Changes in rats' meal patterns as a function of the caloric density of the diet

Rats in a laboratory foraging paradigm were offered each of four diets which differed in caloric density, and intakes, meal frequencies, meal sizes, and eating rates were monitored. The rats maintained a constant daily caloric intake by eating more frequent, larger meals of the lower density foods. However, caloric meal size was not regulated, and significant correlations between meal size and the length of the post-meal interval were rarely found. The 24-hour pattern of calorie intake was the same regardless of diet. Higher-calorie foods were consumed at a faster rate within meals than were lower-calorie foods. The feeding patterns observed suggest that caloric intake may be regulated over a time frame of several meals rather than on a meal-to-meal basis.     

Taste, intake rate, and food choice in rats

The contribution of taste to the food choices of foraging rats was examined. Rats in a laboratory foraging paradigm searched for sequential opportunities to eat at two feeders containing chow-based food pellets that were plain or flavored with saccharin or citric acid. Pellets cost the same number of bar presses at both feeders. Saccharin adulteration had no effect on intake parameters. Citric-acid-flavored pellets were eaten more slowly and in smaller meals. If there was no alternative food, daily intake was slightly reduced. When the alternative food was plain, fewer meals and fewer pellets were taken of the citric-acid-flavored than plain pellets. When we gradually increased the price of the plain pellets at one feeder, while the price at the alternative feeder (which contained either plain or citric-acid-flavored pellets) remained low, the rate of eating (profitability) decreased at the higher-price feeder, and the rats shifted their intake toward the less-costly, more profitable pellets. We compared the relationship between the relative eating rate at each feeder and the relative meal size (or daily intake) at each feeder when the low-priced food was plain and when it was flavored with citric acid, and found no differences. This indicates that taste may influence choice via its effect on rate of intake.     

Effect of food deprivation and maintenance diet composition on fat preference and acceptance in rats

High-fat diets typically elicit greater kcal intake and/or weight gain than low-fat diets. Palatability, caloric density, and the unique postingestive effects of fat have each been shown to contribute to high-fat diet hyperphagia. Because long-term intake reflects the sum of many individual eating episodes (meals), it is important to investigate factors that may modulate fat intake at a meal. The present studies used high-fat (hi-fat) and high-carbohydrate (hi-carb) liquid diets (both 2.3 kcal/mL) to assess the effect of hunger level (0 versus 24-h food deprivation) and fat content of the maintenance diet (12 versus 48%) on fat preference (when a choice among foods is offered in a two-bottle test), and acceptance (only one food offered) in male rats. Preference for hi-fat relative to hi-carb (two-bottle test) was enhanced by 24-h food deprivation, and by a high-fat maintenance diet. In contrast, neither deprivation nor maintenance diet composition influenced relative meal size (one-bottle test) of hi-fat and hi-carb: irrespective of test conditions, meal size of hi-fat was bigger than meal size of hi-carb.     

Caloric regulation in the rat: control by two factors

These experiments demonstrate that rats can immediately adjust their meal size in response to variations in the caloric density of a novel diet. However, this immediate caloric sensitivity only seems to appear when rats have been adapted to small, calorically insufficient meals. Rats in Experiment 1 were given timed access to unlimited quantities of an oil/water diet during baseline, and they showed no indication of compensating for changes in the caloric density of the oil/water diet during a test meal. Instead, they consumed about the same amount they had consumed during the preceding baseline meal, suggesting that a learned habit of consuming a certain volume of food controlled their meal size. In contrast, rats that were accustomed to receiving only a very small quantity of food for one of their daily meals during baseline immediately responded to the caloric density of an oil/water test diet by consuming a larger meal if the diet was dilute than if it was calorically more concentrated (Experiments 2 and 3). This immediate sensitivity to caloric density occurred whether or not the rats were exposed to the oil/water diet during baseline, suggesting that rats have some way of directly "metering" the caloric density of new foods. Thus, rats' caloric intake during a meal appears to be controlled by two factors: under certain conditions, control is by caloric learning, under other conditions control is by a caloric metering mechanism.     

