Cyclic estradiol treatment normalizes body weight and restores physiological patterns of spontaneous feeding and sexual receptivity in ovariectomized rats

Hypothalamic-pituitary-gonadal axis function strongly influences feeding and body weight in cycling females in many species. To test the sufficiency of cyclic variations in plasma estradiol to reproduce normal patterns of spontaneous feeding, food intake, and body weight, ovariectomized Long-Evans rats were subcutaneously injected every fourth day with 2 microg estradiol benzoate or with the oil vehicle alone. Cyclic estradiol treatment completely normalized the trajectory of body weight gain and total food intake through seven treatment cycles. The hyperphagia of ovariectomized rats was expressed as an increase in spontaneous meal size. Meal frequency decreased, but not enough to compensate for the increase in meal size. Estradiol treatment normalized both parameters. In addition, cyclic estradiol treatment produced a further phasic decrease in meal size (and increase in meal frequency) and a decrease in food intake during the second night after injection. This phasic change is similar to the feeding changes occurring during estrus in intact rats. Sexual receptivity was measured during the eighth estradiol treatment cycle, 4 h after injection of 0.5 mg progesterone. Lordosis scores at the time of the treatment cycle modeling estrus were maximal, and scores at the time modeling diestrus were slightly increased over those of rats that did not receive estradiol. Finally, plasma estradiol levels, measured during the ninth treatment cycle, revealed a near-normal cyclic pattern of plasma estradiol levels. These results provide the first demonstration that the induction of a cyclic, near-physiological pattern of plasma estradiol is sufficient to maintain normal levels of body weight, spontaneous feeding patterns, total food intake, and (together with progesterone) sexual receptivity in ovariectomized rats.     

Oscillators entrained by food and the emergence of anticipatory timing behaviors

Circadian rhythms are adjusted to the external environment by the light-dark cycle via the suprachiasmatic nucleus, and to the internal environment of the body by multiple cues that derive from feeding/fasting. These cues determine the timing of sleep/wake cycles and all the activities associated with these states. We suggest that numerous sources of temporal information, including hormonal cues such as corticoids, insulin, and ghrelin, as well as conditioned learned responses determined by the temporal relationships between photic and feeding/fasting signals, can determine the timing of regularly recurring circadian responses. We further propose that these temporal signals can act additively to modulate the pattern of daily activity. Based on such reasoning, we describe the rationale and methodology for separating the influences of these diverse sources of temporal information. The evidence indicates that there are individual differences in sensitivity to internal and external signals that vary over circadian time, time since the previous meal, time until the next meal, or with duration of food deprivation. All of these cues are integrated in sites and circuits modulating physiology and behavior. Individuals detect changes in internal and external signals, interpret those changes as "hunger," and adjust their physiological responses and activity levels accordingly.     

Patterns in feeding: a behavioural analysis using Locusta migratoria nymphs

ABSTRACT. Feeding patterns of Locusta migratoria L. nymphs, kept under LD 12:12 at 30°C with ample food, were analysed over the first 6 days of the fifth instar. A number of differences were found between the light and dark phases. (1) More was eaten during the light than the dark, with the differences becoming progressively larger over the 6 days. (2) During light phases most insects showed an increasing tendency to feed with time since the last meal, while in the dark the probability of starting to feed remained constant. (3) There was little variation in feeding patterns within the light phase, but marked changes occurred within the dark phase, feeding declining to a low level in the last 4h. The amount eaten during a meal was regulated similarly in the light and dark, with time rather than amount eaten being of paramount importance in terminating a meal. The time when feeding started was somewhat influenced by the size of the preceding meal, but once feeding began meal size was largely independent of the preceding meal. The importance of food quality in determining the pattern of feeding is discussed.     

Ecological consequences of scaling of chew cycle duration and daily feeding time in Primates

Feeding systems and behaviors must evolve to satisfy the metabolic needs of organisms. This includes modifications to feeding systems as body size and metabolic needs change. Using our own data and data from the literature, we examine how size-related changes in metabolic needs are met by size-related changes in daily feeding time, chew cycle duration, volume of food processed per chew, and daily food volume intake in primates. Increases in chew cycle duration with body mass in haplorhine primates are described by a simple power function (cycle time α body mass^0.181). Daily feeding time increases with body mass when analyzed using raw data from the “tips” of the primate phylogenetic tree, but not when using phylogenetically independent contrasts. Whether or not daily feeding time remains constant or increases with body mass, isometry of ingested bite size and the slow rate of increase in chew cycle time with body size combine to allow daily ingested food volume to scale faster than predicted by metabolic rate. This positive allometry of daily ingested food volume may compensate for negative allometry of nutrient concentration in primate foods. Food material properties such as toughness and hardness have little impact on scaling of chew cycle durations, sequence durations, or numbers of chews in a sequence. Size-related changes in food processing abilities appear to accommodate size-related changes in food material properties, and primates may alter ingested bite sizes in order to minimize the impacts of food material properties on temporal variables such as chew cycle duration and chew sequence duration.     

