Diet choice, cortisol reactivity, and emotional feeding in socially housed rhesus monkeys

Chronic psychosocial stress produces an array of adverse health consequences that are highly comorbid, including emotional eating, affective disorders, and metabolic syndrome. The consumption of high caloric diets (HCDs) is thought to provide comfort in the face of unrelenting psychosocial stress. Using social subordination in female rhesus monkeys as a model of continual exposure to daily stressors in women, we tested the hypothesis that subordinate females would consume significantly more calories from a HCD compared to dominant females, and this pattern of food intake would be associated with reduced cortisol release and reduced frequency of anxiety-like behaviors. Food intake, parameters of cortisol secretion, and socio-emotional behavior were assessed for 3 weeks during a no choice phase when only a low caloric diet (LCD) was available and during a choice condition when both a LCD and HCD were available. While all animals preferred the HCD, subordinate females consumed significantly more of the HCD than did dominant females. A flattening of the diurnal cortisol rhythm and a greater increase in serum cortisol to an acute social separation occurred during the diet choice condition in all females. Furthermore, the rate of anxiety-like behavior progressively declined during the 3-week choice condition in subordinate but not dominant females. These data provide support for the hypothesis that daily exposure to psychosocial stress increases consumption of calorically dense foods. Furthermore, consumption of HCDs may be a metabolic stressor that synergizes with the psychosocial stress of subordination to further increase the consumption of these diets.     

Dietary fat-induced hyperphagia in rats as a function of fat type and physical form

The influence of dietary fat on food intake and weight gain was assessed by feeding adult female rats diets that differed in the type and form of fat, as well as in the availability of other macro- and micronutrients. Compared to chow-fed controls, the various fat diets increased total food intake by 4% to 27%. Specifically, rats fed chow and a separate source of fat (fat option diet) consumed more fat and total calories, and gained more weight when the fat source was emulsified corn oil rather than pure corn oil or was vegetable shortening rather than corn oil. However, corn oil and shortening had similar effects on caloric intake and weight gain when presented as emulsified gels. Also, pure and emulsified-gel forms of shortening did not differ in their effects on caloric intake and weight gain. Supplementing the vegetable shortening with micronutrients, however, enhanced its hyperphagia-promoting effect. The results of two-choice tests revealed that the rats' preferences for the orosensory properties of the various fat sources did not account for the differential hyperphagias obtained. Rather, it appears that long-term fat selection and caloric intake are influenced primarily by postingestive factors. Fat selection and total intake were determined not only by the fat source itself, but also by the other diet options. That is, rats selected more fat and consumed more calories when chow was the alternative food than when separate sources of carbohydrate and protein were available.     

Effect of dietary macronutrients on food intake, body weight, and tail width in the marsupial S. crassicaudata

The role of dietary fat, as opposed to total energy intake, in the etiology of obesity is controversial. The aim of this study was to determine the effect of macronutrient content, specifically changes in dietary fat on body weight, fat stores, and food intake in S. crassicaudata, a marsupial that stores about 25% of total body fat in its tail. Female animals were divided into three groups (n = 7-9) matched for food intake per gram of body weight. Each group of animals was fed, ad lib an isocaloric diet (1.01 kcal/g), which contained either 10, 20, or 40% of calories from fat. Body weight, food intake, and tail width (an index of body fat stores) were measured daily. Over 21 days, cumulative energy intake was less (p = 0.026) in the 40% fat group compared to the 10% fat group. Despite the differences in food intake, body weight in each group remained stable throughout the study, so that at day 21 there were no differences in the body weights between the three groups. In contrast, tail width increased in the animals who received the 40% fat diet compared to either the 10% (p = 0.016) or 20% (p = 0.001) fat intake groups, whereas there was no significant change in tail width in either of these two groups. These observations indicate that macronutrient composition has a role, independent of total calories in the regulation of food intake and body fat stores, specifically that dietary fat promotes adiposity, independent of total caloric intake.     

Caloric compensation to gastric loads in rats with hypothalamic hyperphagia

Rats with hypothalamic hyperphagia were treated with gastric loads of an amount of calories in the form of glucose solution or vegetable oil, while daily food intake, weight change, and carcass fat content were used as indices of caloric compensation. This treatment revealed: (a) Rats responded to daily loads of either substance by reducing food intake without affecting weight gain. The suppression of food intake persisted as long as loading continued. However, loads of fat produced more variable results than glucose. (b) Loads of either substance did not enhance fat deposit as compared to the control loads of water or liquid paraffin. Finally, (c) there was no relaxation of caloric compensation even during the acute increase of food intake shortly following the placement of lesions in the ventromedial hypothalamus.     