Macronutrient selection by foraging rats

Rats were studied in a laboratory simulation of foraging for separate sources of carbohydrate, fat, and protein. Fixed ratios of bar-presses were required to search for a cue signalling the availability of a meal and to procure each meal. In the first phase, the costs of procuring all three foods were raised equally for three rats. Meal frequency declined and meal size increased for all three foods. Protein opportunities were accepted more frequently than those for other foods, and the proportion of calories taken from protein remained low and constant. Two rats were studied in the remaining three phases. When protein meal cost was raised while the costs of energy foods were low, protein intake was gradually reduced and at the highest cost was dropped from the diet. There were individual differences in the preferred source of energy calories; when each animal's preferred energy source was made more costly, these preferences persisted over the range of costs studied in one rat, but shifted abruptly in the other rat. Finally, when search costs were raised, the animals became less selective in accepting meal opportunities and composing their diets. These results suggest that diet selection in the rat is flexible and responsive to changes in the costs of acquiring food. This ability is consistent with the foraging behaviors required by omnivorous generalized feeders.     

Testing food‐related inhibitory control to high‐ and low‐calorie food stimuli: Electrophysiological responses to high‐calorie food stimuli predict calorie and carbohydrate intake

Maintaining a healthy diet has important implications for physical and mental health. One factor that may influence diet and food consumption is inhibitory control—the ability to withhold a dominant response in order to correctly respond to environmental demands. We examined how N2 amplitude, an ERP that reflects inhibitory control processes, differed toward high‐ and low‐calorie food stimuli and related to food intake. A total of 159 participants (81 female; M age = 23.5 years; SD = 7.6) completed two food‐based go/no‐go tasks (one with high‐calorie and one with low‐calorie food pictures as no‐go stimuli) while N2 amplitude was recorded. Participants recorded food intake using the Automated Self‐Administered 24‐hour Dietary Recall system. Inhibiting responses toward high‐calorie stimuli elicited a larger (i.e., more negative) no‐go N2 amplitude; inhibiting responses toward low‐calorie stimuli elicited a smaller no‐go N2 amplitude. Participants were more accurate during the high‐calorie than low‐calorie task, but took longer to respond on go trials toward high‐calorie rather than low‐calorie stimuli. When controlling for age, gender, and BMI, larger high‐calorie N2 difference amplitude predicted lower caloric intake (β = 0.17); low‐calorie N2 difference amplitude was not related to caloric intake (β = −0.03). Exploratory analyses revealed larger high‐calorie N2 difference amplitude predicted carbohydrate intake (β = 0.22), but not protein (β = 0.08) or fat (β = 0.11) intake. Results suggest that withholding responses from high‐calorie foods requires increased recruitment of inhibitory control processes, which may be necessary to regulate food consumption, particularly for foods high in calories and carbohydrates.     

Regulation of calorie intake in rats with rostral zona incerta lesions: effects of caloric density or palatability of the diet

Following an initial period of hypophagia, the body weights of rats with zona incerta lesions approached asymptotic levels that were significantly lower than those of controls although daily food intake was not reliably depressed. In spite of persisting deficits in feeding efficiency, the ad libitum food intake of the ZI-lesioned rats (relative to body weight) recovered to preoperative levels and was regulated accurately with respect to the caloric density of the diet. However, rats with ZI lesions were finicky eaters and did not maintain stable calorie intake when given unpalatable diets. The water intake of ZI-lesioned rats remained consistently below that of neurologically intact rats but was normal relative to their own food intake and lowered body weight.     

Energetics of meal patterns in rats

Meal patterns in rats were examined as functions of the caloric density and availability of the diet. Three diets were used, a standard laboratory diet (3.6 kcal/g), a calorically diluted diet (2.7 kcal/g), and a calorically concentrated diet (4.5 kcal/g). After obtaining ad lib measures of meal patterns on each diet availability of food was constrained by requiring the rats to complete fixed ratio requirements of barpresses to obtain access to a meal. On all 3 diets, meal frequency decreased, while meal size and duration increased as functions of the ratio requirement. Under ad lib conditions and low ratio requirements, in comparison to the standard diet, meal frequency was greater on the diluted diet and less on the concentrated diet. Meal size did not vary as a function of diet on low ratio schedules. At high ratio requirements, rats continued to maintain caloric intake on the diluted diet by increasing meal frequency. On the concentrated diet, however, rats maintained intake at high ratio values by decreasing meal size rather than meal frequency. The results indicate that the rat can adopt a variety of strategies to solve the problem of controlling energy intake constant across the daily feeding cycle.     

