Physiological mechanisms underlying the control of meal size in Manduca sexta larvae

Abstract. Fifth stadium larvae of the tobacco hornworm, Manduca sexta (L.), ate larger meals than usual when they had been deprived of food for periods of time longer than the usual intermeal interval (c. 45 min). Meal size increased with time since the last meal until 180 min, when it was about 3 times normal. There was no evidence of a role for volumetric feedback from the gut in controlling meal size. Injections of a paraffin oil/wax mixture, or of petroleum jelly (Vaseline) into the foregut, midgut or rectum failed to decrease meal size. Cutting the recurrent nerve failed to alter meal size compared to sham-operated controls (although both groups took smaller meals than unoperated controls). By contrast, injections of an extract of soluble nutrients from the diet into the midgut inhibited feeding in some insects and reduced subsequent meal size in others. Appropriate controls showed that these effects were not due to the volumetric or osmotic effects of the injections. These results imply that nutrient feedback plays an important role in controlling meal size in Manduca caterpillars, while volumetric feedback is probably unimportant.     

Countercurrent flows,water movements and nutrient absorption in the locust midgut

Using amaranth dye as a marker solute, the movements of fluids in the gut of Schistocerca gregaria was studied, either by feeding a meal containing the dye or by injecting the dye into the haemolymph, and by comparing the distribution of amaranth with those of naturally-occurring solutes in the alimentary tract.In animals deprived of food for more than 2–4 hr, some of the fluid from the Malpighian tubules moves forward through the solid food matrix in the midgut carrying solutes into the anterior midgut and gastric caeca, where water is absorbed. After a meal the crop empties at a rate which saturates the absorptive capacity of the anterior caeca, producing a net movement of fluid down the midgut and so such a countercurrent system is not observed in animals fed ad lib., where dye introduced into the gut always moves posteriorly.A countercurrent fluid movement confers several advantages on the alimentary system which act to maximise the efficiency of nutrient absorption: the principal disadvantage of the countercurrent system is that noxious solutes, as well as nutrients, will accumulate at high concentrations near the permeable site of nutrient uptake. Thus a countercurrent flow of solutes is observed only when the insect is deprived of food and the need to conserve nutrient resources exceeds that of excretion of noxious substances. Ways in which the site of nutrient absorption may be protected from noxious solutes are discussed.The anterior caeca gradually become bloated with dark fluid as digestion proceeds; this is expelled into the midgut when a fresh meal is ingested.     

Gut morphology,feeding rate and gut clearance in five species of caddis larvae

Feeding rate, the rate of movement of food through the gut and gut morphology of large larvae of five caddis species (Halesus radiatus, Hydropsyche instabilis, Polycentropus kingi, Rhyacophila dorsalis and Potamophylax cingulatus) were investigated in the laboratory. Following 72 hr starvation, P. cingulatus and H. radiatus larvae became satiated (refused prey offered directly to the mouthparts) after consuming 8–11 and 9–13 mayfly nymphs (Baetis rhodani, 3.5–4.6 mm) respectively. Hunger level affected prey consumption. In P. cingulatus, the number of prey consumed over 24 h (at 9.5–12 °C) increased with starvation periods from 0–72 h, but declined following longer starvation periods. Six clearly recognisable gut states (defined by the position of food material in various parts of the gut) can be identified at different times since commencement of a meal. As environmental temperature increased (from 8–12 °C to 15–17 °C), the rate of change of the gut state increased and the food retention time decreased in all species. Feeding periodicity (i.e. nocturnal/diurnal activity) in the field was estimated based on the evacuation rate and the gut state and environmental temperature at the known time of collection. Initiation of consumption of prey appeared to coincide with emptying of the foregut and proximal midgut, whereas actual feeding continued until complete satiation when some threshold fullness of the foregut had been reached. The length of time food was held in the foregut was positively correlated with increasing specialisation of the foregut (particularly elaboration of the proventriculus).     

The functional morphology of the alimentary canal of larval stages of the parasitic copepod Lepeophtheirus salmonis

The functional morphology of the alimentary canal of copepodite and chalimus stages of Lepeophtheirus salmonis (Krøyer, 1837) is described and compared with that found in other copepods studied to date.
The buccal cavity passes into a gut comprising three major regions: foregut (oesophagus), midgut and hindgut. The foregut and hindgut both posscss a cuticular lining whereas the midgut is lined with specialized epithelial cells. The midgut is divided into three recognizable zones, namely anterior midgut caecum, anterior midgut and posterior midgut. Three main types of epithelial cell are recognizable in the midgut: vesicular cells, microvillous cells and basal cells which correspond to the cell types normally described in other parasitic and free-living copepod species.
Digestion is thought to occur in the midgut and be mediated by the epithelial cells that line it. Although several glands appear to discharge into the area of the buccal cavity, none was seen to interface to any other area of the gut. There was no evidence for the involvement of commensal gut bacteria in food digestion.     