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.     

Dietary self-selection of golden hamsters in response to acute food deprivation and chronic food restriction

Adult male golden hamsters were maintained on either Purine Rat Chow (Chow diet) or a self-selection diet consisting of high-protein chow, pure fat, and pure carbohydrate (Choice diet). In Experiment 1, animals were deprived of food for single periods of up to 48 hr. Animals on the Chow diet did not increase intake at any time after deprivation; animals on the Choice diet selectively increased their consumption of fat-derived calories and increased their total caloric intake during the first 6 hr of refeeding, but not thereafter. The nature of the diet did not influence the rate at which animals regained weight following deprivation. In Experiment 2, hamsters were placed on food-restriction schedules (access to food either for 1 hr/day only or on alternate days only) until they lost 20% of starting body weight. Chow-fed animals demonstrated little or no change in food intake either during or after food restriction. Hamsters on the Choice diet consumed more calories and lost weight more slowly than did chow-fed animals during 1-hr/day feeding; intake of fat-derived calories was elevated during restriction. Choice hamsters increased total caloric intake only towards the end of the alternate-days restriction schedule. Choice hamsters were hyperphagic following both types of food-restriction schedules, but no increased preference for fat-derived calories was observed. Factors influencing food consumption of hamsters in response to deprivation and restriction are discussed.     

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.     

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.     

Sucrose intake as a function of its cost and the cost of chow

The avid consumption of pure carbohydrate solutions, which often results in a distortion of nutrient balance, is generally presumed to be driven by their taste. In the first of two experiments, we examined the effect of consumption cost on rats' intake of three concentrations of sucrose solution (8%, 16%, and 32%) when a nutritionally complete chow was concurrently freely available. In the second experiment, we examined the intake of 24% sucrose solution and chow as the consumption costs of both were varied. Increasing the cost of sucrose resulted in a reduction in the percent calories taken from sucrose; the steepness of the decline in intake with price was inversely related to the sucrose concentration and to the cost of chow. Chow calories were substituted for relatively expensive sucrose calories. An increase in the cost of chow resulted in a reduction in the percent of calories taken from chow and a protein-poor diet. The cost of sucrose did not affect the slope of the chow intake curve, presumably because, despite its sweet taste, sucrose was not a substitute for the protein, fat, and micronutrients in chow. Total caloric intake was conserved in all cases.

Thus, the avid consumption of sucrose solution is curtailed when it is costly; but the degree of change in intake with cost depends on the cost of an alternative food. These results suggest that diet selection involves a comparison not only of the taste and post-ingestive consequences of available foods, but also of the cost of calories and nutrients in the foods. Selection appears to be guided first by caloric requirements and the relative cost of calories, then by nutrient requirements and the relative cost of nutrients, and finally by taste.     

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.     

A role for sweet taste: calorie predictive relations in energy regulation by rats

Animals may use sweet taste to predict the caloric contents of food. Eating sweet noncaloric substances may degrade this predictive relationship, leading to positive energy balance through increased food intake and/or diminished energy expenditure. These experiments were designed to test the hypothesis that experiences that reduce the validity of sweet taste as a predictor of the caloric or nutritive consequences of eating may contribute to deficits in the regulation of energy by reducing the ability of sweet-tasting foods that contain calories to evoke physiological responses that underlie tight regulation. Adult male Sprague-Dawley rats were given differential experience with a sweet taste that either predicted increased caloric content (glucose) or did not predict increased calories (saccharin). We found that reducing the correlation between sweet taste and the caloric content of foods using artificial sweeteners in rats resulted in increased caloric intake, increased body weight, and increased adiposity, as well as diminished caloric compensation and blunted thermic responses to sweet-tasting diets. These results suggest that consumption of products containing artificial sweeteners may lead to increased body weight and obesity by interfering with fundamental homeostatic, physiological processes.     

