The temporal structure of feeding behavior

Meals have long been considered relevant units of feeding behavior. Large data sets of feeding behavior of cattle, pigs, chickens, ducks, turkeys, dolphins, and rats were analyzed with the aims of 1) describing the temporal structure of feeding behavior and 2) developing appropriate methods for estimating meal criteria. Longer (between-meal) intervals were never distributed as the negative exponential assumed by traditional methods, such as log-survivorship analysis, but as a skewed Gaussian, which can be (almost) normalized by log-transformation of interval lengths. Log-transformation can also normalize frequency distributions of within-meal intervals. Meal criteria, i.e., the longest interval considered to occur within meals, can be estimated after fitting models consisting of Gaussian functions alone or of one Weibull and one or more Gaussian functions to the distribution of log-transformed interval lengths. Nonuniform data sets may require disaggregation before this can be achieved. Observations from all species were in conflict with assumptions of random behavior that underlie traditional methods for criteria estimation. Instead, the observed structure of feeding behavior is consistent with 1) a decrease in satiety associated with an increase in the probability of animals starting a meal with time since the last meal and 2) an increase in satiation associated with an increase in the probability of animals ending a meal with the amount of food already consumed. The novel methodology proposed here will avoid biased conclusions from analyses of feeding behavior associated with previous methods and, as demonstrated, can be applied across a range of species to address questions relevant to the control of food intake.     

The role of the stomach in the control of appetite and the secretion of satiation peptides

It is widely accepted that gastric parameters such as gastric distention provide a direct negative feedback signal to inhibit eating; moreover, gastric and intestinal signals have been reported to synergize to promote satiation. However, there are few human data exploring the potential interaction effects of gastric and intestinal signals in the short-term control of appetite and the secretion of satiation peptides. We performed experiments in healthy subjects receiving either a rapid intragastric load or a continuous intraduodenal infusion of glucose or a mixed liquid meal. Intraduodenal infusions (3 kcal/min) were at rates comparable with the duodenal delivery of these nutrients under physiological conditions. Intraduodenal infusions of glucose elicited only weak effects on appetite and the secretion of glucagon-like peptide-1 (GLP-1) and peptide YY (PYY). In contrast, identical amounts of glucose delivered intragastrically markedly suppressed appetite (P < 0.05) paralleled by greatly increased plasma levels of GLP-1 and PYY (≤3-fold, P < 0.05). Administration of the mixed liquid meal showed a comparable phenomenon. In contrast to GLP-1 and PYY, plasma ghrelin was suppressed to a similar degree with both intragastric and intraduodenal nutrients. Our data confirm that the stomach is an important element in the short-term control of appetite and suggest that gastric and intestinal signals interact to mediate early fullness and satiation potentially by increased GLP-1 and PYY secretions.     

Time of day type of food--relation to mood and hunger during 24 hours of constant conditions

A six-day high-carbohydrate meal (HC; 65 E% (energy percent) carbohydrates, 20 E% fat and 15 E% protein) and a six-day high-fat meal (HF; 40 E% carbohydrates, 45 E% fat and 15 E% protein) were given to seven healthy subjects in a crossover design. On the last day subjects were kept awake for 24 hours in a metabolic laboratory while substrate utilisation and energy expenditure were measured by indirect calorimetry. The subjects were given isocaloric meals every four hours. Results showed that hunger decreased at night (F = 4.2, p < 0.05) and linearly increased after meal intake. Macronutrient composition (fat/carbohydrates) seemed to be of less importance for hunger. Hunger and thirst were found to be strongly associated with gastrointestinal substances, for hunger the strongest being a negative correlation with triacylglycerol (partial correlation = -0.39). It is suggested that it might not be necessary for shift workers to eat full portions at night but that satiation will occur with less food. Possibly lack of adjustment of nocturnal food intake might be one reason why overweight is common in shift work populations.     