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.     

Developmental changes in the patterns of feeding in fourth- and fifth-instar Helicoverpa armigera caterpillars

Data are presented for developmental changes in feeding behaviour within and across the fourth and fifth stadium of Helicoverpa armigera (Lepidoptera, Noctuidae) caterpillars fed nutritionally homogeneous semi‐synthetic foods. We recorded the microstructure of feeding over continuous 12‐h periods on consecutive days throughout the two stadia, and in one experiment recorded continuously for 21 h. Larvae in the two stadia showed the same general pattern of macro‐events in feeding, including a similar duration of post‐ecdysis fast, which was usually broken by consumption of the exuviae, and then a sustained period in which discrete meals on the experimental food were taken regularly. There were, however, some distinct differences in the patterns of meal‐taking both between stadia and across different one‐third time segments within stadia. Considering between‐stadium differences, the proportion of time spent feeding differed significantly only in the last segment of the feeding period of the two stadia, with the value for the fourth‐instar larvae being substantially greater than for fifth‐instar larvae. As regards within stadium changes, the proportion of time feeding increased from the first to the second segment of both stadia. However, whereas the proportion of time feeding increased from the second to the final segment of the fourth stadium, it decreased across the same period in the fifth stadium. These patterns of changes in the proportion of time feeding within and between stadia, and their behavioural mechanisms (combination of meal durations and meal frequencies), can be explained only partially with reference to increasing food requirements with development. Three areas are identified where further study might help elucidate the reasons for the observed developmental changes in the microstructure of feeding: allometric constraint, the dynamic links between ingestion and post‐ingestive processing, and ecological factors such as predation.     

Temporal control of feeding behaviour and its association with gastrointestinal function

This paper summarises knowledge about temporal control of ad libitum feeding in poultry, from minute to minute, hour to hour and day to day, and about how it relates to aspects of gastrointestinal function. Evidence is presented of only loose control over initiation and termination of spontaneous meals, and it is proposed that degrees of hunger and satiety determine probabilities of feeding starting and stopping. Voluntary regulation of food intake can be considered in terms of adjustments in mean meal size, meal frequency or both. Short-term variation is associated more with meal frequency and longer-term changes more with meal size. Short-term adjustments appear to depend more on alimentary control and longer-term adjustments more on metabolic control (not considered here). Long-term changes affecting meal size are associated with changes in capacity of parts of the alimentary tract. Food can accumulate in the crop and gizzard, and meal initiation and termination are associated with varying degrees of emptying and filling of these diverticula during most of the day. Later in the day there is usually a conditioned change to cumulative filling of the crop (and gizzard) with food that is digested overnight. Possible roles of osmo-/chemoreceptors and gut peptides are discussed.     

Intracerebroventricular C75 decreases meal frequency and reduces AgRP gene expression in rats

3-Carboxy-4-alkyl-2-methylenebutyrolactone (C75), an inhibitor of fatty acid synthase and stimulator of carnitine palmitoyltransferase-1, reduces food intake and body weight in rodents when given systemically or centrally. Intracellular molecular mechanisms involving changes in cellular energy status are proposed to initiate the feeding and body weight reductions. However, effectors that lie downstream of these initial steps are not yet fully identified. Present experiments characterize the time courses of hypophagia and weight loss after single injections of C75 into the lateral cerebroventicle in rats and go on to identify specific meal pattern changes and coinciding alterations in gene expression for feeding-related hypothalamic neuropeptides. C75 reduced chow intake and body weight dose dependently. Although the principal effects occurred on the first day, weight losses relative to vehicle control were maintained over multiple days. C75 did not affect generalized locomotor activity. C75 began to reduce feeding after a 6-h delay. The hypophagia was due primarily to decreased meal number during 6-12 h without a significant effect on meal size, suggesting that central C75 reduced the drive to initiate meals. C75 prevented the anticipated hypophagia-induced increases in mRNA for AgRP in the arcuate nucleus at 22 h and at 6 h when C75 begins to suppress feeding. Overall, the data suggest that gene expression changes leading to altered melanocortin signaling are important for the hypophagic response to intracerebroventricular C75.     