Effects of dietary caloric density on feeding behavior in Mongolian gerbils (Meriones unguiculatus)

Total daily energy intake, water intake, body weight, and meal patterns were studied in Mongolian gerbils as a function of dietary caloric density. On diets ranging in caloric density from 2.25 kcal/g to 6.09 kcal/g, gerbils consumed an average of 40 kcal per 100 g of body weight per day. In comparison to gerbils fed ground Purina Laboratory Chow (4.2 kcal/g), gerbils presented with diets diluted with nonutritive cellulose increased food intake in proportion to the percentage of cellulose added. Gerbils given diets in which the caloric density was increased by the addition of fat, decreased food intake as a direct function of the added fat. Water intake was increased on the calorically diluted diets, and decreased on the concentrated diets. Body weight did not vary as a function of dietary conditions. On a standard pelleted diet (Noyes), gerbils ate approximately 18 meals a day. Average meal size was 0.4 g. When presented with calorically diluted pellets, gerbils maintained daily energy intake by increasing both meal frequency and meal size. There were no differences in food intake, meal frequency or meal size between the light and dark portions of the 24-hr cycle.     

Inverse relationship between brown fat thermogenesis and meal size: The thermostatic control of food intake revisited

This study examines a new hypothesis whereby heat production from brown fat in response to eating may serve as a feedback signal for satiety. To test this hypothesis, in vitro respiration rate of brown adipose tissue (BAT) was determined in relation to the voluntary caloric intake of the preceding test meal. This relationship was examined as a function of meal composition and of obesity. It was found that in rats fed a high fat diet, as well as in two types of obese rats (VMH and Zucker), respiration rate per 100 mg tissue was significantly reduced, and energy intake of the preceding test meal increased compared to rats receiving a low fat diet or to respective lean rats. These data lend support to a brown fat mediated thermostatic hypothesis for the control of food intake.     

The satiating efficiency of foods

Experiments were undertaken to test the general hypothesis that some foods are more satiating than others, to find a mechanism for their differential satiating efficiencies, and to determine whether certain soups had a high enough satiating efficiency to recommend their addition to a meal as a way of reducing total caloric intake of that meal. In the first experiment it was found that intake of a test meal was lower after a large preload of tomato soup than after a small preload in women, but not in men. However, the total energy intake (soup plus test meal) was no less with meals which included the large soup preload than it was with meals that did not include a preload. Therefore adding a normal portion of tomato soup to a meal would not reduce its total energy intake. We noted the interesting incidential finding that total energy intake (i.e., preload plus test meal) of the meals which contained the larger amount of soup was less than the total energy intake of the meals which contained a combination of crackers, jelly, and juice. In the second experiment we confirmed this finding by showing that when equal weights of tomato soup preloads and a preload of crackers, cheese, and apple juice, which contains more energy, were given, total energy intake was less in meals which included soup. Therefore, substituting tomato soup for a more calorically dense first course could reduce total energy intake of that meal. In the third experiment, the hypothesis suggested by the second was confirmed. Two soups were more satiating than crackers, cheese, and juice. When two calorie levels were used for each preload, it was shown that calorie for calorie, these soups decreased intake of the test meal more than crackers, cheese, and juice. In the fourth experiment we showed that the mechanism for this differential satiating efficiency is not readily attributable to either bulk related factors or fat content. We suggest that the differential satiating efficiencies are related to differences in nutrient dispersion, orosensory cues, or temperature. Finally, reductions in intake were accompanied by reductions in the initial rate of eating and not by increases in the rate of deceleration. This reduction was small but consistent and suggests that foods which are more satiating reduce intake by decreasing desire to eat (i.e., hunger), not by accelerating the onset of meal termination (i.e., satiety). In fact the duration of meals was unaffected by the preloads.     

Fat substitutes promote weight gain in rats consuming high-fat diets

The use of food products designed to mimic the sensory properties of sweet and fat while providing fewer calories has been promoted as a method for reducing food intake and body weight. However, such products may interfere with a learned relationship between the sensory properties of food and the caloric consequences of consuming those foods. In the present experiment, we examined whether use of the fat substitute, olestra, affect energy balance by comparing the effects of consuming high-fat, high-calorie potato chips to the effects of consuming potato chips that sometimes signaled high calories (using high-fat potato chips) and that sometimes signaled lower calories (using nonfat potato chips manufactured with the fat substitute olestra). Food intake, body weight gain and adiposity were greater for rats that consumed both the high-calorie chips and the low-calorie chips with olestra compared to rats that consumed consuming only the high-calorie chips, but only if animals were also consuming a chow diet that was high in fat and calories. However, rats previously exposed to both the high- and low-calorie chips exhibited increased body weight gain, food intake and adiposity when they were subsequently provided with a high fat, high calorie chow diet suggesting that experience with the chips containing olestra affected the ability to predict high calories based on the sensory properties of fat. These results extend the generality of previous findings that interfering with a predictive relationship between sensory properties of foods and calories may contribute to dysregulation of energy balance, overweight and obesity.     

