Aspects of feeding,including estimates of gut residence time,in three mytilid species (Bivalvia,Mollusca) at two contrasting sites in the Cape Peninsula,South Africa

Individuals of three mytilid species (Choromytilus meridionalis; Perna perna; Aulacomya ater) from two sites characterised by different qualities of ration available to these suspension feeders showed different rates in some components of the physiological energy budget. These differences included higher feeding (=clearance) and respiration rates, but lower absorption efficiencies, in individuals from the site with the higher quality ration. A novel technique was employed to estimate the residence time of food particles in the digestive gland. Individuals showing higher feeding rates had shorter residence times than those feeding more slowly and a significant positive correlation was demonstrated between residence time and absorption efficiency. These relationships, together with an exponential increase in rates of respiratory heat loss with an increase in ingested ration, are suggested to provide these animals with a physiological flexibility to compensate for reduced food quality in a way consistent with some theoretical predictions. Such compensations are made more effective if the total gut capacity can also change in response to the quality of the ration.     

Longer guts and higher food quality increase energy intake in migratory swans

1. Within the broad field of optimal foraging, it is increasingly acknowledged that animals often face digestive constraints rather than constraints on rates of food collection. This therefore calls for a formalization of how animals could optimize food absorption rates. 2. Here we generate predictions from a simple graphical optimal digestion model for foragers that aim to maximize their (true) metabolizable food intake over total time (i.e. including nonforaging bouts) under a digestive constraint. 3. The model predicts that such foragers should maintain a constant food retention time, even if gut length or food quality changes. For phenotypically flexible foragers, which are able to change the size of their digestive machinery, this means that an increase in gut length should go hand in hand with an increase in gross intake rate. It also means that better quality food should be digested more efficiently. 4. These latter two predictions are tested in a large avian long-distance migrant, the Bewick's swan (Cygnus columbianus bewickii), feeding on grasslands in its Dutch wintering quarters. 5. Throughout winter, free-ranging Bewick's swans, growing a longer gut and experiencing improved food quality, increased their gross intake rate (i.e. bite rate) and showed a higher digestive efficiency. These responses were in accordance with the model and suggest maintenance of a constant food retention time. 6. These changes doubled the birds' absorption rate. Had only food quality changed (and not gut length), then absorption rate would have increased by only 67%; absorption rate would have increased by only 17% had only gut length changed (and not food quality). 7. The prediction that gross intake rate should go up with gut length parallels the mechanism included in some proximate models of foraging that feeding motivation scales inversely to gut fullness. We plea for a tighter integration between ultimate and proximate foraging models.     

Nutmeg mannikins (<Emphasis Type="Italic">Lonchura punctulata</Emphasis>) reduce their feeding rates in response to simulated competition

Group feeding animals experience a number of competitive foraging costs that may result in a lowered feeding rate. It is important to distinguish between reductions in feeding rates that are caused by reduced food availability and physical interactions among foragers from those caused by the mere presence of foraging companions that may be self-imposed in order to obtain some benefit of group membership. Starlings (Sturnus vulgaris) reduce their feeding rates when in the company of simulated competitors located in an adjacent cage that cannot affect the food availability or interact with the forager. In the present study, we investigate whether the presence of simulated competitors in another species of passerine, nutmeg mannikins (Lonchura punctulata), can result in self-imposed reductions in feeding rates. When feeding in the company of simulated competitors, mannikins spent more non-foraging time near them, fed more slowly, reduced travel times between patches, reduced their scanning time and pecked more slowly. These results provide evidence that simulated competitors induce a reduction in pecking rate: behavioural interference. These self-imposed responses to competitors may have resulted from attempts to remain close to the non-feeding companions. Such self-imposed reductions in feeding rates may be a widespread yet generally unrecognised foraging cost to group feeding individuals.     

