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© 2004 The American Society for Nutritional Sciences J. Nutr. 134:2263-2269, September 2004

Nutrient Metabolism

Cis-9, Trans-11 and Trans-10, Cis-12 Conjugated Linoleic Acid Isomers Do Not Modify Body Composition in Adult Sedentary or Exercised Rats1

Philippe Patureau Mirand2, Marie-Agnès Arnal-Bagnard, Laurent Mosoni, Yannick Faulconnier*, Jean-Michel Chardigny{dagger} and Yves Chilliard*

Unité Nutrition et Métabolisme Protéique and * Unité Recherches sur les Herbivores, Centre Inra de Clermont-Ferrand-Theix, 63122 Theix, France and {dagger} Unité Nutrition Lipidique, Centre Inra de Dijon, 21065 Dijon, France

2To whom correspondence should be addressed. E-mail: patureau{at}clermont.inra.fr.


   ABSTRACT
TOP
 ABSTRACT
MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 LITERATURE CITED
 
Dietary CLA isomers were shown to reduce adipose tissues in growing animals, mainly in mice, but their effects in adult animals remain unclear. This study was conducted to determine whether these effects depend on the isomer fed, on physical activity, or on the initial level of body fat. Male Wistar rats (4 mo old) were fed for 6 wk diets containing either no CLA, the cis-9, trans-11 CLA isomer (10 g/kg), the trans-10, cis-12 CLA isomer (10 g/kg), or both isomers (10 g/kg each). Half of the rats were assigned to exercise by treadmill running (1 h/d, 22 m/min). The initial body fat level was normal (12.7%) in a first trial, and high (18.9%) in a second trial. Chemical and anatomical body compositions were determined by chemical analysis and organ dissection. In both trials, the CLA diets, whatever the isomer, had no effect on food intake and body weight changes, on body chemical composition (fat, protein and water contents or gains), or on the body anatomical composition (weights or gains in epididymal and perirenal adipose tissues, in liver and in 4 muscles). There was no interaction between CLA treatment and physical activity. In conclusion, adult male rats do not appear to be responsive to the fat-to-lean partitioning effect of CLA described in growing rats. This was not affected by exercise or initial body fat level.

KEY WORDS: • CLA isomers • exercise • body composition • rats

The consequences of feeding conjugated linoleic acid isomers (CLA) on whole-body composition appear to be unclear, probably because they depend on numerous factors related either to the subjects (genotype, age, gender, physiologic and nutritional status), or to the treatment (isomers, level of feeding, duration of treatment). In mice, since the first experiment by Park et al. in 1997 (1), consistent data indicate that dietary CLA decreases adiposity in growing animals (2–6) and in adults (7–10), both male and female (2–5). CLA feeding was described as slightly increasing lean body mass in growing mice (11). The fat-reducing effect was associated with the trans-10, cis-12 isomer of CLA (5,12), whereas the cis-9, trans-11 had a growth-promoting effect (10).

Less consistent effects were described in rats, hamsters, pigs, and humans. In growing female rats, CLA significantly decreased adipose tissues (13–15). In growing male rats, the reported effects were less constant, i.e., a significant decrease was reported for retroperitoneal pad (15,16) or perirenal pad (17–19) but not for epididymal pad (16–18). CLA feeding did not affect fat pad weights in obese Zucker male rats (16) or in growing male rats (14), but soleus weight was increased in the latter study (14). In growing hamsters, a CLA isomer mixture had a slight fat-reducing effect without effect on protein mass (20); it could also prevent an increase in adiposity, whereas the cis-9, trans-11 isomer increased it (21). In growing-finishing pigs, CLA intake reduced adiposity linearly (22) or quadratically (23,24). In this case, the maximum effect was obtained with different doses according to age: 5 g/kg food during the first 4 wk of the growing-finishing period and 2.5 g during the last 4 wk. This may explain why other studies did not detect any effect of CLA feeding on body composition (25,26). In humans, conflicting results were described. In a recent review (27), 7 studies conducted among overweight subjects were compiled. In 3 studies, CLA feeding reduced the body fat percentage and the sagittal abdominal diameter, and it had no significant effect in the other 4 studies. Recently, CLA effects on weight regain and body composition were studied in overweight subjects after weight loss (28). CLA did not prevent weight regain but increased fat-free mass regain. In 2 other studies, the effects of CLA were tested in exercising healthy humans of normal body weight, and conflicting results were described. CLA supplementation reduced the body fat percentage in 1 experiment (29) but had no significant effect on total body mass, fat-free mass, fat mass, percentage of body fat, bone mass, or strength in another experiment (30).

