The role of taste in dietary self-selection in rats

In order to study the role of taste in dietary self-selection, rats were subjected to two degrees of gustatory deafferentation. The chorda tympani was sectioned alone or in combination with the glossopharyngeal nerve and the pharyngeal branch of the vagus. The animals chose from two diets, only one of which contained protein. After surgery, deficits were observed in body weight, food and water intake, and diet selection, proportional to the extent of deafferentation. Some 76% of the animals increased protein and decreased carbohydrate intake, but all rats continued to select a nutritionally balanced diet (Experiment 1). When subjected to a nutritional challenge of intragastric protein or carbohydrate supplementation, the rats compensated for calories and nutrients by selectively adjusting oral intake (Experiment 2). In saccharin preference tests, the preference as well as the total consumption of the test solutions was reduced (Experiment 3). The findings are contrasted to the effects of oral somatosensory deafferentation and are interpreted as a change in preference within the limits of metabolic requirements. The different roles/mechanisms of the two sensory systems in regard to dietary self-selection are discussed.     

Dietary self-selection in intact, ovariectomized, and estradiol-treated female rats

The effects of estrogenic stimulation on diet selection were examined in intact, estrous cycling rats, ovariectomized (OVX) rats, and OVX rats given estradiol benzoate (EB) hormone replacement therapy. In Experiment 1, OVX was associated with the nearly exclusive choice of the more calorically dense diet of a pair of diets varying in the concentration of one of the three basic macronutrients (i.e., fat, carbohydrate, and protein), an effect that was decreased by EB administration. In the second experiment, dietary self-selection was examined in intact, estrous cycling rats given access to an isocaloric diet triplet of fat, carbohydrate (CHO), and protein. Total caloric intake and body weight did not vary across the estrous cycle. However, diet selection did vary. Fat intake increased; CHO and, to a lesser extent, protein intake decreased during estrus. An opposite diet selection occurred during diestrus. In Experiment 3, OVX resulted in progressive increases in CHO and protein intake, with a concurrent decrease in fat consumption. The EB treatment partially reversed this diet selection profile (Experiment 4). These results were confirmed by diet pairs with both naturally occurring and experimentally produced estrogenic stimulation (Experiments 5 and 6). These data are consistent with the findings of previous research demonstrating estrogenic reduction in CHO intake with standard high-CHO commercial diets. In addition, an increase in fat intake during estrogenic stimulation was found.     

Protein selection, food intake, and body composition in response to the amount of dietary protein

Though not universally observed, moderately low-protein diets have been found to increase caloric intake and body fat. It appears that animals overeat in calories in order to obtain more dietary protein. For animals to control protein intake, they must be able to distinguish between two isocaloric diets containing different percentages of protein and make the appropriate dietary selection on the basis of their previous history of protein intake. Experiment 1 examined the 24-h diet selection (5 vs. 35% casein) of Sprague-Dawley rats that had been previously fed diets containing various percentages of dietary protein (5, 10, 20, 35, or 60% casein). Animals fed 5, 10, or 20% dietary protein showed a preference for the higher protein selection diet. In contrast, no significant diet preference was found in animals pre-fed the two higher levels of dietary protein (35 or 60% casein). In this study, daily food intake and body fat of rats fed the low-protein diets (5 and 10% casein) were similar to rats fed the 20% casein diet. Experiment 2 examined the effects of the level of methionine supplementation on rats fed 10% casein. In this study, food intake and body fat were increased by approximately 20% in rats fed 10% casein diets, regardless of the level of methionine supplementation (0.3 vs. 0.15%). Together, the results suggest that the presence of low-protein-induced hyperphagia helps maintain body protein levels in the face of moderately low dietary protein and promotes an increase in the amount of body fat and energy.     

Oral somatosensory factors in dietary self-selection in rats

For studying the mechanisms of the control of protein intake, rats were subjected to partial trigeminal deafferentation. Oral somatosensory afferents from the lower anterior portion of the oral cavity were selectively sectioned, but somatosensation of the other parts of the mouth as well as gustatory and olfactory afferents were left intact. Animals were allowed to choose from two diets, only one of which contained protein. Before deafferentation, the rats chose a constant portion of their daily intake in the form of protein (12.2%; SE = 0.5). After deafferentation, protein intake was all but eliminated at first but was resumed later; carbohydrate intake was reduced. When total intake had recovered to nearly preoperative values, the protein/total-intake ratio remained impaired, with deficits that ranged from seemingly random selection to extreme preferences. The variability of dietary selection between days was considerably higher than before surgery. The discussion considers the involvement of learning processes in homeostasis.     