Identification of leucomyosuppressin in the German cockroach, Blattella germanica, as an inhibitor of food intake

The feeding pattern of the adult female of Blattella germanica peaks in the middle of the vitellogenic cycle. Following the hypothesis that a factor inhibiting gut peristalsis also inhibits food intake and is involved in the regulation of feeding, we searched for the most powerful myoinhibitory peptide in brain extracts from B. germanica females collected after the peak within the feeding cycle. Through HPLC purification and sequence analysis, we obtained the peptide leucomyosuppressin (LMS): pQDVDHVFLRFamide. LMS elicited a powerful myoinhibitory effect on B. germanica foregut and hindgut, with ED(50) values around 10(-10) M. In addition, it inhibited food intake in vivo in a dose-dependent manner at doses between 5 and 50 microg. The study of the distribution of ingested food in the foregut, midgut and hindgut of B. germanica females treated with LMS showed that food accumulates in the foregut, which may be due to the myoinhibitory effects of the peptide. We propose that this accumulation inhibits food intake because of the persistence of the signals from gut stretch receptors.     

Food deprivation and the regulation of meal size in larvae of Chrysopa carnea

ABSTRACT. The course of repletion and the effects of food deprivation on meal size were explored in three experiments on larvae of Chrysopa carnea (Neuroptera). Feeding to repletion was found to occur within the first 30 min of exposure to food. Meal size increased as an ogival function of deprivation, up to the limit of gut capacity. Behavioural components involved in the initiation of feeding were little affected by deprivation and did not appear to be inhibited by distention of the gut. Termination of a meal may be mediated by the stimulation of prey-release behaviour, rather than by inhibition of feeding.     

Reduced feeding during water deprivation depends on hydration of the gut

Removal of drinking water at the start of the dark period reduced food intake in freely feeding rats within 45 min. Both first and later meals were smaller during 7.5 h of water deprivation, but meal frequency did not change. Ingestion of a normal-sized meal (3 g) rapidly increased plasma tonicity when drinking water was withheld, but intravenous infusions of hypertonic NaCl causing similar increases in plasma tonicity did not reduce feeding. Feeding during 6 h of water deprivation was restored by slowly infusing the volume of water normally drunk into the stomach, jejunum, or cecum, but not in the vena cava or hepatic portal vein. The infusions did not alter water or electrolyte excretion or affect food intake in rats allowed to drink. We conclude that the inhibition of feeding seen during water deprivation is mediated by a sensor that is located in the gastrointestinal tract or perhaps in the mesenteric veins draining the gut, but not the hepatic portal vein or the liver. In the absence of drinking water, signals from this sensor provoke the early termination of a meal.     

Feeding behaviour of a new worm (Priapulida) from the Sirius Passet Lagerstätte (Cambrian Series 2, Stage 3) of North Greenland (Laurentia)

Singuuriqia simoni gen. et sp. nov. represents the first record of a priapulid worm from the Sirius Passet Lagerstätte (Cambrian Series 2, Stage 3) of North Greenland (Laurentia). It is defined by an unusually broad, longitudinally folded, foregut which tapers through the pharynx towards the anterior mouth; posteriorly, the same longitudinal folding is evident in the narrow gut. The slender, smooth, trunk in the unique specimen passes anteriorly into an oval proboscis which culminates in a smooth, extensible, pharynx with pharyngeal teeth. The capacity for substantial expansion of the foregut permitted rapid ingestion of food prior to digestion at leisure. Cololites suggest both carnivorous and deposit feeding behaviour, indicating that Singuuriqia, like the present day Priapulus, was probably omnivorous.     

Hoarding versus the immediate consumption of food among hamsters and gerbils

This project dealt with a comparative analysis of the effects of food deprivation on feeding and hoarding in hamsters and gerbils. The animals were given food in their home cage and their food intake was measured during a 30-min period after which they were transferred to an apparatus in which hoarding behaviour was assessed. The results indicated interesting species differences in the animals' reactions in the test situations. Whereas food-deprived gerbils ate more food than nondeprived gerbils, the food intake of hamsters was not significantly enhanced by deprivation. In the hoarding test, although significant differences were observed between deprived and nondeprived gerbils, the absolute level of hoarding in gerbils was very low in comparison to that of hamsters tested under the same conditions. These results suggest that hamsters and gerbils respond to challenges to their food reserves with different strategies.     