Effects of dietary sugar and of dietary fat on food intake and body fat content in rats

The long term ingestion of a sugar-rich diet (low fat) caused severe obesity in adult rats. In a separate experiment, the habitual consumption of a fat-rich diet (40% kcal from fat) also caused severe obesity. Severe obesity developed in both groups of animals even though they did not overeat. Voluntary food intake for the sugar-fed rats averaged 28,314 +/- 756 calories/rat per 55 wks which was similar to the value of 28,884 +/- 953 calories/rat per 55 wks for the fat-fed rats. However, both values were lower than that of 32,869 +/- 588 for the control rats eating Purina chow. Despite a lower caloric intake, carcass fat averaged 45 +/- 1% for rats eating the sugar-rich diet and 46 +/- 2% for rats eating the fat-rich diet, but only 33 +/- 2% for rats eating a diet of Purina chow. These results provide evidence that severe obesity can develop in the absence of hyperphagia in animals eating a sugar-rich or fat-rich diet. Finally, a rat model for severe obesity is presented in which carcass fat ranged from 18% (lean) to 61% (severe obesity) using dietary intervention alone at critical stages of the animal's life.     

Flavor-cued modulation of intake in rats: role of familiarity and impact on 24-h intake.

Following training with distinctively flavored nutritive solutions that differ in concentration and thus in caloric value, rats demonstrate flavor-postingestive consequence learning by preferentially consuming one of the flavors in two-bottle tests (both flavors in nutrient-identical solutions.) Experiment 1 investigated whether the relative familiarity of the flavor-nutrient combinations encountered in two-bottle tests contributes to the observed preference. One of the training concentrations (rather than the customary intermediate concentration) was used to present the flavors in testing; thus, one of the flavors was in a familiar context while the other was in an unfamiliar context. The results of two independent trials (rats trained with 1 and 5% sucrose; rats trained with 5 and 40% sucrose) confirmed that two-bottle test preference was not a preference for the familiar flavor-nutrient combination. Experiment 2 examined whether caloric expectancies based upon a previously learned flavor-postingestive consequence association would affect total daily intake. On alternating days, rats consumed 30 mL of dilute (5%) and concentrated (40%) sucrose, each distinctively flavored. When given 30 mL of 22.5% sucrose containing each of the flavors on separate test days, they ate less chow and thus fewer total calories over 24 h when given the flavor previously paired with concentrated sucrose. Experiment 3 replicated the design of Experiment 2 except that fat calories were used instead of sucrose; no significant adjustment of chow intake in extinction tests was noted, even when the number of fat calories used in training was increased (Experiment 4). Thus, rats did not exhibit flavor-cued modulation of chow intake when trained with fat, in contrast to responsivity to flavor cues when trained with sucrose. This differential responding to fat versus carbohydrate calories is consistent with previous observations, in a variety of paradigms, that modulation of caloric intake is less energetically appropriate when ingested foods are high in fat relative to high-carbohydrate foods.     

Effect of intravenous nutrient infusions on food intake in rats

To assess the effect of gut signals on food intake two types of nutrients were infused intravenously for 17.5 hours in 17 hour fed rats. In the first experiment a solution of 25% d-glucose and 4.25% amino acids (Travasol) was infused at levels of 26 and 52 kcal/day for two consecutive four-day periods. During infusion periods, food intake was reduced from saline baseline levels by 18.9 +/- 1.7 and 34.8 +/- 1.8 kcal/day, respectively. This represents an oral intake reduction of approximately 70% of the infused calories. In contrast, food intake was reduced 17.4 +/- 1.7 kcal/day below saline baseline levels when 40 kcal of Nutralipid were infused. The reduction in food intake was only 43% of the lipid calories infused. These results indicate that infusions of glucose and amino acids are more effective than infusion of fats in inhibiting daily food intake, that gut signals associated with absorption of fat provide important satiety signals and that removal of fat from the bloodstream has relatively little effect on daily food intake.     

Food selection changes under stress

Two studies investigate the effect of stress on food choice. Experiment 1 demonstrates experimentally that stress causes changes in food choice away from healthy low fat foods (grapes) to less healthy high fat foods (M&Ms), confirming previous survey research. Experiment 2, a survey study, finds that more females than males report increasing food consumption when stressed. A much larger percentage of those who report increasing their food consumption when stressed (71%) are restrained eaters (i.e., dieters) than are people who undereat or who do not change the amount they eat when stressed (35%). The foods that they report overeating when stressed are foods they normally avoid for weight-loss or health reasons (i.e., highly caloric high fat snack foods). They report eating these foods to feel better. Both studies show that stress not only increases consumption in certain individuals but also shifts their food choice from lower fat to higher fat foods.     