Changes in blood histamine and bradykininogen levels during different periods of the oestrous cycle in goats.

Blood histamine and plasma bradykininogen levels were estimated in eighteen normally cyclic goats at different periods of the oestrous cycle. The histamine level increased significantly (P smaller than 0.05) on Day 18 of the oestrous cycle. The bradykininogen level decreased significantly (P smaller than 0.05) towards the end of oestrus.     

The patterning of compensatory sugar feeding in the Australian sheep blowfly

Abstract. The pattern of feeding is described for males of the Australian sheep blowfly, Lucilia cuprina , with ad libitum access to either 0.1 M or 1.0 M glucose solution. Flies given the 0.1 M solution ingested nearly 3 times the volume taken by flies given the 1.0 M solution by eating meals of, on average, twice the size about 1.5 times as frequently. Flies were usually relatively inactive following a meal, with the extent of this post-prandial quiescence being a function both of meal size and concentration of sugar. Quiescence lasted only about 20% of the average intermeal interval, however, and there was no correlation between meal size and time to the next meal. The crop emptied more slowly when it contained 1.0 M rather than 0.1 M glucose solution and the crop was, on average, fuller at the beginning of a meal on the higher concentration. The volume of solution imbibed during a meal was positively correlated with time since the end of the preceding meal. The average crop volume at the end of a meal was similar in flies feeding on 0.1 M and 1.0 M solutions. The results are considered in relation to published information on control of feeding and compensation in the blowfly Phormia regina.     

Postprandial responses in the African rhombig egg eater (Dasypeltis scabra)

The African rhombic egg eater (Dasypeltis scabra) is a colubrid snake feeding exclusively on bird eggs. Frequency of feeding is governed by the seasonal availability of bird eggs; i.e., long fasting intervals change with relatively short periods when plenty of food is available. Intermittent feeding snakes show a remarkable postprandial increase of metabolic rate and digestive organ size. The postprandial increase in metabolic rate (specific dynamic action, SDA) in snakes is affected by meal size, temperature, and meal composition. A major portion of SDA in snakes is allocated to gastric function and the breakdown of the meal. We hypothesize that SDA in egg eaters is lower than in other snake species, because egg eaters feed on “liquid” food that does not require enzymatic breakdown in the stomach. We also hypothesized that other components of the postprandial response of egg eaters (e.g., size changes of the intestine and the liver) do not differ from other snakes. The standard metabolic rate and metabolic response to feeding were measured using closed-chamber respirometry. Size changes of small intestine and liver were measured using high-resolution transcutaneous ultrasonography. Standard metabolic rates of fasting egg eaters were in the same range of mass specific values as known from other snakes. Within 24 h after feeding, oxygen consumption doubled and peaked at 2 days after feeding. At the same time, the size of the small intestine and the cross-sectional diameter of the liver increased. Within 2 days after feeding, the size of the mucosal epithelium doubled its thickness. Liver size increased significantly within 24 h reaching maximum size 2–4 days after feeding. The size of both organs returned to fasting values within 7–10 days after feeding. The postprandial response of African rhombic egg eaters shows the same pattern and dynamics as known from other snake species. However, the factorial increase of metabolic rate during SDA is the lowest reported for any snake. A comparison with literature data supports the idea that SDA is mainly determined by gastric function and that it is low in egg eaters because they do not have to break down solid meals in the stomach as other snake species do.     

Gastric emptying in Limanda limanda (L.) and the return of appetite

Limanda limanda (35–225 g) fed during daytime when trained to use demand feeders under natural photoperiods. Under continuous illumination different feeding rhythms developed which were detected by periodogram analysis. Interfeeding periods (P hours) varied with body weight (W grams) and temperature (T° C) over the range 6.5–15° C as: In P=4.4 – 0.167 (In W) – 0.05 T. Daily food intake (D grams increased with both temperature and size (In D = 0.099 T + 0.579 In W – 3.49). Relative daily intake (R as % body weight, b.w) increased with temperature but decreased with size (In R = 0.099T–0.41 In W + 1.05). These changes in daily intake are primarily caused by changes in feeding frequency; the amount eaten at a meal is relatively constant save at low temperatures (6.5° C) when appetite is suppressed. Dilution of the food with kaolin (from 4.8 to 2.2 kcal/g) did not cause a compensatory increase in meal size or frequency. Gastric emptying time (GET hours) measured by X-radiography changed with body weight, temperature and relative meal size (M % b.w.) as In GET = 0.68 In M + 0.39 In W – 0.035T + 1.46. With the exception of the lowest temperature (6.5° C) interfeeding periods closely followed GET (P = 1.09 GET – 2.6) supporting the proposal that stomach fullness is a major factor controlling appetite in the dab.     