Costs and benefits of within-group spatial position: a feeding competition model

An animal's within-group spatial position has several important fitness consequences. Risk of predation, time spent engaging in antipredatory behavior and feeding competition can all vary with respect to spatial position. Previous research has found evidence that feeding rates are higher at the group edge in many species, but these studies have not represented the entire breadth of dietary diversity and ecological situations faced by many animals. In particular the presence of concentrated, defendable food patches can lead to increased feeding rates by dominants in the center of the group that are able to monopolize or defend these areas. To fully understand the tradeoffs of within-group spatial position in relation to a variety of factors, it is important to be able to predict where individuals should preferably position themselves in relation to feeding rates and food competition. A qualitative model is presented here to predict how food depletion time, abundance of food patches within a group, and the presence of prior knowledge of feeding sites affect the payoffs of different within-group spatial positions for dominant and subordinate animals. In general, when feeding on small abundant food items, individuals at the front edge of the group should have higher foraging success. When feeding on slowly depleted, rare food items, dominants will often have the highest feeding rates in the center of the group. Between these two extreme points of a continuum, an individual's optimal spatial position is predicted to be influenced by an additional combination of factors, such as group size, group spread, satiation rates, and the presence of producer-scrounger tactics.     

Distribution of Gammarus lacustris Sars (Amphipoda, Gammaridae) in Lake Shira (Khakasia, Siberia) and laboratory study of its growth characteristics

Spatial distribution of Gammarus lacustris in Lake Shira and growth of young specimens that feed on lake biota were studied. The amphipods have been shown to inhabit the littoral, sublittoral and the upper aphytal zones of the lake on stony-sandy soil and silted sand. The young and adult individuals stay apart. The young live in submerged or semi-submerged vegetation in the littoral, the adults in the sublittoral and upper aphytal zones. Maximum density of amphipods was encountered in the areas influenced by human activity.The feeding experiments revealed that the lake plankton is a more important food source for G. lacustris than any other food species. The specific growth rate measured was 0.039 d^–1, with a length increment 0.095 mm d^–1.     

Usefulness of multiple chalk‐based food colorings for inducing better gene silencing by feeding RNA interference in planarians

Planarians have become widely recognized as one of the major animal models for regeneration studies in invertebrates. To induce RNA interference (RNAi) by feeding in planarians, the widely accepted protocol is one in which animals undergo two or three feedings of food containing double‐stranded RNA (dsRNA) plus visible food coloring (e.g., blood) for confirmation of feeding by individual animals. However, one possible problem is that incorporated food coloring is often retained within the gut for several days, which makes it difficult to confirm the success of each round of dsRNA feeding based on the difference of the color density within the gut before and after feeding. As a consequence, the difference of appetite levels among individuals undergoing dsRNA feeding leads to phenotypic variability among them due to insufficient knockdown. In our attempts to overcome this problem, we have developed a novel method for achieving robust confirmation of the success of dsRNA feeding in individuals fed multiple times by means of including a combination of three different colored chalks (pink, yellow and blue) as food coloring. Notably, we found that this method is superior to the conventional method for positively marking individuals that actively consumed the dsRNA‐containing food during four times of once‐daily feeding. Using these selected animals, we obtained stable and sufficiently strong RNAi‐induced phenotypes. We termed this improved multi‐colored chalk‐spiked method of feeding RNAi “Candi” and propose its benefits for gene function analysis in planarians.     

Effect of feeding frequency on food intake and growth of Arctic charr, Salvelinus alpinus L.

Social interactions and dominance hierarchy effects are important factors governing rates of growth of Arctic charr, Salvelinus alpinus L. The effects of hierarchy were increased as access to food became more restricted, i.e. feeding frequency was reduced, but these effects could not be attributed to direct competition for food since fish were fed to satiation at each feeding period. The results suggest that, whilst some fish on the restricted feeding regime were able to maintain good rates of growth, feeding by the majority of the fish was inhibited by the presence of larger individuals. Due to the importance of these hierarchy effects it was not possible to demonstrate physiological adaptations in fish allowed infrequent access to food.     