Increased short-term food satiation and sensitivity to cholecystokinin in neurotrophin-4 knock-in mice

Neurotrophin-4 (NT-4) knockout mice exhibited decreased innervation of the small intestine by vagal intraganglionic laminar endings (IGLEs) and reduced food satiation. Recent findings suggested this innervation was increased in NT-4 knock-in (NT-4KI) mice. Therefore, to further investigate the relationship between intestinal IGLEs and satiation, meal patterns were characterized using solid and liquid diets, and cholecystokinin (CCK) effects on 30-min solid diet intake were examined in NT-4KI and wild-type mice. NT-4KI mice consuming the solid diet exhibited reduced meal size, suggesting increased satiation. However, compensation occurred through increased meal frequency, maintaining daily food intake and body weight gain similar to controls. Mutants fed the liquid diet displayed a decrease in intake rate, again implying increased satiation, but meal duration increased, which led to an increase in meal size. This was compensated for by decreased meal frequency, resulting in similar daily food intake and weight gain as controls. Importantly, these alterations in NT-4KI mice were opposite, or different, from those of NT-4 knockout mice, further supporting the hypothesis that they are specific to vagal afferent signaling. CCK suppressed short-term intake in mutants and controls, but the mutants exhibited larger suppressions at lower doses, implying they were more sensitive to CCK. Moreover, devazepide prevented this suppression, indicating this increased sensitivity was mediated by CCK-1 receptors. These results suggest that the NT-4 gene knock-in, probably involving increased intestinal IGLE innervation, altered short-term feeding, in particular by enhancing satiation and sensitivity to CCK, whereas long-term control of daily intake and body weight was unaffected.     

The amount of food ingested in a single meal by rainbow trout offered chopped herring, dry and wet diets

Two-year-old 1·5-kg rainbow trout were held in cages and conditioned by feeding either on low-fat chopped herring (H trout) or dry pellets (P trout) for 15 weeks. Their satiation amounts were then determined under standard conditions. On a wet weight basis H trout ate 2·5-3·5 times more food than P trout; this was sufficient to compensate for the high water content of herring and thereby maintain the dry matter intake. When P trout were offered herring (PH trout) they consumed more food than when offered dry pellets but not as much as H trout. Stomach capacity restricted the intake and their dry matter intake was reduced by c. 40%. When H trout were offered dry pellets (HP trout) they adjusted their intake immediately close to the level of P trout although their larger stomachs could have accommodated more than twice this volume of dry food. The return of appetite after a satiation meal was almost linear with time. Appetite increased at c. 556 mg g^-1 body weight h^-1 for H trout and at 142 mg g^-1 bw h^-1 for P trout. The return of appetite in PH trout was significantly slower (c. 370 mg g^-1 bw h^-1) than in H trout; the previous dietary history of the PH trout limited their capacity to process larger volumes of wet food in a single meal. Fish offered dry diet (P and HP trout) had similar rates of appetite return despite their previous feeding history suggesting that the property of the dry feed itself might limit meal size. The total gastric emptying time of diets of similar dry matter content (with and without large amounts of water) was similar, but the delay time before gastric emptying starts tended to be longer for dry diets. Dry pellets appear to impose a demand for water that prolongs the gastric delay. This water demand is met partly by drinking since the trout fed on dry pellets drank significantly more (436 ± 189 mg kg^-1 h^-1) than unfed and herring-fed trout which drank little or not at all (65 ± 113 and 70 ± 66 mg kg^-1 h^-1 respectively). Dietary water facilitated food processing and increased daily dry matter intake of trout when fed four times a day. When only one satiation meal per day was allowed, dietary water had no effect. It is concluded from this work that, in addition to gastric volume, a short-term limitation on the size of satiation meals in the rainbow trout is the availability of water to moisturize the food and thus to promote gastric digestion and emptying.     

Estradiol, CCK and satiation.