Patterns of respiration in Locusta migratoria nymphs when feeding

Flow‐through respirometry was used to investigate patterns of respiration of fifth‐instar Locusta migratoria L. nymphs fed a chemically defined, synthetic food. Each animal was recorded for up to 2.7 h, during which they had access to food and water ad libitum, and at least one meal was taken. The start of feeding was coincident with a sudden and rapid rise in respiration. Both carbon dioxide (CO₂) production and oxygen (O₂) consumption rose, the traces for the two gasses showing a high degree of alignment. The end of a meal correlated with a sudden and rapid decrease in respiratory rate towards resting levels. When feeding was interrupted by an intra‐meal pause, respiratory rate tended to drop marginally and then stabilize, before rising rapidly upon the resumption of feeding within the meal. Maximal rates of respiration during feeding represented a 3–4‐fold increase over those at rest. Walking and climbing within the chamber were not associated with any noticeable change in respiratory rate above baseline. When locusts were quiescent between feeding episodes, respiration was steady and continuous, rather than discontinuous. Possible causes for large changes in respiration during feeding are discussed.     

Meal Pattern Analysis of Macronutrient Intake Aafter PVN Norepinephrine and Peripheral Clonidine Administration

Norepinephrine (NE) injected into the paraventricular nucleus (PVN) of the hypothalamus of rats is a potent stimulant of food intake, more specifically ingestion of the carbohydrate nutrient. In 2 experiments of the present study, this effect was found to be dose-dependent, and the effectiveness of NE in potentiating total food consumption was greatly reduced when the carbohydrate diet was removed. In addition, experiments using a computer-automated data acquisition apparatus were performed to characterize, in detail, the impact of PVN injection of NE and peripheral administration of the α2-nor-adrenergic agonist clonidine (CLON) on the macrostructure of feeding behavior in animals given 3 pure macronutrient diets. These 2 compounds, injected at the onset of the nocturnal feeding cycle, had very similar effects on meal patterns, with both affecting nutrient intake by increasing meal size and duration rather than by increasing meal frequency. They both affected primarily the first meal of the dark cycle, selectively enhancing carbohydrate ingestion by increasing Kcal intake, percent composition in the total diet and feeding time, and also by decreasing the satiating impact of this macronutrient. These stimulatory effects of NE and CLON on carbohydrate ingestion during the first meal were followed by complete recovery over the next 1 to 2 hours after injection. In addition to these predominant effects on carbohydrate intake, PVN NE at the highest doses tested (10 and 20 nmoles) produced a small increase in fat intake, whereas peripheral CLON actually decreased intake of fat and protein over the 12-hour cycle. The similarities in the impact of NE and CLON on carbohydrate feeding patterns support the hypothesis that both agonists may be acting via the same PVN α2-noradrenergic system controlling ingestion of the carbohydrate-rich meals which predominate at dark onset.     

Energetic costs at different levels of feeding in juvenile cod, Gadus morhua L.

The energetic costs associated with feeding by juvenile cod were determined by means of an open-circuit respirometer. Fish acclimated to several temperatures (7, 10, 15 and 18°C) were kept at natural lighting levels, and fed inside their individual respirometers. They consumed a diet compounded from natural foods, at five different ration levels, their oxygen consumption being monitored continually over an 11–16 day period.
After each meal the rate of oxygen consumption increased to above the pre-feeding level, reaching a peak 8–10 h later. With each successive meal the oxygen consumption showed a cumulative increase, reaching a maximum usually after the last meal.
The elevation in metabolic rate associated with feeding was dependent upon ration size, increasing linearly as the food intake increased. The effect was also dependent upon temperature; for fish fed to satiation the total energy cost was equivalent to 11.9, 10.9, 16.4 and 17.1% of the ingested energy at 7, 10, 15 and 18°C respectively. For resting satiated fish the rate of oxygen consumption was close to the maximum rate for active fish.     