Predator gut state and prey detectability using electrophoretic analysis of gut contents

Abstract. I. An investigation of the effects of position of food in the gut (gut state), time since feeding, meal size and environmental temperature on the detection of prey using the electrophoretic separation of proteins with esterase activity was undertaken. The waterbug Notonecta glauca Linn (Hemiptera/Heteroptera) was used as the model predator.
2. Seven easily recognizable gut states can be identified at different times since commencement of a meal.
3. Specific prey esterase 'fingerprint' bands can be detected whilst food remains in the foregut.
4. Meal size and environmental temperature (affecting digestion and assimilation rates) both affect the rate of filling and emptying of the foregut, and hence the time period over which prey proteins can be detected by electrophoresis following a meal. Small meals and high temperatures lead to quite rapid changes in gut state with time and thus reduce the time period over which prey can be identified from gut contents (20 h). Large meals and low temperatures extend this period (48 h).
5. Predator gut state is the most important parameter in prey detectability from gut contents, and from a visual examination of the gut state it is immediately evident whether or not prey detection by electrophoresis will be possible.
6. There is further evidence in support of a twocompartment food processing model for Nofoonecru , and the significance of this type of processing in relation to gut content analysis of sucking predators is discussed.     

Mucous contribution to gut nutrient content in American gizzard shad Dorosoma cepedianum

This study developed and applied an approach to calculate the proportion of fish gut content composed of mucus secreted by the oropharyngeal cavity and gut. The amount of nitrogen in the contents of the foregut (oesophagus and gizzard) and the epibranchial organs of suspension‐feeding American gizzard shad Dorosoma cepedianum was significantly higher than the nitrogen in the homogeneous food source. Using data collected from suspension‐feeding experiments and the nitrogen content of D. cepedianum mucus, a series of equations illustrated that mucus constituted c. 10% of D. cepedianum foregut content and 12% of epibranchial organ content by dry mass. Future quantification of fish feeding selectivity and absorption efficiency can use this approach to take into account the contribution of fish mucus to the nutrients in the gut contents. This study supports the conclusion that suspension‐feeding D. cepedianum in a heterogeneous environment selectively ingest nutrient‐rich particles, even when gut nutrient content is adjusted to take into account the contribution of mucus.     

The regulation of feeding in Locusta migratoria: Internal inhibitory mechanisms

During the course of a meal by insects with an empty gut, food is held in the foregut and the midgut remains empty. The last part of the foregut to fill is the extreme anterior end of the crop, and stretch receptors in this region control the amount eaten. Cutting the posterior pharyngeal nerves isolates these receptors from the frontal garglion and results in hyperphagia pharyngeal nerves isolates these receptors from the frontal garglion and results in hyperphagia during one meal. On unfavourable food, when smaller amounts are eaten, distension of the foregut is not involved in regulating meal size.Feedback from receptors in the body wall is not important in regulating meal size; most of the increase in gut volume resulting from feeding is taken up by the collapse of the air sacs so that there is little or no change in the body volume.

---

Zusammenfassung Alle Hauptnerven des Mund- und Rachennervensystems wurden kontrolliert reseziert. Erst wenn die rückwärtigen Rachennerven zerschnitten werden, nehmen die Insekten wesentlich mehr Nahrung zu sich. Solche Insekten versuchen längere Zeit zu fressen.Wenn sich während des Fressens der Vorderdarm füllt, wird keine Nahrung an den Mitteldarm weitergegeben und sein Vorderende ist der letzte Teil, der sich vollständig ausdehnt. Nach Durchtrennung der rückwärtigen Rachennerven sind alle Teile des Vorderdarms offen-sichtlich mehr gedehnt. Eine solche erhöhte Nahrungsaufnahme tritt aber nicht ein, wenn die Nahrung relativ ungünstig ist. Das Abbinden des Kropfes verursacht beim nachträglichen Fressen keine übermäßige Ausdehnung des Vorderteiles.Die Nahrungsaufnahme hat keinen Unterschied im Körpervolumen zur Folge und das Durchtrennen des ventralen Nervenstranges bleibt ohne Einfluß auf die nachträglich aufgenommenen Nahrungsmengen. Das vergrößerte Darmvolumen wird durch Kollaps des Luftsackes kompensiert.

