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.     

To split behaviour into bouts, log-transform the intervals

Analysis of behaviour that is displayed in bouts depends crucially on quantitative estimates of bout criteria, that is, the lengths of the shortest intervals between bouts. Current methods estimate bout criteria by modelling the log-transformed (cumulative) frequency distributions of intervals between events. For analysis of feeding behaviour, these models will not result in biologically meaningful quantitative estimates (Tolkamp et al. 1998, Journal of Theoretical Biology194, 235-250). We proposed a method that models the frequency distribution of log-transformed interval lengths instead. Applying this method to a single data set showed that the log-transformed lengths of intervals between feeding events were distributed as two Gaussians. Here we test this model using a data set of 35 171 intervals between feeding that was obtained during an experiment with 38 cows in three dietary treatment groups. No meaningful bout criterion could be obtained for some individuals, which casts doubt on the general validity of the proposed model. Addition of a third log-normal improved the fit of the model and we hypothesized that this third population represents intervals including drinking. In a second experiment, we found the measurements to be consistent with this hypothesis. We obtained meaningful meal criteria for all individuals by fitting either a double, or a triple, log-normal model to the frequency distributions of the lengths of intervals between feeding. These log-normal models appear to be not only more biologically meaningful than log (cumulative) frequency models but also far more flexible. Copyright 1999 The Association for the Study of Animal Behaviour.     

Measuring meals: structure of prandial food and water intake of rats

Attempts to understand ingestion have sought to understand the control of meals. The present study evaluated a meal definition that included prandial drinking (drinking-explicit meals). The spontaneous nocturnal intake of male Wistar rats was studied. The meal breakpoint was defined as the interval between feeding or drinking events providing the most stable estimate of meal structure. Alternative breakpoints derived from prevailing methodology, log-survivorship, or frequency histogram analysis of interfeeding intervals without respect to drinking were compared (drinking-naive meals). Threshold interfeeding intervals that accounted for drinking indirectly were evaluated as surrogate breakpoints (drinking-implicit meals). Definitions were compared by determining which criterion better conformed to predictions of satiety. Microstructural differences resulting from the definitions were also studied. Under the drinking-explicit definition, rats averaged nine or ten 13-min meals/night, during which they consumed food and water equally in duration (5-6 min) and quantity (2.3 g). Individual differences were observed in microstructure measures. Meals defined by drinking-informed, but not drinking-naive, methods were followed by the behavioral satiety sequence and by an initially low likelihood of resuming feeding that monotonically increased with time. The drinking-explicit definition uniquely revealed preprandial and postprandial correlations of similar magnitude. Under drinking-informed definitions, food restriction increased meal size, whereas drinking-naive definitions increased meal frequency. Drinking-implicit definitions provided workable approximations of meal frequency and size but inferior estimates of feeding duration, eating rate, and the preprandial correlation. Thus log-survivorship analysis is not appropriate for identifying meal breakpoints, and the consideration of drinking in meal definitions can provide a better estimate of meal structure.     

A novel flexible method to split feeding behaviour into bouts

Before meal patterns can be analysed properly, a biologically relevant meal criterion must be determined in order to group short-term feeding behaviour into meals. Existing methodologies are based on modelling of the frequency distributions of intervals between feeding events but these methods cannot be used if the proper distributions cannot be clearly identified. For such cases we developed two new methods – (1) based on the analysis of the distribution of between-meal interval lengths only and (2) based on the analysis of changes in the probability of animals starting to feed with time since the last feeding event. Both methods were developed using a data set of over 700,000 records of visits to feeders obtained with broilers (Gallus gallus) aged between 2 and 5 weeks. The two methods resulted in meal criteria estimates of 20.1 and 17.5 min, respectively, which, when applied to the data set, gave statistically significant but very small differences in meal characteristics. The new methods were tested against an independent cow (Bos taurus) data set and the resultant meal criteria compared with those predicted by an existing method. The two novel methods estimated meal criteria for cows at 27.9 and 35.5 min, compared with 28.9 min for the existing method. Again, these differences in meal criteria resulted in only very small differences in meal characteristics. Even though meal criteria were relatively similar for birds and cows, characteristics such as average daily number of meals (10.9 and 5.9), meal size (12.5 g and 7.4 kg) and meal duration (7.7 and 31.4 min) were very different. The analyses show that, if the appropriate distributions of intervals cannot be identified, meal criteria can still be estimated for species as diverse as mature cows and young birds by the novel methodologies developed here.     