Estradiol has long been known to inhibit feeding in animals, but the mechanism(s) mediating its effects have not been clear. Demonstrations that estradiol's feeding effects are expressed as decreases in meal size coupled with the emerging consensus that cholecystokinin (CCK) released from the small intestines during meals is a physiological negative-feedback signal controlling meal size (i.e. satiation) suggested a new approach to the problem of the mechanisms of estradiol's inhibitory effect on feeding. Progress on this approach is reviewed here. Experimental manipulations of exogenous and endogenous CCK and estradiol have produced converging evidence that estradiol cyclically increases the activity of the CCK satiation-signaling pathway so that meal size and food intake decrease during the ovulatory or estrus phase of the ovarian cycle. This is a striking example of the modulation of the operation of a control of meal size by the physiological context in which the meal occurs. Estradiol also produces a tonic decrease in meal size, but this apparently does not involve the CCK satiation-signaling pathway. Where and how estradiol acts to increase the potency of the CCK satiating-signaling pathway are not known. Several possible sites are suggested by the observations that estradiol treatment increases feeding- and CCK-induced expression of c-Fos in ovariectomized animals in brain areas including the nucleus tractus solitarius, paraventricular nucleus of the hypothalamus, and central nucleus of the amygdala. Tests with null mutation mice indicate that estrogen receptor-alpha is necessary for estradiol's feeding effects. Finally, the possibilities that estradiol exerts important influences on normal or disordered eating in women are discussed. It is concluded that estradiol exerts a biologically significant action on CCK satiation in animals. Further research to determine whether this action of estradiol has a role in the pathogenesis, course, or treatment of disordered eating in women is indicated.     

Effect of Alginate on Satiation,Appetite, Gastric Function,and Selected Gut Satiety Hormones in Overweight and Obesity

Lack of control of food intake, excess size, and frequency of meals are critical to the development of obesity. The stomach signals satiation postprandially and may play an important role in control of calorie intake. Sodium alginate (based on brown seaweed Laminaria digitata) is currently marketed as a weight loss supplement, but its effects on gastric motor functions and satiation are unknown. We evaluated effects of 10 days treatment with alginate or placebo on gastric functions, satiation, appetite, and gut hormones associated with satiety in overweight or obese adults. We conducted a randomized, 1:1, placebo‐controlled, allocation‐concealed study in 48 overweight or obese participants with excluded psychiatric comorbidity and binge eating disorder. All underwent measurements of gastric emptying (GE), fasting, and postprandial gastric volumes (GVs), postprandial satiation, calorie intake at a free choice meal and selected gut hormones after 1 week of alginate (three capsules vs. matching placebo per day, ingested 30 min before the main meal). Six capsules were ingested with water 30 min before the GE, GV, and satiation tests on days 8–10. There were no treatment group effects on GE or volumes, gut hormones (ghrelin, cholecystokinin (CCK), glucagon‐like peptide‐1 (GLP‐1), peptide YY (PYY)), satiation, total and macronutrient calorie intake at a free choice meal. There was no difference detected in results between obese and overweight patients. Alginate treatment for a period of 10 days showed no effect on gastric motor functions, satiation, appetite, or gut hormones. These results question the use of short‐term alginate treatment for weight loss.     

Uncertainty as wealth

Emotional satisfaction cannot be increased above "normal"-the same normal as the caveman's-for any length of time, but the wealth and consumption style of modern civilization may systematically reduce some people's satisfaction below normal. Hyperbolic discounting of delayed, expected rewards suggests causes for this reduction in humans, and for how we often respond to it, while conventional exponential discounting does not. Hyperbolic discounting has been well demonstrated by four experimental routes; and there is moderate evidence that it motivates impulse control by an intertemporal bargaining technique, proposed as the mechanism of willpower.A theoretical model is described in which emotion is a reward-dependent behavior rather than a stimulus-bound respondent. Positive emotion is then limited by premature satiation of the appetite for it, a relentless process motivated by the impatience that is described by hyperbolic discount curves. This satiation can be restrained only by using adequately rare and unpredictable occasions as cues for the emotion. Willpower not only is helpless against the urge for premature satiation, but it exacerbates the satiation problem by making anticipation more thorough. The result is an asymmetrical contest between systematic attempts to vouchsafe satisfying events and impetuous attempts to put them at risk. Despite their adversarial relationship, both may to some extent be in the person's long range interest.     