Thus, the specific effects of feeding CLA isomers on body composition in adults and whether these effects might depend on initial fatness or on physical training remain unclear. The present study was designed to determine the effects of CLA intake (cis-9, trans-11 and/or trans-10, cis-12 isomers) on body chemical and anatomical compositions in adult rats and to establish whether physical training or initial body fat level could interact with CLA feeding. Two trials with different initial body fat levels (normal or high) were performed. Half of the animals in each trial were assigned to treadmill exercise to test the possibility that exercise would affect the efficiency of CLA treatments.


   MATERIALS AND METHODS
TOP
 ABSTRACT
 MATERIALS AND METHODS
RESULTS
 DISCUSSION
 LITERATURE CITED
 
    Animals, diets, and exercise. In each trial, male Wistar rats (n = 55) were purchased from Iffa-Credo/Charles River. In trial 1, they were 15-wk-old rats and had been fed a 20% protein diet since weaning (lean rats). In trial 2, the rats were 17 wk old and the protein content of the diet they were fed since weaning was 17.5% (fat rats). They were maintained in individual wire-bottom cages at 21°C with a 12-h light:dark cycle (lights on at 2000 h) and free access to water. The adaptation period lasted 7 d. In each trial, a group of 7 rats (group 0) was killed and dissected at the end of the adaptation period to verify the body composition at the beginning of the experimental period.

The 48 remaining rats were divided into 2 groups. The 24 trained rats were exercised by treadmill running. The rats were progressively adapted during wk 1 to run for an hour at ~22 m/min (i.e., no >50% VO2 max). They were exercised for 6 wk, 6 d/wk during the dark period (at 1500 h). The other group (n = 24) was not assigned to exercise (sedentary rats).

Four diets were tested: the control diet (Control), the control diet with 10 g/kg cis-9, trans-11 CLA (c9,t11), the control diet with 10 g/kg trans-10, cis-12 CLA (t10,c12), and the control diet with the same amounts (10 g/kg) of each isomer (Mixture). The 4 diets had the same basal composition (g/kg): casein 180, cornstarch 430, sucrose 210, cellulose 20, mineral mixture 50, vitamin mixture 10 (31), oil mixture 100. They differed in the composition of the oil mixture (Table 1). The CLA isomers, provided as triglycerides (Natural Lipids), were substituted for high-oleic sunflower oil. The diets were fed in a semiliquid form to allow better control of food intake.


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  		TABLE 1 Composition of the oil mixtures fed to male Wistar rats for 6 wk

 
Each diet was consumed ad libitum for 6 wk by 6 rats in each of the sedentary and exercised groups. During this period, daily food intake was measured 5 d/wk. The rats were weighed 3 times/wk. All procedures were performed according to current legislation on animal experimentation in France.

    Whole-body composition. General anesthesia was induced by i.p. injection of pentobarbital sodium (Sanofi). Liver, digestive tract, heart, 4 skeletal muscles (left and right gastrocnemius, extensor digitorum longus, tibialis anterior, and soleus) and 2 adipose tissues (perirenal and epididymal) were quickly removed. All organs were weighed; the digestive tract was weighed twice, before and after it was emptied. The remaining carcass was frozen and stored at –20°C until determination of its chemical composition. The frozen carcass of each rat was pulverized with a grinder (Robot-Coupe) to obtain a homogenous frozen powder. Two representative samples of each carcass were freeze-dried. Residual water, protein, lipids, and minerals were determined in these samples. Residual water was measured by desiccation at 103°C for 48 h. The nitrogen content was measured by the Kjeldahl method. The protein content was calculated as the nitrogen content x 6.25. Total lipids were determined by the method of Folch (32). Minerals were measured as the ash content after incineration at 500°C for 6 h. Lipids and water in the adipose tissues sampled were added to the value for the carcass. The biochemical composition of the empty body was calculated from the values obtained for carcass and adipose tissues, assuming that the mean composition of liver, blood collected, intestines, and the 4 muscles was not different from whole-body composition. This could not affect the global composition because this group of organs and tissues represented <6% of whole-body weight.