Influence of anterior subdiaphragmatic vagotomy and TPN on rat feeding behavior.

Total parenteral nutrition (TPN) inhibits food intake and feeding behavior. Whether caloric sensory function of the liver contributes to this food intake and feeding behavior regulation via vagal-afferent innervation was tested after performing anterior hepatic vagotomy or sham operation in rats infused with a TPN solution providing 100% of daily energy needs, given continuously for 4 days. Food intake, meal number, size, duration, meal and intermeal sniffs, and eating activity were measured using an automated computerized rat eater meter (ACREM). TPN infusion resulted in a significant decrease of food intake and feeding indexes in both groups. The vagotomized rats showed a significantly higher food consumption, achieved by greater meal frequency, larger meal size, and longer meal duration. Thus, vagotomized rats consumed more than their controls by eating larger meals more often and of longer duration. Data suggest that anterior hepatic vagotomy interrupts hepatic caloric sensory feedback loop, diminishing inhibitory vagal effects on food intake with TPN, leading to an overall increase in food intake.     

Influence of early dietary experience on energy regulation in rats

The environmental mechanisms that have contributed to the rapid increases in overweight and obesity in the US over the past 25-30 years have yet to be fully specified. One hypothesis that has been forwarded is that increased consumption of calories in liquid form may be a contributing factor, since some studies support the idea that caloric compensation is less adequate for liquid calories compared to calories delivered in more solid form. Work from our laboratory using rats has examined the role that differences in diet viscosity may play in altering energy intake and body weight regulation. This work has suggested that when offered diets matched on caloric density, macronutrient and micronutrient composition, and differing only in viscosity, adult rats fail to compensate for calories delivered in low-viscosity form in short-term intake tests. Further, long-term consumption of low-viscosity diets leads to enhanced weight gain in adult rats. In the present studies, we examined whether short- or long-term exposure to varying relationships between viscosity and calories led to altered food intake or body weight regulation in juvenile rats. The results demonstrated that animals given either short- or long-term experience with direct relationships between viscosity and calories (high viscosity, high calorie meals and low viscosity, low calorie meals) did not differ in food intake or body weight gain compared to animals given short- or long-term experience with indirect relationships between viscosity and calories (high viscosity, low calorie meals and low viscosity, high calorie meals). When juvenile rats were given long-term (9 weeks) exposure to a single, high or low viscosity diet supplement, matched on caloric density and differing only in viscosity, there were no effects on body weight gain. However, analysis of body composition using DEXA demonstrated that animals consuming the low viscosity supplements had significantly greater body fat than animals that consumed either the high viscosity supplement or no dietary supplement at all. These differences in body fat persisted for at least 3 months following the cessation of dietary supplements; during this 3-month period, animals previously exposed to the low viscosity supplement also gained significantly more weight than animals previously exposed to the high viscosity supplement. Taken together, the results suggest that consuming calories in low viscosity form may contribute to poor intake regulation over the short-term and to increased adiposity over the long term. When animals experience these diets as juveniles, these effects may persist into adulthood.     

Controlling the intake of calories with the intake of food

The author presents a dietary program for weight reduction based not on the substitution of a low calorie meal for a high calorie meal, but on the pre-meal ingestion of low calorie bulk beverages that are pleasant to taste and space filling, thereby compromising the capacity of the stomach to accommodate much of the higher calorie foods of the regular meals. This method makes any form of fasting unnecessary, as the intake of calories is already limited. Because of this method, few changes need to be made in food selection for regular meals.     