     

Is the food‐entrainable circadian oscillator in the digestive system?

Food-anticipatory activity (FAA) is the increase in locomotion and core body temperature that precedes a daily scheduled meal. It is driven by a circadian oscillator but is independent of the suprachiasmatic nuclei. Recent results that reveal meal-entrained clock gene expression in rat and mouse peripheral organs raise the intriguing possibility that the digestive system is the site of the feeding-entrained oscillator (FEO) that underlies FAA. We tested this possibility by comparing FAA and Per1 rhythmicity in the digestive system of the Per 1-luciferase transgenic rat. First, rats were entrained to daytime restricted feeding (RF, 10 days), then fed ad libitum (AL, 10 days), then food deprived (FD, 2 days). As expected FAA was evident during RF and disappeared during subsequent AL feeding, but returned at the correct phase during deprivation. The phase of Per1 in liver, stomach and colon shifted from a nocturnal to a diurnal peak during RF, but shifted back to nocturnal phase during the subsequent AL and remained nocturnal during food deprivation periods. Second, rats were entrained to two daily meals at zeitgeber time (ZT) 0400 and ZT 1600. FAA to both meals emerged after about 10 days of dual RF. However, all tissues studied (all five liver lobes, esophagus, antral stomach, body of stomach, colon) showed entrainment consistent with only the night-time meal. These two results are inconsistent with the hypothesis that FAA arises as an output of rhythms in the gastrointestinal (GI) system. The results also highlight an interesting diversity among peripheral oscillators in their ability to entrain to meals and the direction of the phase shift after RF ends.     

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.     

Meal size in nymphs of Locusta migratoria

Meal size in fifth-instar nymphs of Locusta migratoria L. has been investigated by weighing the gut and its contents immediately after feeding. Larger meals are taken in mid-instar, while females eat more in one meal than males under the same conditions. Bigger meals are also eaten after longer periods of food deprivation, but after deprivation for two hours or less, corresponding with the normal interfeed length, meals are smaller than would be expected from the potential maximum foregut volume. The maximum volume attained varies with the type of food available to the insects and is influenced by previous conditioning. Temperature and humidity do not affect meal size except at 40°, at which temperature the usual pattern of feeding is completely altered.

---

Zusammenfassung Die Nahrungsmenge, die von einer Larve der Wanderheuschrecke, Locusta migratoria L., im fünften Entwicklungsstadium bei einer einzigen Mahlzeit verzehrt wird, wurde ermittelt, indem ihr Darm und Darminhalt sofort nach dem Fressen gewogen wurden. In der Mitte des Entwicklungsstadiums wurden größere Mengen verzehrt, Weibchen fraßen mehr als Männchen, jedoch sind die weiblichen Larven ja auch größer. Die Menge der aufgenommenen Nahrung vergrößerte sich mit der Dauer künstlichen Futterentzuges. Sie erreichte ihren Höhepunkt nach sechs Stunden, nach welcher Zeit der Vorderdarm absolut leer war.Die verzehrten Mengen waren je nach der Art der Nahrung unterschiedlich. Auch stellten die Insekten sich so ein, daß sie von einer bestimmten Nahrung mehr zu sich nahmen, wenn sie daran aufgezogen worden waren. Nach längeren Zeitspannen der Isolierung hatten sie die Tendenz, weniger zu fressen.Temperatur und Feuchtigkeit hatten keinen Einfluß auf die Mengen, die verzehrt wurden, außer bei 40°, bei welcher Temperatur sich das gesamte Verhalten zum Fressen ändert.

     

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.     

Modifications in DARPP‐32 phosphorylation pattern after repeated palatable food consumption undergo rapid habituation in the nucleus accumbens shell of non‐food‐deprived rats