Leptin-induced satiation mediated by abdominal vagal afferents

Leptin is a hormone secreted into the systemic blood primarily by white adipose tissue. However, leptin also is synthesized and stored by cells in the gastric mucosa. Because gastric mucosal leptin is secreted in response to ingestion of a meal, we hypothesized that it might contribute to satiation (meal termination) by acting on gastrointestinal vagal afferent neurons. To test whether leptin is capable of acutely reducing short-term food intake, we measured consumption of a liquid meal (15% sucrose) following low-dose leptin administration via the celiac artery, which perfuses the upper gastrointestinal tract. Leptin (1, 3, 10 mug) was infused via a chronically implanted, nonocclusive celiac arterial catheter or via a jugular vein catheter with its tip in the right cardiac atrium. Fifteen percent sucrose intake was then measured for 30 min. We found that leptin dose dependently inhibited sucrose intake when infused through the celiac catheter but not when infused into the general circulation via a jugular catheter. Plasma leptin concentrations in the general circulation following celiac arterial or jugular leptin infusions were not significantly different. Celiac arterial leptin infusion did not reduce meal size in vagotomized or capsaicin-treated rats. Finally, we also found that reduction of meal size by celiac leptin infusion was markedly enhanced when coinfused with cholecystokinin, a gastrointestinal satiety peptide whose action depends on vagal afferent neurons. Our results support the hypothesis that leptin contributes to satiation by a mechanism dependent on gastrointestinal vagal afferent innervation of the upper gastrointestinal tract.     

How Many Calories Are on Our Plate? Expected Fullness,Not Liking,Determines Meal‐size Selection

The availability of highly palatable food is thought to stimulate the selection of larger meals (leading to weight gain and obesity). In this article, we explore aspects of this proposition. Specifically, we scrutinize two basic assumptions: (i) palatable energy‐dense foods are more rewarding (desired), and (ii) these palatable foods are selected in relatively larger portions. In combination with palatability, we also consider the relative role for “expected satiation”—the extent to which a food is expected to deliver satiation. A total of 17 commonly consumed foods were assessed by 28 normal‐weight participants at lunchtime. Critically, our measure of food reward and expected satiation involves comparisons between foods based on equicaloric portions. When assessed in this way, we find that food reward and ideal portion sizes (in kcal) are both closely associated with expected satiation, but not with “expected liking.” Low expected satiation (not expected liking) predicts the selection of large portion sizes (in kcal) and foods with this characteristic tend to be more energy dense and are regarded as less (not more) rewarding (when compared calorie for calorie). Together, these findings challenge the role of palatability in meal‐size selection and they highlight the importance of expected satiation, a “nonaffective” component of food reward.     

The temporal pattern of feeding over the oestrous cycle of the mouse.

The detailed temporal pattern of feeding in female mice was measured by an automated method over the oestrous cycle whilst feeding on either of two foods differing in stimulus characteristics, but of similar calorific value. The total time spent feeding varied systematically over the cycle, being increased for sweet food and decreased for laboratory chow at the time of sexual receptivity. and increased for both foods towards the end of the cycle. The temporal pattern is analysed into a number of variables, which are seen to alter in different ways over the oestrous cycle. The pro-oestrus disturbance is due to alterations in meal size brought about solely by changes in mean bout length, themselves products of variations in the initial bouts of the meal. Inter-meal interval is unrelated to changes in meal size at this time, allowing the latter to affect food intake. The changes towards the end of the cycle are produced by alterations of meal frequency at constant meal size. It is argued that separable 'phasic' and 'tonic' effects of reproductive condition on feeding may therefore be identified, and that they might possibly have different functions.     

Effects of oral preload, CCK or bombesin administration on short term food intake of melanocortin 4-receptor knockout (MC4RKO) mice

We investigated whether either heterozygous (HET) or homozygous (knockout, KO) disruption of the melanocortin type 4 receptor (MC4R) gene alters post ingestive responsiveness of mice. Specifically, we tested the hypothesis that hyperphagia in MC4RKO mice might be due to a deficit in processes that sustain intermeal intervals (satiety) and/or processes that terminate ongoing episodes of eating (satiation). To test satiety, mice drank an oral preload and then we monitored intake of a subsequent liquid diet test meal. To test satiation, we examined the effect of exogenous administration of cholecystokinin (CCK) and bombesin (BN) on the size of a liquid diet meal. Experiment 1 was comprised of two studies. In the first, we determined that the intake of all three genotypes following fasts of either 6, 12, or 24 h were comparable, and so chose 12 h deprivation for the subsequent studies. In the second, 12 h fasted mice were allowed to consume a fixed preload, approximately 50% of their expected mean intake and, following delays of either 30 or 60 min, were allowed to consume to satiation. Compared with no preload, the preload significantly reduced meal size comparably in all three genotypes. The reduction in intake was greater when the test meal was presented 30 compared with 60 min after the preload, again with no genotype differences in this decay of satiety. In experiment 2, we administered either CCK or BN and examined suppression of meal size after a 12 h fast. Mice were tested repeatedly with CCK-8 (2, 6, or 18 μg/kg ip) or BN (2, 4 or 8 μg/kg ip) with vehicle injection days intervening. The 30 min intakes of HET and KO mice were suppressed more than those of WT following either CCK or BN. These experiments suggest that diminished responsiveness to nutrients or gut satiety hormones is not responsible for hyperphagia in MC4RKO mice.     