Meal pattern of male rats maintained on histidine-, leucine-, or tyrosine-supplemented diet

Objective: Food intake is known to be affected by macronutrient composition of the diet, and protein manipulation has been reported to alter food intake, but the effect of individual amino acids on eating behavior has not been fully studied. This study investigated the effect of diet supplementation with three individual amino acids on meal pattern in male rats. Research Methods and Procedures: Thirty‐two Sprague‐Dawley rats were randomly divided into four equal groups and fed control diet or histidine (5%)‐, leucine (5%)‐, or tyrosine (5%)‐supplemented diet for 2 weeks and were monitored for their meal pattern. Results: Total food intake and feeding rate of the different groups were not affected, although other components of meal pattern were altered. Histidine supplementation reduced diurnal meal size by 42% (p < 0.05), whereas that of leucine increased nocturnal meal size by ～35% (p < 0.05). Tyrosine supplementation increased food intake of the nocturnal period and decreased that of the diurnal period. Both histidine and tyrosine supplementation elevated fasting plasma insulin levels and suppressed fasting glucose significantly. Discussion: Individual amino acids were found to alter meal pattern differently. Further investigations are required to dissect the involvement of central and peripheral factors in these alterations.     

The effect of stomach fullness on food intake of whiting in the North Sea

The probability of a North Sea whiting Merlangius merlangus stomach containing fresh food was depressed when partially digested food was already present in the stomach. The lowered probability was detected even at levels where the fish was physiologically able to ingest an average meal. The feeding probability of c . 15% of the fish caught was predicted to be severely decreased at the level of partially digested food found in the stomachs. No effect of stomach fullness on meal size was found, indicating that the saturation is affecting search activity rather than prey or meal size selection. The diurnal pattern in food intake varied between the five sampling locations, presumably as a result of differences in prey availability.     

Does the Anopheles blood meal-fecundity curve, curve?

We determined the blood meal size-fecundity relationship in Anopheles gambiae in the laboratory. Our primary interest was to determine whether the fecundity curve has a non-linear component, i.e.. does it decelerate towards an asymptote? Small and large adult females in their second gonotrophic cycle were fed on human hosts for predetermined lengths of time. Blood meal size was ascertained by weight and subsequent fecundity was determined by the number of eggs laid on filter paper. In both small and large-bodied females there was a significant curvilinear component. We used this relationship to show that marginal returns from feeding decline as a function of current blood load. This means that further fecundity payoff for continued feeding declines over time.     

Effects of testosterone upon feeding in male mice.

Food intake and pattern of feeding were recorded in intact and castrate male mice. Intact adults are larger and eat more than pre-pubertal castrates, due to rapid growth at puberty. Castration in adulthood has little effect upon food intake, but feeding becomes concentrated into large, infrequent meals. The change in meal size appears primary and is reversed by testosterone. During rapid growth at puberty, both meal size and frequency are high. It is suggested that testosterone affects feeding in two ways. Rapid weight gain at puberty leads to high meal size and frequency, but in adulthood meal size is depressed. Meal size in adults is further depressed by stimuli from females, and it is argued in this case that testosterone affects a general mechanism for altering feeding when other activities have a high priority.     

Feeding: Models of Costs and Benefits in Energy Regulation

Models to predict feeding behavior at the level of consumptionand use of energy involve either details of internal (physiological)controls or economic principles of regulation based on optimal(evolutionary) foraging theory. These two approaches will ultimatelybe related, but the former requires more information for specificpredictions. The latter can provide predictions based on selectedcriteria for regulation. Meal sizes and feeding frequencies of hummingbirds are examinedrelative to two regulatory, criteria: maximizing rate of netenergy gain and maximizing efficiency (intakes/expenditures)through a "crop emptying" model that incorporates energy intakefrom food and energy expenditures for short-term (meal to meal)maintenance and longer-term (overnight) energy storage. Experimentalresults suggest that the feeding behavior of hummingbirds isdifferentially sensitive to short-term and daily uses of energy.Changes in overnight energy storage requirements result primarilyin changes in meal size, while changes in meal to meal maintenancerequirements result primarily in feeding frequency changes.The economic models predict these responses. The feeding behaviorof hummingbirds also appears to be sensitive to food quality,time spent flying to and from a food source, and costs associatedwith the weight of ingested food.