L-tryptophan injection fails to alter nutrient selection by rats

Nutritional studies on rats given a choice between two diets differing in protein content have led to the proposal that brain 5-HT content regulates protein intake [2]. Pharmacologic studies under similar conditions of dietary self-selection suggest that brain 5-HT controls carbohydrate intake [41]. We tested the effect of elevating brain 5-HT via tryptophan injection (100 mg/kg) on short-term food intake and selection by rats choosing between two diets differing in protein and carbohydrate content. Under these conditions neither total food intake nor protein and carbohydrate selection were affected despite increases of 50% in brain concentrations of 5-HT and 5-HIAA. The effect of Trp administration was selective to serotonin metabolism as brain concentrations of NE, DA and DOPAC were not affected. These results suggest that alterations in brain 5-HT content which may occur following meal ingestion may not be of physiological importance in regulating nutrient intake and selection.     

Feeding behavior in rats fed diets containing medium chain triglyceride.

The effect of dietary medium chain triglyceride (MCT) on short-term food intake was compared with the effect of long chain triglyceride (LCT) in rats. Corn oil and glyceryl tricaprylate were used as LCT and MCT sources, respectively. Rats were given diets containing 200 g MCT/kg diet (MCT diet), 100 g MCT + 100 g LCT/kg diet (ML diet), or 200 g LCT/kg diet (LCT diet) in Experiment 1. Cumulative food intake was determined every h for the first 12 h, then at 2-h intervals thereafter during the subsequent 12 h. As early as 1 h after feeding, cumulative food intake significantly decreased in MCT-fed animals in a dose-dependent fashion. In Experiment 2, rats were given a choice between MCT and LCT diets for 1 h to confirm whether or not the palatability of diets was influenced by dietary fat sources. There was no difference in food intake between the two diets. In Experiment 3, the responsibility of endogenous cholecystokinin (CCK) for the difference in food intake between the two diets was investigated for 6 h by using a CCK-A receptor antagonist, Devazepide (DVZ, 1 mg/kg b. wt.). Food intake in the MCT diet and also in the LCT diet was improved by DVZ. It is concluded that the satiety, but not the palatability, is affected by carbon chain length in dietary triglyceride sources, although the responsibility of endogenous CCK is very small.     

Dietary self-selection and intake of protein and energy is altered by the form of the diets

The technique of dietary self-selection has been widely used to investigate proposed mechanisms of protein and carbohydrate regulation and to substantiate theories relating nutrients, brain chemistry and behaviour. These experiments have produced conflicting results which might be accounted for by differences in experimental variables or in the different techniques used to measure self-selection. The possibility that the sensory qualities of diets might alter dietary self-selection was examined in three groups of rats self-selecting protein and carbohydrate from pairs of isocaloric diets in which the physical form (finely-milled casein/corn starch powder, granular casein/corn starch powder and gelled finely-milled casein/corn starch), but not the nutrient composition of each pair was varied. In addition, since the serotonergic drug fenfluramine has been frequently used to influence dietary selection, the effects of this drug on the selection of protein and carbohydrate from the three diets were also examined. The selection of particular diets and consequently protein and carbohydrate intake was markedly altered by the form of the diet. Furthermore, the effects of fenfluramine on diet selection were also modified significantly by the form of the diet. These results suggest that intake of certain nutrients is not strictly regulated at a particular fixed value and that drug-induced changes of self-selection behaviour must be interpreted cautiously.     

Estrogen reduces total food and carbohydrate intake,but not protein intake,in female rats

Female rats were given simultaneous access to two isocaloric, isocarbohydrate diets that contained 5% or 45% protein. During four consecutive estrous cycles, rats reduced their total food and carbohydrate intake at estrus but maintained protein intake at levels found during the other stages of the estrous cycle. Administration of estradiol benzoate (EB) to ovariectomized rats given the same 5% or 45% protein diet choices also reduced total food and carbohydrate consumption while maintaining protein consumption. By contrast, administration of EB to ovariectomized rats given isocaloric, isonitrogenous diets containing 25% or 70% carbohydrate significantly reduced both total food and carbohydrate intake. The effects of estrogen on food and protein intake reported here are similar to those previously found after administration of drugs that enhance serotoninergic neurotransmission. Thus, the effects of estrogen on nutrient selection may depend, in part, on activation of a serotoninergic mechanism.     