    Calculations and statistical analysis. The empty body weight was determined as the difference between the whole-body weight and that of the digestive contents. The body composition was expressed/100 g empty body weight. The variations in organ weights during the experimental period were calculated from the difference in organ weights measured at the end of the trial and estimated at d 0. Organ weight at d 0 was estimated in all rats by multiplying their respective body weight by the mean organ weight per unit of body weight measured directly in the 7 rats killed on d 0. The biochemical composition of weight gain was calculated from the difference in the amount of constituents measured at the end of the experimental period and that estimated at d 0. Whole-body biochemical composition at d 0 was estimated in all of the rats by multiplying their mean body weight by the empty body composition in the 7 rats killed on d 0. A Student’s t test was performed to determine whether variation in body composition or organ weights during the trial differed from zero or to compare the initial body composition in lean and fat rats. In each trial, data were subjected to a 2-factor ANOVA to detect the effects of diets, exercise, and their interaction. When the effects of the diets were significant (P < 0.05), differences between diets were determined using Fisher’s protected multiple comparison test. Means of main effects (diet and exercise) and pooled SEM are reported. Differences are considered significant when P < 0.05. The StatView statistical software package (version 5 SAS Institute) was used for all of the statistical analyses.


   RESULTS
TOP
 ABSTRACT
 MATERIALS AND METHODS
 RESULTS
DISCUSSION
 LITERATURE CITED
 
    Initial body composition. The fat rats (trial 2), which had an initial body lipid content higher (+48%) than that of the lean rats (trial 1), also had lower water, protein and mineral contents (Table 2). These differences were confirmed by anatomical measurements. The relative weights (expressed as a percentage of empty body weight) of perirenal adipose tissue and liver were higher in the fat group than in the lean group and soleus muscle relative weight was lower. However, epididymal adipose tissue weights did not differ.


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  		TABLE 2 Initial whole-body composition of male Wistar rats fed diets with or without various CLA isomers for 6 wk1

 
    Food intake and growth rates. Two rats died for undetermined reasons in trial 2. One was in the exercised group and was fed the control diet; the other was in the sedentary group and was fed diet t10,c12 (with trans-10, cis-12 isomer). They were withdrawn from the study and not replaced.

In both trials, food intake, weight gains, and food efficiency were not affected by diets but they were significantly lower in the exercised rats than in the sedentary rats (Table 3). There was no significant interaction between diets and exercise for dry matter intake, weight gain, or food efficiency whatever the initial body fat level.


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  		TABLE 3 Effects of conjugated linoleic acid and exercise on food intake, weight gain, and food efficiency in lean and fat rats fed diets with or without various CLA isomers for 6 wk1

 
    Body composition. In both trials, the diets had no influence on water, protein, and lipid contents in whole body (Table 4). Exercised rats had a lower lipid content and higher water and mineral contents than the sedentary rats. There was no significant interaction between diet and exercise on body chemical composition in the lean and the fat rats.


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  		TABLE 4 Effects of conjugated linoleic acid and exercise on whole-body biochemical composition in lean and fat rats fed diets with or without various CLA isomers for 6 wk1

 
In the fat rats, as in the lean rats, the diets did not affect the relative weights of the 2 adipose tissues studied, the liver, or the 4 muscles studied (Table 5). Exercise induced higher relative muscle weights (except in the soleus of the fat rats). On the contrary, the relative weights of perirenal and epididymal adipose tissues were lower in exercised rats than in sedentary rats. Exercise did not affect relative liver weights. There was no significant interaction between diet and exercise on organ weights whatever the initial body fat level.