Stomach temperature telemetry reveals temporal patterns of foraging success in a free-ranging marine mammal

1. We studied feeding frequency in free-ranging grey seals using stomach temperature telemetry to test if previously reported sex differences in the diving, movement and diet were reflected in the temporal pattern of foraging success. 2. Data were retrieved from 21 of 32 grey seals from 1999 to 2001, totalling 343 days and 555 feeding events, with individual record length varying from 2 to 40 days (mean: 16.33 +/- 2.67 days/seal). 3. Seals fed on 57.8 +/- 6.46% of days sampled and had an average of 1.7 +/- 0.26 meals per day, but individual variability was apparent in the temporal distribution of feeding as evidenced by high coefficients of variation (coefficient of variation = 69.0%). 4. Bout analysis of non-feeding intervals of six grey seals suggests that feeding intervals of individuals were varied and probably reflect differences in prey availability. Grey seals tended to have many single feeding events with long periods separating each event, as would be expected for a large carnivore with a batch-reactor digestive system. 5. We found significant sex differences in the temporal distribution of feeding. The number of feeding events per day was greater in males (2.2 +/- 0.4 vs. 1.0 +/- 0.2), as was time associated with feeding per day (56.6 +/- 5.8 min vs. 43.9 +/- 9.4 min). 6. The number of feeding events varied with time of day with the least number occurring during dawn. Feeding event size differed significantly by time of day, with greater meal sizes during the dawn and the smallest meals during the night. 7. The length of time between meals increased with the size of the previous meal, and was significantly less in males (541.4 +/- 63.5 min) than in females (1092.6 +/- 169.9 min). 8. These results provide new insight into the basis of sex differences in diving and diet in this large size-dimorphic marine predator.     

The structure of feeding behavior in the colorado potato beetle,Leptinotarsa decemlineata (Coleoptera: Chrysomelidae)

Feeding behavior, in an ad libitum situation on potato plants in the laboratory, was continuously observed for approximately 7 h/day on 2 successive days for 18 adult femaleLeptinotarsa decemlineata. Additional behaviors were also recorded including resting, walking, biting, local movements, grooming, defecating, and regurgitating. These data were used to calculate a time budget for the various behaviors. The feeding data were analyzed to describe the structure of feeding for young adult females on their normal host plant. The criterion for a meal (minimum intermeal interval) was determined to be 286 s. This criterion was used to distinguish between intra- and intermeal interruptions in feeding for all subsequent analyses. Meals taken from leaves that were young, medium aged, or old did not differ, but on average beetles took 60% of their meals from young leaves. Meal size and meal duration were equally good predictors of when a meal would end. Feeding from stems was a prominent feature for most beetles. The stem meals had much longer durations than leaf meals, but stem feeding did not affect subsequent leaf feeding. The structure of feeding by these beetles is compared with that found in other insects, especiallyLocusta migratoria.     

Correlations between meals and inter-meal intervals in Japanese quail and their significance in the control of feeding

Significant correlations (P<0.05) between meals and preceding intervals were shown more often by Japanese quail when fed on diluted mash (40% cellulose) than with undiluted mash or pellets. They showed significant correlations between meals and succeeding intervals with about the same frequency on all three foods. Most of the correlation coefficients were low, but experiments in which interval length and meal size were manipulated artificially confirmed that close relationships between meals and intervals can occur, and appear to verify the existence of short-term hunger and satiety mechanisms. Possible explanations for the low correlation coefficients shown by several bird species are discussed, and it is concluded that meal-eating is controlled by a very flexible system. There is no evidence that the timing of meals depends on fixed set points, and it is suggested instead that degrees of hunger and satiety may determine the probabilities of a meal starting or stopping, such a system being associated with emptying and filling of parts of the digestive tract.     