The adaptive value of gluttony: predators mediate the life history trade‐offs of satiation threshold

Animals vary greatly in their tendency to consume large meals. Yet, whether or how meal size influences fitness in wild populations is infrequently considered. Using a predator exclusion, mark–recapture experiment, we estimated selection on the amount of food accepted during an ad libitum feeding bout (hereafter termed ‘satiation threshold’) in the wolf spider Schizocosa ocreata. Individually marked, size‐matched females of known satiation threshold were assigned to predator exclusion and predator inclusion treatments and tracked for a 40‐day period. We also estimated the narrow‐sense heritability of satiation threshold using dam‐on‐female‐offspring regression. In the absence of predation, high satiation threshold was positively associated with larger and faster egg case production. However, these selective advantages were lost when predators were present. We estimated the heritability of satiation threshold to be 0.56. Taken together, our results suggest that satiation threshold can respond to selection and begets a life history trade‐off in this system: high satiation threshold individuals tend to produce larger egg cases but also suffer increased susceptibility to predation.     

The interpretation of exponential dose-effect curves

This paper deals with the following question: Which distributions of radiosensitivity in a population can lead to an exponential survival curve? The problem is solved exactly, with statistical fluctuations in dose fully accounted for. It is shown that only an exponential distribution of sensitivities can give rise to an exponential survival curve.     

Weight of food and time required to satiate brown trout, Salmo trutta L.

Brown trout of different weights (range 8-358 g) were fed to satiation at fifteen different water temperatures (range 3.8–21.6°C. Both the weight of the trout (Wg) and the water temperature (T°C) affected the maximum weight of food (Q mg) consumed in a meal, and the relationship between the three variables was well described by a multiple regression equation which can be used to estimate the value of Q (with 95% confidence limits) for trout of different weights at different temperatures between 3.8°C and 21.6°C. The satiation time (with 95% confidence limits) can also be estimated from a multiple regression equation for trout of different weights at temperatures between 6.8°C and 18.1°C. Estimates from the multiple regression equations were applicable to a wide range of food organisms with the exception of larvae ofTenebrio molitor (mealworms). Appetite (measured by voluntary food intake) varied with temperature and was greatest between 13.3°C and 18.4°C. From comparisons with the results of other workers, it was concluded that the maximum amount of food consumed in a meal may provide sufficient calories for both the daily metabolic requirements and the daily maintenance requirements of a trout at temperatures between 3.8°C and 18.4°C, but not at temperatures above 18.4°C.     

Apparent digestibility of crude protein and gross energy in some plant and animal-based feedstuffs by Clarias isheriensis (Siluriformes: Clariidae) (Sydenham 1980)

Apparent digestibility coefficient (ACD) of dry matter, crude protein and gross energy of white fish meal, blood meal, poultry by-product meal, hydrolysed feather meal, shrimp head meal, meat and bone meal, soybean meal, cottonseed meal, peanut meal, yellow maize, corn starch, cassava meal, sorghum, rice bran and cocoa pod meal were determined for Clarias isheriensis (47.5–51.2 g) fed to satiation with diets containing each test feedstuff in a 30:70 mixture with a reference diet, and chromium oxide as an indicator using the dissection technique. ADC_protein values ranged from 50 to 93 % while ADC_energy ranged from 42 to 98 %. Protein and energy content of animal or plant feed-stuffs were equally digestible to C. isheriensis.     

The effects of food type, meal size and body weight on digestion and gastric evacuation in turbot, Scophthalmus maximus L.