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  		TABLE 5 Effects of conjugated linoleic acid and exercise on organ weights in lean and fat rats fed diets with or without various CLA isomers for 6 wk1

 
    Weight gain and changes in organ weight during the experimental periods. In the lean and the fat groups, diets did not affect the biochemical composition of weight gains (Table 6) except for minerals. Mineral gains during the experiment were significantly lower in rats fed diet c9,t11 than in those fed the control diet. Exercised rats had lower lipid, water, and protein gains than sedentary rats. Changes in organ weights during the experimental period were not affected by the diets whatever the initial body fat level (Table 7). In both trials, exercise lowered the empty body weight gain as well as the increase in the weight of perirenal and epididymal adipose tissues, and of liver (nonsignificant in fat rats, P = 0.18) but it did not affect the weight changes in gastrocnemius, extensor digitorum longus, and tibialis anterior muscles.


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  		TABLE 6 Effects of conjugated linoleic acid and exercise on the biochemical composition of body weight gain in lean and fat rats fed diets with or without various CLA isomers for 6 wk1

 

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  		TABLE 7 Effects of conjugated linoleic acid and exercise on the variations of empty body and organ weights in lean and fat rats fed diets with or without various CLA isomers for 6 wk1

 

   DISCUSSION
TOP
 ABSTRACT
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
LITERATURE CITED
 
In these experiments, feeding high-fat diets (100 g/kg) with 10 g/kg of cis-9, trans-11, trans-10, cis-12, or both CLA isomers for 6 wk to 4-mo-old rats as triacylglycerols had only minor effects on their body weight changes and on their body composition tested at 2 initial body fat levels and 2 levels of physical activity. To our knowledge, this is the first time that the effects of CLA on body composition obtained in rats > 13 wk old are reported. Moreover, the consequences of the initial fat status or exercise training interaction were tested.

The initial fat status was either normal or high for 16- to 18-wk-old Wistar rats (33). This difference was not the only consequence of the slight age difference (2 wk) between the 2 groups. Indeed the body fat content of the 22-wk-old lean rats (at the end of the experimental period) did not equal that of the 18-wk-old fat rats (at the beginning of this period) despite the high-fat diet they were fed for 6 wk. This allowed us to test the effects of CLA feeding and exercise in the same strain of rats at 2 levels of body fat status due primarily to the diets fed after weaning. Physical exercise in this study (treadmill running at 22 m/min for 1 h) can be considered to be low-intensity endurance training, <55% VO2 max for male rats (34,35). It decreased food intake and weight gain and induced a decrease in body lipid content and adipose tissue weights. Consequently, the lipid and adipose tissue gains were lower in exercised rats than in sedentary rats. On the contrary, body water concentrations and also muscle relative weights were increased by exercise. Furthermore, it can be calculated that the amount of dietary protein not used to increase whole-body protein content was slightly higher (P = 0.05) in exercised rats than in sedentary rats (2.96 ± 0.4 and 3.06 ± 0.4 g/d, respectively). This demonstrates that exercise had no sparing effect on protein in this context of generous protein feeding (165 mg/g food dry matter) and restricted energy (–10% compared with sedentary rats). All of these results are consistent with data from the literature on the consequences of endurance training in rats (36–38).