Compensatory feeding in response to variable food quality by Melanoplus differentialis

Abstract. The behavioural mechanisms driving compensatory food consumption in response to dietary dilution as well as the relationship between feeding time and food residence time (i.e. digesta retention time in the gut) were studied using the non-diapausing strain of the grasshopper Melanoplus differentialis (Thomas) (Orthoptera: Acrididae). 3-day-old, sixth-instar nymphs and 3-day-old adults were fed artificial diets containing 1%, 3% and 5% total nitrogen (N) at 30C, LD 14:10h; the feeding behaviour was recorded using electronic monitoring devices connected to microcomputers for 24 h. The percentage of time spent feeding increased linearly as diets were diluted using non-digestible cellulose from 5% to 1% N. This response was due to an increase in the number of meals while the meal duration of a feeding bout was unaltered. Sixth-instar nymphs spent about 40% more time feeding than the larger adults. The increased feeding time in nymphs resulted from both more frequent feeding bouts and longer meal duration. Feeding time and food residence time were highly negatively related.     

Circadian clocks for all meal-times: anticipation of 2 daily meals in rats

Anticipation of a daily meal in rats has been conceptualized as a rest-activity rhythm driven by a food-entrained circadian oscillator separate from the pacemaker generating light-dark (LD) entrained rhythms. Rats can also anticipate two daily mealtimes, but whether this involves independently entrained oscillators, one 'continuously consulted' clock, cue-dependent non-circadian interval timing or a combination of processes, is unclear. Rats received two daily meals, beginning 3-h (meal 1) and 13-h (meal 2) after lights-on (LD 14:10). Anticipatory wheel running began 68±8 min prior to meal 1 and 101±9 min prior to meal 2 but neither the duration nor the variability of anticipation bout lengths exhibited the scalar property, a hallmark of interval timing. Meal omission tests in LD and constant dark (DD) did not alter the timing of either bout of anticipation, and anticipation of meal 2 was not altered by a 3-h advance of meal 1. Food anticipatory running in this 2-meal protocol thus does not exhibit properties of interval timing despite the availability of external time cues in LD. Across all days, the two bouts of anticipation were uncorrelated, a result more consistent with two independently entrained oscillators than a single consulted clock. Similar results were obtained for meals scheduled 3-h and 10-h after lights-on, and for a food-bin measure of anticipation. Most rats that showed weak or no anticipation to one or both meals exhibited elevated activity at mealtime during 1 or 2 day food deprivation tests in DD, suggesting covert operation of circadian timing in the absence of anticipatory behavior. A control experiment confirmed that daytime feeding did not shift LD-entrained rhythms, ruling out displaced nocturnal activity as an explanation for daytime activity. The results favor a multiple oscillator basis for 2-meal anticipatory rhythms and provide no evidence for involvement of cue-dependent interval timing.     

An analysis of the pattern of feeding in the Costa Rican weevil, Exopthalmus jekelianus, on the tropical tree, Cedrela odorata

Feeding patterns were recorded and analysed for adult female weevils, Exopthalmus jekelianus (White) (Coleoptera: Curculionidae), feeding on Central American mahogany, Cedrela odorata L., in the field in Costa Rica. The study forms part of an investigation into the relationship between feeding patterns and the fine-scale variation in leaf chemistry occurring within the host plant. The weevils’ feeding patterns were the simplest in temporal structure of any reported to date for an insect herbivore. Weevils spent an average of only 3% of their time feeding during the 10-h observation periods. Meals lasted an average of 2.8 min and occurred at a mean intermeal interval of 84 min. The feeding patterns gave the appearance of a short-term rhythm underlying the onset of feeding (as has been found in locusts and caterpillars), although there were insufficient meals taken by individuals over the 10-h period to test this suggestion. Meals were notable in apparently lacking intrameal pauses and also commencing without preliminary sampling behaviours, such as palpating or biting. Whether the combination of short, infrequent meals, ingested without pauses and not preceded by sampling behaviour, represents an adaptation reducing apparency to natural enemies, or else simply reflects low nutritional needs, is discussed. Correlations between meal durations and following and preceding intermeal intervals suggested that variation in intermeal intervals stemmed largely from variation in meal duration, not vice versa, with variation in meal duration resulting from an external influence such as leaf nutritional and/or allelo-chemistry. The latter suggestion is currently being tested.     