In non‐food‐deprived rats a palatable meal induces a transient increase in dopamine output in the prefrontal cortex and nucleus accumbens shell and core; habituation to this response develops with a second palatable meal, selectively in the shell, unless animals are food‐deprived. A palatable meal also induces time‐dependent modifications in the dopamine and cAMP‐regulated phosphoprotein of Mr 32 000 (DARPP‐32) phosphorylation pattern that are prevented when SCH 23390, a selective dopamine D₁ receptor antagonist, is administered shortly after the meal. This study investigated whether dopaminergic habituation in the shell had a counterpart in DARPP‐32 phosphorylation changes. In non‐food‐deprived rats, two consecutive palatable meals were followed by similar sequences of modifications in DARPP‐32 phosphorylation levels in the prefrontal cortex and nucleus accumbens core, while changes after the second meal were blunted in the shell. In food‐deprived rats two consecutive meals also induced similar phosphorylation changes in the shell. Finally, SCH 23390 administered shortly after the first palatable meal in non‐food‐deprived rats inhibited DARPP‐32 phosphorylation changes in response to the first meal, and prevented the habituation to a second meal in terms of dopaminergic response and DARPP‐32 phosphorylation changes. Thus, dopamine D₁ receptor stimulation plays a role in the development of habituation.     

State‐dependent attacks in a mosquito

Mosquito biting frequency and meal size are considered to be important parameters in the epidemiology of insect‐vectored diseases such as malaria. Because both parameters are likely to depend on the size and energetic state of adult mosquitoes, the present study investigates the effects of body size and energy state on attack behaviours in the malaria mosquito, Anopheles gambiae. Attack rates are measured as well as total time spent before giving up for individual females when provided with an unobtainable human hand (i.e. mosquitoes are dislodged every time that they land). The factorial design considers two body sizes, small and large, as well as three sugar deprivation states, 0, 1 and 2 days. The results reveal a positive effect of size on attack rate and a nonlinear effect of energy state, where mosquitoes of intermediate energy state show lower attack rates than either 2‐day food‐deprived or nondeprived mosquitoes. Moreover, attack rate is negatively associated with persistence time in nondeprived and 2‐day food‐deprived Anopheles but is unrelated to persistence time in 1‐day food‐deprived mosquitoes, Interestingly, although persistence times are generally inversely related to attack rates, they are not significantly influenced by either energetic or size states.     

The interactive effects of exercise and feeding on oxygen uptake, activity levels, and gastric processing in the graceful crab Cancer gracilis

Exercise and digestive processes are known to elevate the metabolic rate of organisms independently. In this study, the effects of simultaneous exercise and digestion were examined in the graceful crab Cancer gracilis. This species exhibited resting oxygen uptake levels between 29 and 42 mg O(2) kg(-1) h(-1). In postprandial crabs, oxygen uptake was approximately double that of unfed crabs. During exercise, oxygen uptake increased three- to fourfold, reaching maximal levels of more than 130 mg O(2) kg(-1 ) h(-1). However, there was no difference in oxygen uptake during activity between unfed and postprandial animals. There was also no difference in exercise endurance levels between unfed and postprandial animals; both sets of animals were unable to right themselves after being turned on their backs, reaching exhaustion after 13-15 attempts. To determine whether increased activity affected gastric processes, the passage of a meal through the digestive system was followed using a fluoroscope. Passage of digesta through the gut system was slower in active animals than in resting crabs. Resting crabs cleared the foregut after approximately 18 h, which was significantly faster than the 34.5 h for constantly active animals. Likewise, the midgut region of resting animals was cleared at a faster rate than that of active animals. Because of residual amounts of digesta remaining in the hindgut, no difference in clearance rates of this section of the gut was evident. The slower clearance times of the foregut were due to a significantly slower rate of mastication of food, as evidenced by a lower cardiac stomach contraction rate. Contraction of the pyloric region of the foregut functions to move the digesta along the midgut, and there was a direct correlation between slower contraction rates of this region and the increased time of passage for digesta through the midgut of active animals. Because increased activity levels affected gastric processing, the crabs exhibited a behavioral response. During a 24-h period after feeding, there was a significant reduction in locomotor activity. The findings of this study suggest a prioritization of metabolic responses toward activity at the expense of digestion. This is discussed in relation to the ability of the crabs to balance the demands of competing physiological systems.     

Ontogeny of endocrine cells in the gut of the insect Periplaneta americana

Summary The ontogeny of the endocrine cells of the gut of the cockroach Periplaneta americana was studied by immunohistochemistry. During embryogenesis, the midgut begins to be formed as an outgrowth of the foregut and hindgut invaginations. Gut endocrine cells with pancreatic polypeptide (PP)-like immunoreactivity begin to appear at the anterior and posterior ends of the forming midgut. These cells are restricted to the midgut epithelium, and no mitotic cells with PP-like immunoreactivity are observed. These results strongly suggest that the gut endocrine cells, at least those with PP-like immunoreactivity, are derived from precursor cells they have in common with other epithelial cells of the midgut.