The effects of CLA supplementation were less marked. Food intake was not affected by CLA feeding. Depressive effects were described mainly in growing mice fed diets supplemented with 5–10 g CLA isomer mixtures/kg (1,2) but not in growing rats whose diets were supplemented with 5 g CLA isomer mixture/kg (13,14,16,19). Body weight changes were not affected by CLA feeding, which agrees with most results obtained in growing rats consuming feed ad libitum (13,14,19). Similarly, no effect of CLA supplementation on body chemical composition could be detected, in keeping with the few studies in which this composition was determined in rats (20,39). CLA supplementation had no effect on anatomical composition either. This is consistent with the literature concerning rats for liver (13,15,16,18,19,39,40), epididymal adipose tissue (14,16–19), and gastrocnemius muscle (13–16,40). For soleus muscle, it was reported that the CLA mixture induced an increase in muscle weight in male rats but not in females (14); however, in other studies (13,15,41) no effect was reported, as is the case in the present experiment. It is more surprising that no effect of CLA isomer feeding was detected on the weight of perirenal adipose tissue because it was reported to be decreased in the 4 studies in which it was measured in rats (17–19,42). Furthermore a lowering effect of CLA feeding was usually described in rats for most other adipose tissues (inguinal, retroperitoneal, parametrial), except for epididymal. In summary, the lack of an effect of CLA on food intake, growth, and CLA-non responsive parameters of body composition (lean body mass, muscles, liver, epididymal adipose tissue) confirms in adult male rats what was already observed in young rats. On the contrary, the decrease in perirenal adipose tissue induced by CLA feeding in young male rats was not found in the present study with adult male rats. It is still unknown whether the same age-related discrepancy exists for other CLA responsive adipose tissues such as inguinal or retroperitoneal tissues. However, it was shown in 11-wk-old female rats (14) and in 26- to 30-wk-old mice (9) that CLA feeding reduced fat pads, as in younger animals. This indicates that adult male mice or female rats are more responsive than adult rats. Concerning potential specific effects of each isomer or of an interaction between both isomers, none could be detected on body composition. This contrasts with data from mice (5,12,43), from obese rats (41), and from hamsters (44) that demonstrated a specific fat-lowering effect of the trans-10, cis-12 isomer. The absence of CLA effect at both initial body fat levels in these male Wistar rats is also different from what was reported in mice and in female Zucker rats. Indeed, a CLA mixture reduced adipose tissue similarly in lean and fat mice (9) and in the lean line of rats, but increased it in the obese line (16). This discrepancy could result from the species difference, from the very high level of body fat in obese Zucker rats, and/or from the specific cause of obesity in these rats. A synergistic effect with exercise was also hypothesized because exercise increases energy losses, thus limiting energy available for fat deposition and fat mass gain. Furthermore, CLA supplementation proved to be effective in restricted subjects such as growing rats (39) and adult dogs (45). This may contribute to a part of its fat-lowering effect in exercising men (29). However, in the present study, CLA feeding had the same consequences on body composition in exercised and in sedentary rats. Globally, the parameters related to food intake, growth rate, and chemical and anatomical composition that were not modified by CLA feeding in young male rats were not affected in adults either. However, 4 main discrepancies were found between this study and data in the literature, including perirenal adipose tissue weight, specific effect of cis-10, trans-12 isomer on body composition, effect of initial fat status, and exercise. It seems unlikely that these discrepancies can be explained by the fact that CLA isomers were fed as triacylglycerols rather than FFA because it was shown in mice that both forms have similar effects on body composition (46). These discrepancies are likely the consequence of age or result from species differences.

In conclusion, this experiment confirms the effects of physical exercise on body composition changes that were already known, but it has not been possible to detect any effect of the 2 major CLA isomers on body fat and body fat-free masses in lean or fat adult rats. The comparison with growing rats suggests that adipose tissues (epididymal excepted) in adult male rats could be less responsive to the fat-reducing effect of the CLA mixture (or of the trans-10, cis-12 isomer). However, this does not mean that CLA isomers had no effect on lipid, protein, or energy metabolisms in adult rats because metabolic rates can be altered without any consequence on body composition; such aspects deserve further studies to elucidate the real effect of these compounds on major metabolic pathways.


   ACKNOWLEDGMENTS
 
Christine Cubizolles is specially acknowledged for animal facilities, Christian Lafarge for animal management, and Danielle Bonin and Hélène Lafarge for help in the literature search.


   FOOTNOTES
 
1 Conducted with financial support from the Commission of the European Communities specific RTD programme "Quality of Life and Management of Living Resources" QLK1-CT99–00076 "Conjugated Linoleic Acid (CLA) in functional food: a potential benefit for overweight middle-aged Europeans (FunCLA)." It does not necessarily reflect its view and in no way anticipates the Commission’s future policy in this area. Back

Manuscript received 8 April 2004. Initial review completed 28 April 2004. Revision accepted 7 June 2004.


   LITERATURE CITED
TOP
 ABSTRACT
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 LITERATURE CITED
 

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