Experiments were conducted on a size range of turbot using whole prey organisms. Stomach contents were serially sampled at intervals after feeding on a range of meal sizes of sandeels and sprats. Gastric evacuation was essentially linear with time, though some fluctuations were observed. Control over evacuation appeared to be by regulation of the processes leading to chyme production. Close agreement between evacuation rate and the satiation feeding rate of fish in laboratory experiments was observed. Implications of these findings for techniques of estimating the feeding rates of natural fish populations are discussed.     

Sucking behaviour of dairy calves fed milk ad libitum by bucket or teat

The milk intake and sucking behaviour of Dutch and Holstein-Friesian crossbred calves fed milk replacer ad libitum by either the bucket method (n=8) or an artificial teat (n=6) were compared. The animals were observed for 3 weeks from the age of 2 weeks, penned individually and all provided with a “dummy” (artificial) teat near the milk source within the pen.

The teat-fed calves ingested significantly more milk than the bucket-fed calves (11.9 vs. 8.0 kg/day; P<0.05), and this intake took much longer (44.2 vs. 17.7 min/day; P<0.05). In both treatment groups, milk intake was organized in “meals”. The meal criterion, separating the withinmeal and between-meal non-feeding intervals, was set at 5 min. Frequency of meals and daily total meal duration did not differ significantly. Meals occurred rather randomly throughout a 24-h period.

On average, the dummy teat was used for 13 min per day by bucket-fed calves, but for only 1 min by teat-fed calves (P<0.05). Sucking of the dummy teat was largely clustered within the meal periods.

It is concluded that in the young calf a need for sucking exists independently of milk satiation. However, the level of satiation depends on whether the calf drinks or sucks the milk. Nutritive sucking is clearly more reinforcing than non-nutritive sucking.     

Satiation and the functional response: a test of a new model

Abstract. 1. A model of the functional response to prey density is derived to include the reduction in time available for search, T_s , resulting from predator satiation.
2. For larger prey items predator satiation occurs at each prey capture and T_s is reduced by the attack time and digestive pause of a series of attack cycles. For small prey items predator foraging is continuous at low densities with T_s reduced solely by attack time. At higher densities predator satiation occurs after the capture of several small prey items and T_s is reduced by the attack time and digestive pause of a series of foraging cycles.
3. A comparison of the predicted asymptotic level of prey capture using experimentally estimated parameter values, with the maximum consumption of aphids by larval and adult coccinellids provides a test of the satiation model.
4. The limitation of prey capture by predator satiation is discussed with reference to handling time and the success of coccinellids in biological control.     

Effects of temperature and feed intake on astaxanthin digestibility and metabolism in Atlantic salmon, Salmo salar

The effects of feed intake, growth rate and temperature (8 and 12 °C) on apparent digestibility coefficients (ADC), blood uptake of individual astaxanthin E/Z isomers and metabolism of astaxanthin (3,3′-dihydroxy-β,β-carotene-4,4′-dione) were determined in Atlantic salmon. Accumulation of idoxanthin (3,4,3′-trihydroxy-β,β-carotene-4-one) in plasma was used to indicate metabolic transformation of astaxanthin. Quadruplicate groups of fish were subjected to three different treatments; one treatment was kept at 12 °C and fed to satiation. Another treatment kept at 12 °C was pair-fed with fish fed to satiation at 8 °C, resulting in a restricted feeding regime for the former treatment. After 2 months of feeding, the fish were fed a single meal containing ballotini glass beads to determine individual feed intake and Y₂O₃ as an inert marker to determine ADCs. The faeces samples were pooled into 6 categories according to individual meal size (range 0.2–1.5% of body weight) and the ADCs for different meal sizes were determined. ADCs of astaxanthin ranged from 20% to 60% but were not significantly correlated with meal size. However, fish kept at 12 °C had approximately 10% higher ADC than fish kept at 8 °C (p = 0.032). Growth rate and plasma astaxanthin concentration were higher at higher temperature and higher ration. Plasma concentration of idoxanthin was not affected by temperature or by meal size. The incidence of fin erosion and non-feeding individuals was significantly higher among fish fed a restricted ration indicating more aggressive interactions. Fish with visible fin damage had a tendency for having higher idoxanthin content in plasma than fish without noticeable fin damage. It is concluded that temperature but not individual meal size affect ADC of astaxanthin, whereas both influence plasma astaxanthin levels and may therefore affect the efficiency of astaxanthin utilization.