre M, Mendy F. Effects of different diets rich in polyunsaturatedfatty acids on plasma phospholipids in the human. Progr Lipid Res 1986;25:185

23. Vapaatalo H, Laustiola K, Seppala E, et al. Exercise, ethanol and arachidonic acidmetabolism in healthy men. Biomed Biochem Acta 1984;43:S413

24. Leyton J, Drury PJ, Crawford MA. Differential oxidation of saturated andunsaturated fatty acids in vivo in the rat. Br J Nutr 1987;57:383

25. Jones PJH, Pencharz PB, Clandinin MT. Whole body oxidation of dietary fattyacids: implications for energy utilization. Am J Clin Nutr 1985;42:769

26. Helge JW, Ayre K, Chaunchaiyakul S, et al. Endurance in high-fat-fed rats: effectsof carbohydrate content and fatty acid pro le. J Appl Physiol 1998;85:1342

27. Shimomura Y, Tamura T, Suzuki M. Less body fat accumulation in rats fed asaf ower oil diet than in rats fed a beef tallow diet. J Nutr 1990;120:1291

28. Haglund O, Wallin R, Wretling S, Saldeen T. Effects of sh oil alone andcombined with long chain (n-6) fatty acids on some coronary risk factors in malesubjects. J Nutr Biochem 1998;9:629

29. Lipworth L, Martinez ME, Angell J, Hsieh C-C, Trichopoulos D. Olive oil andhuman cancer: an assessment of the evidence. Prev Med 1997;26:181

30. Masumura S, Furui H, Hashimoto M, Watanabe Y. The effects of season andexercise on the levels of plasma polyunsaturated fatty acids and lipoproteincholesterol in young rats. Biochim Biophys Acta 1992;1125:292

368 Quiles et al. Nutrition Volume 19, Number 4, 2003