Leptin resistance does not induce hyperphagia in the rat

Leptin has been thought to work as a mediator for body weight control by inhibiting food intake. Leptin, however, cannot prevent obesity induced by a high-fat diet (HFD) probably because of leptin resistance. We investigated daily feeding and weight gain when ordinary chow (OC) was changed to a HFD in male rats. Food intake, by weight, significantly increased the next day, but gradually decreased until at 20 days the HFD intake contained the same calories as consumed by the OC-fed control rats. The reduction in food intake occurred only during the night without change of preference for the HFD, even after leptin resistance had developed. Nonetheless, the HFD-fed rats gained more weight than the controls. From the present experiment, it is concluded that leptin resistance does not induce hyperphagia, and suggested that body weight is not regulated to be constant.     

Can we live longer by eating less? A review of caloric restriction and longevity

Caloric restriction, decreasing caloric intake by 20-30%, was first shown to extend life in rats nearly 80 years ago. Since that time, limiting food intake for longevity has been investigated in species from yeast to humans. In yeast and lower animals, caloric restriction has repeatedly been demonstrated to lengthen the life span. Studies of caloric restriction in non-human primates and in humans are ongoing and initial results suggest prolongation of life as well as prevention of age-related disease. There is also data in rodents suggesting that short term caloric restriction has beneficial effects on fertility. Although caloric restriction has many positive effects on health and longevity, quality of life on a restricted diet as well as the ability to maintain that diet long term are concerns that must be considered in humans.     

Basal metabolic rate and dietary seasonality among Tibetan nomads

The results of 51 overnight measurements of basal metabolic rate (BMR) in a sample of pastoral nomads resident permanently in Phala, Tibet Autonomous Region, China, are reported. Past studies demonstrated a culturally driven seasonality of diet, with very low summer and very high winter caloric intake. The study was designed to test the hypothesis that the ability of Phala nomads to subsist on low caloric intake for several summer months without signs of malnutrition is explained by lower summer BMR. However, BMR measurements of 40 nomads 13–69 years of age during the summer and remeasurement of 11 nomads during the winter of 1993 provide no evidence for low summer BMR to compensate for the low summer caloric intake. BMR in both seasons is within the normal range predicted by international equations. The BMR of males does not differ from that of females, and the BMR of females averages 7% higher than predicted. Anthropometric evidence reveals that the Phala nomads accumulate body fat during the winter. It is inferred that this may buffer the summer period of low intake. The pattern of subcutaneous fat accumulation in winter, moreover, may afford slight improvement in physiological cold insulation during the severe winters as a consequence of depositing winter fat on the trunk rather than on the periphery. Thus, the dietary seasonality in Phala is a stress that elicits fluctuation in fat energy stores but not BMR. © 1996 Wiley-Liss, Inc.     

Hunger and satiety in genetically obese mice (C57BL/6J-ob/ob)

To determine the mechanism for hyperphagia in genetically obese mice (C57BL/6J-ob/ob), several experiments were conducted on the ability of these mice to respond to caloric deficits and surpluses. Presentation of food or sugar reduces subsequent operant licking in both obese and lean mice. When given sugar solutions, evaporated milk, or sweetened non-fat milk, both obese and lean mice reduce food intake to compensate for the calories obtained from the solutions. These findings indicate that genetically obese mice respond normally to caloric surpluses. Obese mice respond to food deprivation (caloric deficit) by increasing subsequent food intake but they do so more slowly than controls.     

Voluntary and involuntary maternal separation in guinea pig pups with mothers required to forage

Lactating guinea pigs required to leave a nest box to obtain food were observed to be separated from their pups almost 10% of the time on Days 13-15 postpartum. The duration of separation was greater in the dark than in the light, and was not different for mothers/litters exposed to low (LFD) versus high (HFD) foraging demand. Subsequently, LFD and HFD pups as well as pups reared under standard laboratory conditions (SLC) all exhibited vocalization and plasma cortisol responses to involuntary maternal separation. However, several effects of rearing were also observed. HFD pups vocalized less during separation and had higher resting cortisol levels than did SLC pups; LFD and HFD males spent less time in contact with the mother in a test cage than did SLC males. These effects are discussed in terms of the importance of the physical environment in shaping infant-mother interactions and the infant's perception of control.