The structure of feeding behavior in a phytophagous insect (Hylobius abietis)

Analysis of the feeding behavior of animals using such a high temporal resolution that meals can be defined may improve our understanding of the mechanisms regulating feeding. Meals can be distinguished in an ethologically meaningful manner by using the ‘meal criterion’, the shortest non‐feeding interval between feeding bouts recognized as meals. However, such a criterion has only been determined for a few insect species. Applying a recent method developed for assessing meal criteria for vertebrates, we determined the meal criterion for Hylobius abietis (L.) (Coleoptera: Curculionidae) based on data from video recordings of single individuals feeding on seedlings of Norway spruce, Picea abies (L.) Karst. (Pinaceae). The pine weevil is an economically important pest insect, because it feeds on the stem bark of planted conifer seedlings. Weevils had 4–5 meals per day. Each meal lasted about 24 min during which about 13 mm² of bark per meal were removed. Females had longer total meal durations and longer non‐feeding intervals within meals than males. Girdling seedlings did not affect the weevils' feeding properties. The size of meals was significantly correlated with the duration of non‐feeding intervals before and after them. This study is one of few describing the feeding behavior of an insect at a temporal resolution that allows individual meals to be distinguished. With more meal‐related data from insects available, differences in meal properties may be interpreted based on phylogeny, ecology, and physiology. Our results may also assist in the setup and interpretation of studies of plant‐insect interactions, and facilitate the evaluation and development of methods to protect plants against herbivores.     

The pattern of feeding of first instar nymphs of Schistocerca americana

ABSTRACT The feeding behaviour of nymphs Schistocerca americana (Drury) was recorded throughout the light phases of the first stadium using a behavioural event recording program on a microcomputer. Most food was ingested on day 4 of the stadium, very little on day 1 and none on day 6, the final day of the stadium. Only 26–45% of the total food intake over the stadium occurred during the dark periods and more food was eaten in the last 4 h of each light period than earlier. Feeding occupied only about 5% of the total time. Variation in food intake was a consequence primarily of changes in the numbers of meals. After day 1, the average meal size did not change significantly. Meal length increased relatively slightly compared with meal size as a consequence of an increase in the proportion of time spent feeding in each meal as meal size increased. Meal size was correlated with the length of the previous interfeed suggesting a volumetric regulation. The distribution of pauses within meals was also consistent with the thesis that meal size is governed by the level of excitation in the central nervous system at the start of the meal.     

Provisioning and growth rates of nestling Fulmars Fulmarus glacialis: stochastic variation or regulation?

The pattern of chick feeding in the Fulmar Fulmarus glacialis at St Kilda, Scotland, was examined by repeated weighing of chicks throughout 14 consecutive days during the first half of the chick-rearing period in 1994. After correcting for metabolic weight losses, the sizes of positive mass increments between weighings were used to assess meal sizes and feeding frequency for each chick. Individual meals fed to chicks averaged 80.8 g (s.d. ± 21.0 g), or approximately 10% of adult mass. Each chick received 0 to 4 meals per day, with an average of 1.9 meals per chick per day, giving an average interval of around 25 h between meals delivered by each parent. The distribution of time intervals between feeds for each chick (whether single or double meals) followed a negative exponential function with a maximum value of 80 h. These results are not compatible with the idea that the purpose of large fat deposits in procellariiform chicks is to guarantee survival over long intervals between feeds. Over 14 days, the chicks' mean daily food requirements for zero-growth increased from 98 g to 160 g. This corresponded with an increase in feeding frequency but not meal size. Chicks with lower scores for body condition after feeding by both parents received more meals during the subsequent 16 h and had shorter intervals to the next feed, indicating that adults regulated feeding frequency in accordance with chick condition at the previous feed. This does not agree with the hypothesis that lipid accumulation by nestling Procellariiformes is a response to stochastic variation in food delivery associated with an absence of regulation. In view of the diversity of growth and feeding patterns present among the Procellariiformes, it is possible that lipid accumulation in this group does not have a unitary explanation.     