Selection of protein and fat by diabetic rats following separate dilution of the dietary sources

Diabetic and normal rats were allowed to select their diets from separate sources of protein, carbohydrate and fat. Following the determination of baseline intakes, diabetic and normal rats received dietary components in which either the protein (Experiment 1) or fat source (Experiment 2) was diluted by 25% or 50% by the addition of cellulose. Diabetic rats failed to maintain protein intake at both dilution levels, but made up for the loss of protein-derived calories by consuming more fat. Diabetic rats maintained fat intake at both dilution levels. Dietary dilutions had no effect on total caloric intakes or body weight gain of diabetic rats. Diabetic status, measured by fasting plasma glucose levels and urinary glucose excretion rates, also was unaffected by diet dilutions. These data suggest that diabetic rats maintain total caloric intake following dilution of either the protein or fat source of their diets, but defend intake of fat-derived calories more readily than protein-derived calories. Normal rats maintained both protein and fat intake at the 25% but not at the 50% dilution level. These findings are discussed in terms of the ability of diabetic rats to solve the metabolic problems associated with their diabetic condition.     

Meal composition influences subsequent food selection in the young rat

Food selection one-half hour after the ingestion (2 g) of either a protein containing or a protein-free (carbohydrate) diet was studied in young rats. Following a 12 hr fast the rats were allowed 15 minutes to consume the meal (premeal). Thirty minutes later, they had access to two isocaloric diets that differed only in protein and carbohydrate content. During the first hour of ad lib feeding, protein intake and protein concentration selected were lower in the 45% casein prefed group compared to the carbohydrate group. A further reduction in protein intake and protein concentration occurred when the protein content of the premeal was increased to 70% from 45% casein. Rats prefed with the 70% casein diet significantly reduced their daily total food intake (12 hr) compared to either the 45% casein or carbohydrate prefed group. It is concluded that both quantity and composition of food selected is influenced by the composition of the preceding meal.     

Dietary hyperphagia in rats: role of fat, carbohydrate, and energy content

Dietary energy, fat and carbohydrate content were varied to determine the nutritional factors responsible for hyperphagia induced by feeding rats high-fat diets. In the first experiment, rats were fed isoenergetic high-fat or high-carbohydrate diets for 2 weeks. Weight gain and energy intake were lower in rats given the high-fat diet. When some of the rats were switched to a diet that was high in fat, carbohydrate and energy, gram food intake was initially unchanged, resulting in a substantial increase in energy intake and weight gain. Energy intake gradually declined over the 4 weeks following the switch to the high-energy diet. In the second experiment, rats were fed high-fat diets that were either high or low in carbohydrate content and either high or low in energy content (kcal/g). Rats fed a high-fat diet that was high in energy and carbohydrate ate the most energy and gained the most body weight and carcass fat. In the third experiment, rats were fed high-carbohydrate diets varying in fat and cellulose content. Energy intake and body weight gain varied directly as a function of caloric density regardless of the fat or cellulose content of the diets. It is concluded that hyperphagia induced by feeding high-fat diets is not due to the high dietary fat content alone. Rather, high levels of fat, carbohydrate, and energy interact to produce overeating and obesity in rats fed high-fat diets.     

Dietary self-selection vs. complete diet: body weight gain and meal pattern in rats.

Food intake and body weight gain of male adult Wistar rats were examined in two groups of animals. One group (n = 14) was allowed to select its diet from separate sources of protein (casein, 3.1 kcal/g), fat (lard and sunflower oil, 7.9 kcal/g) and carbohydrate (CHO, starch and sucrose, 3.3 kcal/g). Another group (n = 10) received a nutritionally complete diet (3.3 kcal/g). After 2 weeks of adaptation to the diets, body weights and meal patterns were recorded for at least 4 days. The total caloric intake was nearly identical for the two groups of rats. Rats given dietary choice gained less weight over 4 days than rats fed chow and showed reduced feed efficiency. During the 24-h period, self-selecting rats consumed 20.8% of calories as proteins, 21% as fats and 58.2% as CHO. Self-selecting rats ate significantly less calories during the day than did rats given chow. The chow diet consisting of 17.3% calories as protein, 7.6% as fat and 75.1% as CHO. When comparing the self-selecting group nutrient intakes to those of chow-fed group it was observed that 24-h protein calorie intakes were identical in both groups. Fat intake was significantly higher and CHO reduced as compared to chow-fed rats. During the day, CHO intake was higher in self-selecting rats, and fat intake was not significantly reduced. During the night, protein and fat intakes were significantly higher in self-selecting rats, while CHO intake was significantly decreased, particularly in the last periods of the night.(ABSTRACT TRUNCATED AT 250 WORDS)     

The effects of continuous naltrexone infusions on diet preferences are modulated by adaptation to the diets.