Variety in the diet enhances intake in a meal and contributes to the development of obesity in the rat

Male and female rats were given three palatable, high energy foods either simultaneously or in succession during three 40 min courses. Both simultaneous and successive variety enhanced energy intake compared to the intake of single palatable foods, which was itself enhanced compared to the intake of chow. Rats deprived of food for 24 hr showed a compensatory increase in chow intake (84%) but only a 20% increase in intake in the single palatable food conditions, and no increase in the variety conditions. Male and female rats showed a similar response to variety and deprivation. The effect of variety on body weight was also examined in rats offered either chow, or chow and one palatable food, or chow and three palatable foods in succession (changed every 12 hr) or simultaneously, for seven weeks. All rats offered the palatable foods were hyperphagic compared to chow-fed controls. Rats given the simultaneous but not the successive variety diet were more hyperphagic than the other palatable food groups and showed significantly greater body weight and fat gains. The availability of a variety of foods is an important factor in the amount eaten in the meal and in the etiology of obesity.     

Meal patterns of lean and leptin-deficient obese mice in a simulated foraging environment

C57BL/6J lean and obese (lep -/-) mice were studied in a closed economy operant protocol that simulates foraging. A predetermined number of presses on a procurement lever (PFR) activated a consumatory lever on which presses would produce 20-mg food pellets. Mice could eat as much as they wished but, once no responding occurred for an elapsed 10-min period, the consumatory lever was inactivated and the procurement or foraging cycle began again. Under these conditions, as has been shown for rats and other species, mice initiated relatively discrete meals (about nine per day) at the lowest PFR, and the number of meals initiated per day decreased with increasing PFR. Meal size increased reciprocally, so that total intake was conserved across the range of PFR examined. Obese mice ate larger meals than lean mice at low PFR, and showed further increases but only at the highest PFRs. The small and inconsistent literature on meal patterns in mice is reviewed, and we discuss the utility of the present protocol to study the interactions between genetic and environmental economic factors, and their implications for the etiology of human obesity.     

Sensory stimulation and energy density in the development of satiety

To determine the contribution of sensory stimulation to the changing hedonic response to foods, the effects of consuming very low-calorie and higher calorie versions of soup and jello on the subjective pleasantness of foods were compared. Subjects were 24 normal weight females with low dietary restraint. Half the subjects were given test meals of low- and high-calorie tomato soup and half were given low- and high-calorie orange jello. They rated the pleasantness of the appearance, smell, texture and taste of nine sample foods as well as hunger and stomach fullness before and 2, 20, 40 and 60 min after consumption of the test meal. After the 60 min rating subjects were offered as much as they wanted to eat of a second course of cheese on crackers. Despite differences in the energy density of the test meals, there were no significant differences in the weights of food eaten in either the first or second course and no significant compensation in the second course for the energy differences of the first course. No differential effects of the caloric manipulation were observed in terms of hunger or fullness or in the development, time course and magnitude of sensory-specific satiety. That very low-calorie foods can produce sensory-specific satiety indicates that the sensory properties of foods are important for the changing hedonic response to foods as they are consumed.     

The effect of dehydroepiandrosterone on Zucker rats selected for fat food preference

When allowed to select between macronutrients in a 1-h-a-day meal paradigm, Zucker rats consume 20–80% of their total caloric intake as fat. If they receive an intraperitoneal injection of DHEA 2 h before such a test meal, they consume fewer total calories. The magnitude of this effect on each macronutrient depends upon the animal's initial preference for fat; the higher the initial fat preference, the more profound is the decrease in caloric intake and the more pronounced the effect on fat consumption. Doses as low as 25 mg DHEA/kg body weight are effective. Lean Zucker rats that prefer to consume a high-fat diet have higher epinephrine and dopamine levels in select regions of the hypothalamus known to control food intake. Administration of DHEA to such animals 2 h before decapitation reduces the content of norepinephrine and these monoamines to levels that mimic the values found in the low-fat-preferring animals. It is hypothesized that exogenous DHEA causes the acute release of norepinephrine, epinephrine, and dopamine in select regions of the hypothalamus, and this release causes a decrease in food intake, particularly fat.     

Dietary obesity: differential effects with self-selection and composite diet feeding techniques.

Female rats were given access to casein, cornstarch, and fat in separate cups or a composite diet identical in composition to that chosen by self-selecting rats. After 2 weeks, all rats received granular sucrose ad lib in addition to these other foods. Sucrose availability resulted in increased caloric intake and increased body weight gain in composite-fed rats, but not in self-selectors. Self-selectors maintained caloric homeostasis by decreased consumption of all foods initially, but protein intake recovered to control levels by the third week of treatment.