The effect of deprivation on the microstructure of feeding by the Tobacco hornworm caterpillar

An automated device was used to examine, in detail, feeding on disks of wheat germ medium by fifth-instar Manduca sextacaterpillars. Comparisons were made between some animals which had ad libitum access to food at all times and others which were deprived of food for 1–5 h before being tested. Feeding patterns of both groups indicated regulation of feeding both between and within meals. Deprived animals ate more during their first meal than did nondeprived animals chiefly by increasing (a) the number of chewing bouts (and thus the meal duration) and (b) the bite frequency. Calculations indicated that the deficit caused by deprivation was made up during the first meal. However, deprived animals continued to eat more than nondeprived ones in subsequent feeding also. Passage of food through the gut was examined by dissecting out the contents of each region of the gut at various times after a colored test meal. Food passed through the foregut directly into the anterior part of the midgut. It stayed in the middle third of the midgut longer than in the anterior and posterior thirds, and the first pellet resulting from the test meal appeared 4 h after the meal. The following mechanisms of feeding regulation are proposed: (a) volumetric feedback mediated by stretch receptors of the foregut and anterior third of the midgut which terminates meals; (b) the development and subsequent reduction of satietyspecific behaviors mediated either by stretch receptors or by some other means which, e.g., allow the next meal to begin; and (c) metabolites whose levels drop during deprivation, triggering a series of events which lead to the excess feeding observed.     

Neural ensemble coding of satiety states

The motivation to start or terminate a meal involves the continual updating of information on current body status by central gustatory and reward systems. Previous electrophysiological and neuroimaging investigations revealed region-specific decreases in activity as the subject's state transitions from hunger to satiety. By implanting bundles of microelectrodes in the lateral hypothalamus, orbitofrontal cortex, insular cortex, and amygdala of hungry rats that voluntarily eat to satiety, we have measured the behavior of neuronal populations through the different phases of a complete feeding cycle (hunger-satiety-hunger). Our data show that while most satiety-sensitive units preferentially responded to a unique hunger phase within a cycle, neuronal populations integrated single-unit information in order to reflect the animal's motivational state across the entire cycle, with higher activity levels during the hunger phases. This distributed population code might constitute a neural mechanism underlying meal initiation under different metabolic states.     

Altered Feeding Patterns in Rats Exposed to a Palatable Cafeteria Diet: Increased Snacking and Its Implications for Development of Obesity

Background

Rats prefer energy-rich foods over chow and eat them to excess. The pattern of eating elicited by this diet is unknown. We used the behavioral satiety sequence to classify an eating bout as a meal or snack and compared the eating patterns of rats fed an energy rich cafeteria diet or chow.

Methods

Eight week old male Sprague Dawley rats were exposed to lab chow or an energy-rich cafeteria diet (plus chow) for 16 weeks. After 5, 10 and 15 weeks, home-cage overnight feeding behavior was recorded. Eating followed by grooming then resting or sleeping was classified as a meal; whereas eating not followed by the full sequence was classified as a snack. Numbers of meals and snacks, their duration, and waiting times between feeding bouts were compared between the two conditions.

Results

Cafeteria-fed rats ate more protein, fat and carbohydrate, consistently ingesting double the energy of chow-fed rats, and were significantly heavier by week 4. Cafeteria-fed rats tended to take multiple snacks between meals and ate fewer meals than chow-fed rats. They also ate more snacks at 5 weeks, were less effective at compensating for snacking by reducing meals, and the number of snacks in the majority of the cafeteria-fed rats was positively related to terminal body weights.

Conclusions

Exposure to a palatable diet had long-term effects on feeding patterns. Rats became overweight because they initially ate more frequently and ultimately ate more of foods with higher energy density. The early increased snacking in young cafeteria-fed rats may represent the establishment of eating habits that promote weight gain.     

The temporal pattern of feeding in the zebra finch

Feeding in zebra finches occurs in clearly defined bouts, but strong individual differences have been found in the finer details of its pattern. Some birds showed a constant probability of starting feeding with passage of time between meals and a constant probability of stopping during a meal. In these cases meal length (number of pecks at seed) tended to correlate with the length of the preceding gap. By contrast, in most individuals both meals and gaps tended to be of a typical length, and in some of these autocorrelation showed feeding to follow a cycle approximately 24 to 30 min long. Meal length in most of these birds correlated strongly with the length of the succeeding gap. The individual differences found are discussed and hypotheses put forward for their causation.     