Two groups of male rats were placed on a feeding regimen in which a fat/protein diet and a carbohydrate/protein diet were available ad lib. Naltrexone was infused via osmotic minipumps either at the time the diets were introduced or after one week of adaptation to the diets. In rats adapted to the diets, naltrexone caused a decrease in the intakes of fat/protein and carbohydrate/protein diets. Relative preferences for the two diets were generally unchanged. In contrast, when naltrexone was infused at the time of introduction of the diets, a polarization phenomenon was observed: rats tended to consume nearly all of their daily calories from either one diet or the other. Six rats (out of 10) showed a stronger preference for the carbohydrate/protein diet than did any of the saline-treated rats, while 3 showed a stronger preference for the fat/protein diet than did any of the saline-treated rats. Thus, the effect was not diet- or macronutrient-specific. These preferences became significantly less extreme after termination of naltrexone infusions. Conditioned aversions and naltrexone-induced reductions in exploratory behavior are discussed as potential explanations for this polarization effect. These results indicate that naltrexone has differential effects on the development versus the maintenance of diet preferences. Further, they emphasize the importance of examining individual differences as well as baseline preferences in studies on the control of intake and diet selection.     

Intact regulation of protein intake during the development of hypothalamic or genetic obesity in rats

Adult female rats made hyperphagic with bilateral lesions of the medial hypothalamus as well as spontaneously hyperphagic adult female rats of the Zucker strain were studied for their abilities to regulate dietary protein and energy intakes. This was accomplished by permitting both models of animal obesity to select freely between two isocaloric diets containing 10% vs 60% by weight casein (protein) for at least one month. Compared to appropriate control groups, both experimental groups became obese by consuming more of the low protein diet and less of the high protein diet. The overall effect of this selection pattern was an excessive intake of energy in the form of carbohydrate and fat while maintaining control levels of protein intake. These data imply that the stimulus for hyperphagia in both animal models of obesity is some physiological and/or behavioral error in energy regulation but not protein regulation.     

Sucrose-induced hyperphagia and obesity in rats fed a macronutrient self-selection diet

Adult female rats were allowed to self-select their diet from separate sources of fat, protein, and carbohydrate (starch). Other rats were fed a composite diet that matched the nutrient composition chosen by the self-selecting rats (50% fat, 28% protein, 22% carbohydrate) or a low-fat, high-carbohydrate chow diet. Half of the rats in each diet condition were given access to a 32% sucrose solution for 30 days. Sucrose availability increased total caloric intake (approximately 20%) and body weight gain in all three groups compared to control groups not fed the sucrose solution. The selection animals compensated for their sucrose intake by reducing their fat intake, and to a lesser degree, their starch intake; protein intake was the least affected by sucrose availability. The selection rats consumed less sucrose than the chow-fed rats and displayed a smaller increase in weight, relative to controls, than the chow-fed rats. These differences were attributed to the high-fat intake of the selection animals since similar results were obtained with the rats fed the composite diet. In particular, both the selection and composite diets produced mild obesity in the absence of sucrose. The results demonstrate that sucrose-induced overeating and overweight is not an artifact of restraining the diet choices of rats to a pure sugar and a nutritionally complete diet.     

Patterns of nutrient selection in rats with streptozotocin-induced diabetes

The effects of experimental diabetes on energy intake, patterns of nutrient selection, water intake, body weight and body composition were examined in male Sprague-Dawley rats given ground Purina Chow or a dietary self-selection regime. Following adaptation to dietary conditions, a portion of the animals in each diet group were made diabetic by the administration of 45 mg/kg streptozotocin (STZ). The remaining animals in each group served as vehicle-injected controls. STZ reliably produced diabetes in rats on both dietary regimes. Immediately after the induction of diabetes, rats on the self-selection regime increased carbohydrate and protein intakes and decreased fat intake. Approximately three weeks after STZ administration, diabetic rats reduced carbohydrate intake and increased fat intake. Diabetic animals in both diet groups were hyperphagic and polydipsic relative to non-diabetic controls. During the first three weeks following STZ injections, energy and water intakes of diabetic animals in the two dietary conditions were similar. However, after this initial period, energy and water intakes of diabetic rats given the self-selection regime were significantly lower than those of diabetic animals given Purina Chow.     