Effect of meal viscosity and nutrients on satiety, intragastric dilution, and emptying assessed by MRI

The relationship between the intragastric distribution, dilution, and emptying of meals and satiety was studied using noninvasive magnetic resonance imaging techniques in 12 healthy subjects with four polysaccharide test meals of varying viscosity and nutrient content as follows: 1) low-viscosity nonnutrient, 2) low-viscosity nutrient, 3) high-viscosity nonnutrient, and 4) high-viscosity nutrient. Increasing the nutrient content of the high-viscosity meal delayed gastric emptying from 46 +/- 9 to 76 +/- 6 min (P < 0.004), whereas increasing viscosity had a smaller effect. The volume of secretions within the stomach 60 min after ingestion was higher for the high-viscosity nutrient meal (P < 0.04). A simple model to calculate the total volume of secretion added to the test meal is presented. Color-coded dilution map images showed the heterogeneous process of progressive gastric dilution of high-viscosity meals, whereas low-viscosity meals were uniformly diluted. Fullness was found to be linearly related to total gastric volumes for the nutrient meals (R(2) = 0.98) and logarithmically related for the nonnutrient meals (R(2) = 0.96). Fullness was higher for high- compared with low-viscosity meals (P < 0.02), and with the nutrient meals this was associated with greater antral volumes (P < 0.05).     

Can one unrestricted meal buffer the effects of previous pre-meal intervals on the feeding behaviour of sheep?

Varying the time since the last meal is one means of manipulating feeding motivation. In order to use this method effectively it is necessary to know whether and the extent to which effects of one pre-meal interval are carried over to affect the behaviour during the following meals. Pre-meal interval (PMI) is defined here for practical purposes, for short meals, as the time between the start of two successive meals. The possibility that one unrestricted meal might buffer the effects of an 8h as opposed to a 4h PMI on aspects of feeding behaviour was studied with eight Scottish Blackface sheep. They were fed on a regime in which they were given access to food until they finished their meal and lay down (this always occurred within 60min) at which time the remaining food was withdrawn. Feeding behaviour was recorded during the meal after these 4 and 8h intervals, as well as during the following meal 4h later.At a meal after a PMI of 8h, compared to 4h, sheep had a higher intake per meal (mean+/-S.T.D. for 8 and 4h PMI, respectively: 604.4+/-78.8 and 430.1+/-100.9g; P<0.001), a longer meal duration (27.1+/-7.5 and 21.8+/-8.1min; P<0.001), and a tendency for a higher intake rate (23.8+/-6.2 and 21.9+/-8.2g/min; P=0.11). During the following meal 4h later these differences were smaller, but intake per meal still tended to be higher (430.8+/-81.5 and 338.5+/-45.6g; P<0.06) for sheep who had previously had the 8h PMI. Meal duration (21.9+/-7.2 and 20.6+/-7.08min; P=0.28) and intake rate (21.2+/-6.1 and 18.7+/-7.2g/min; P=0.13) were no longer different.A single meal after the different PMIs reduced differences in all three aspects of feeding behaviour observed during the subsequent meal, 4h later, but differences in intake per meal were still apparent. It is suggested that an additional meal may overcome the carry-over effect.     

Cholecystokinin-33 inhibits meal size and prolongs the subsequent intermeal interval

There are various forms of the satiety gut-brain peptide cholecystokinin (CCK), a short, widely utilized form or CCK-8, and a long, putatively more effective form or CCK-33. The issue of which of these forms is a more effective satiety peptide is not resolved. Here, we compared the satiety responses, including the sizes of the first three meals (MS) and intermeal intervals (IMI) as well as their calculated satiety ratios (SR), evoked by both peptides. CCK-8 and 33 (1, 3 and 5 nmol/kg, i.p) reduced the size of the first meal similarly, only CCK-33 prolonged the first IMI and increased SR and both peptides failed to affect second and third MS and IMI. As such, CCK-33 is a more effective satiety peptide than CCK-8. The current results confirm previous findings which showed that both peptides reduce food intake by inhibiting meal size, whereas only CCK-33 reduces food intake by prolonging the intermeal interval.