Effect of melatonin on total food intake and macronutrient choice in rats

Melatonin, a hormone secreted in a rhythmic manner over 24 h mainly by the pineal gland, is used to alleviate the symptoms of jetlag and treat sleeping problems. The objective of the present study was to examine the effects of a 7-h phase-shift from the natural peak of melatonin secretion on total food intake and macronutrient selection. Forty-eight adult Wistar rats of both sexes were divided in three dietary groups, each group offered a simultaneous and different choice of a carbohydrate- and a protein-rich diet. Macronutrient intakes following intraperitoneal administration of four doses of melatonin (3000, 6000, 10 000 and 15 000 pg/ml blood) at dark onset were examined. Melatonin increased short- (4 h postinjection) and long-term (12 h postinjection) nocturnal total food intake in both male and female rats, mainly with the two highest doses. This effect of melatonin was mainly due to a short-term increase of intake across all carbohydrate-rich diet preparations (dextrin/cornstarch, cornstarch, and sucrose/cornstarch) and across genders. This consistent effect of melatonin on the intake of carbohydrate-rich diets with contrasting sensory attributes rules out the possibility that melatonin acts on sensorymotor pathways, thus suggesting that melatonin's effect on food intake is controlled by the carbohydrate content of the diet. In contrast, melatonin could be affecting some sensory or motor processes peculiar to the ingestion of protein since it increased protein-rich diet intake inconsistently across the various preparations (casein, soy isolate, and egg protein) as well as genders. This evidence supports the view that melatonin acts as a time indicator, reinforcing the animals with a “night cue”, and favors predominant carbohydrate intake normally occurring at the beginning of the activity period.     

Effect of diet textural characteristics on the temporal rhythms of feeding in rats

To examine whether the diurnal rhythms of protein-rich and carbohydrate-rich diet ingestion can be altered by presenting the diets in different textural forms, adult male Wistar rats were assigned to two dietary groups. One group received a two-way choice between high-protein powder and high-carbohydrate granular (HPP-HCG) diets. In the other group the textures were reversed [high-protein granular and high-carbohydrate powder (HPG-HCP) diets]. Rats fed HPP-HCG diets selected significantly less protein (kcal) vs. rats fed HPG-HCP diets, during the 24-h and 12-h dark phase and during the 4-h early and late dark phases. Carbohydrate intakes of the two dietary groups were not significantly different. Total caloric intake for the HPG-HCP dietary group was significantly higher than that of the HPP-HCG dietary group during the 24-h and 12-h dark phase. Body weight was significantly lower in rats fed HPP-HCG diets. In conclusion, macronutrient-rich diets presented in different textural forms alter protein-rich diet ingestion and total energy intake.     

Dietary self-selection,activity and carcass composition of rats fed thiouracil

The interaction of dietary self-selection and spontaneous running wheel activity as fat regulating mechanisms was studied in rats treated with thiouracil. Both hypothyroid and euthyroid rats were given the opportunity to either select their diets, engage in running activity or both. Thiouracil treated rats reduced carbohydrate intake while maintaining protein intake levels equal to those of controls. Thiouracil treatment reduced running activity levels when the rats were given the opportunity to select their diets, but did not affect the running levels of rats fed a control diet. All groups given access to running wheels were at the same carcass fat level except the control diet fed, thiouracil treated rats which were at a lower fat level. It was suggested that, at least for hypothyroid rats, the combination of dietary self-selection and running activity is a more efficient mechanism for regulating carcass composition than is either behavior alone.     

Effect of sibutramine on macronutrient selection in male and female rats

Sibutramine, a serotonin-noradrenaline reuptake inhibitor (SNRI), has been shown to be a safe and effective weight-loss drug. The purpose of the present study was to examine whether sibutramine has an effect on macronutrient selection in both female and male rats in addition to total food intake. Wistar rats of both sexes were divided into three groups, and each group was offered a different set of three sensorily contrasting macronutrient-specific diets, each set including carbohydrate-, protein-, and fat-rich diets. Sibutramine (10 mg/kg) was shown to consistently decrease carbohydrate and fat intake at all data points regardless of gender and diet. Intake of carbohydrate differed between male and female rats at 2 h post administration with 2.5 and 5 mg/kg of sibutramine. The effect of sibutramine on protein intake was diet- and gender-specific. All doses of sibutramine decreased total food intake regardless of gender and diet group beginning at 6 h post administration. In conclusion, sibutramine affected macronutrient selection and emphasis on dietary recommendations, as well as appropriate dosage according to gender should be considered during therapy.