Feeding rate as valuable information in primate feeding ecology

In this review I outline studies on wild non-human primates using information on feeding rate, which is defined as the food intake per minute on a dry-weight basis; further, I summarize the significance of feeding rate in primate feeding ecology. The optimal foraging theory has addressed three aspects of animal feeding: (1) optimal food patch choice, (2) optimal time allocation to different patches, and (3) optimal food choice. In order to gain a better understanding of these three aspects, the feeding rate itself or its relevance indices (e.g., rates of calorie and protein intake) could be appropriate measures to assess the quality of food and food patches. Moreover, the feeding rate plays an essential role in estimation of total food intake, because it varies greatly for different food items and the feeding time is not a precise measure. The feeding rate could also vary across individuals who simultaneously feed on the same food items in the same food patch. Body size-dependent and rank-dependent differences in the feeding rate sometimes cause individuals to take strategic behavioral options. In the closing remarks, I discuss the usefulness of even limited data on feeding rate obtained under adverse observational conditions in understanding primate feeding ecology.     

Size-dependent flight initiation by a lotic mayfly in response to a predatory fish

1. Laboratory experiments were performed to determine whether flight initiation by lotic mayfly larvae of Baetis tricaudatus exposed to the longnose dace ( Rhinichthys cataractae ) is affected by an individual's size and its physiological state.
2. We used a three factorial ANOVA design to test whether flight initiation distances (FIDs) were affected by Baetis size (small, large), starvation level (low, high), and the length of a predator-free foraging period (short, long).
3. FIDs were significantly affected by the interaction between Baetis size and starvation level, and the main level effects of starvation and Baetis size. FIDs of small larvae were unaffected by starvation level and the length of the predator-free feeding period, whereas starvation reduced FIDs of large larvae 2-fold.
4. Subsequent experiments showed that size-dependent differences in FIDs could not be attributed to size-specific differences in the economics of rates of food intake or gut evacuation. For both small and large larvae, food intake rates declined with increasing time spent foraging and both small and large larvae consumed the majority (70–75%) of food within the first 3 h of the 12-h feeding period. Similarly, gut evacuation declined with increasing starvation time and rates of evacuation did not differ between small and large larvae. These data suggest that size-dependent differences in flight initiation by Baetis larvae do not involve the energetics of food intake or processing.     

Gut evacuation of walleye pollock larvae in response to feeding conditions

Gut residence times of first-feeding walleye pollock larvae were measured at 6°C in continuous and discontinuous feeding regimes. Larvae fed tagged copepod prey evacuated their guts more quickly in continuous feeding as compared to the discontinuous feeding treatment. The mean gut residence time was estimated to be 5.0 h for larvae feeding continuously. Gut evacuation by larvae fed tagged prey and then isolated without food (discontinuous treatment) was slower and more variable, with an estimated gut residence time exceeding 8.0 h. Field and laboratory observations suggest that the larval fish gut may be modeled as an intermittent plug-flow reactor (PFR) in response to diel feeding patterns. The cyclical nature of gut dynamics has implications for gut content analyses and the estimation of daily food rations.     

The ingestion in wolf spiders II. The expression of degree of hunger and amount of ingestion in relation to spider's hunger

To express the degree of hunger during both of feeding and unfeeding periods of spiders, equations for the amount of ingestion and food disappearance from the gut were presented using three components: capacity of gut, rate of ingestion and rate of food disappearance. The degree of hunger was expressed by the rate of unfilled capacity of gut to the capacity of gut based on these equations. The rates of ingestion and food disappearance were estimated from the results of experiment with a wolf spider, Pardosa laura. The equations obtained well applied to the experimental result. By changing values of these rates, it was revealed that the ratio of rates of ingestion to food disappearance determine the amount of ingestion.     

Does excess dietary carbon affect respiration of <Emphasis Type="Italic">Daphnia</Emphasis>?

Like many invertebrate herbivores, Daphnia frequently face diets with excess carbon (C) relative to elements like phosphorus (P), and with limited ability to store C-rich compounds. To cope with this relative surplus of C they may either regulate the net uptake of C or dispose of excess assimilated C via increased release of dissolved organic carbon or CO₂. Here we investigate whether juvenile Daphnia magna use respiration as a means of stoichiometrically regulating excess C. Growth rate and respiration were measured under different algal food qualities (P-replete and P-depleted algae). Growth rate was strongly reduced by P-depleted food, implying a stoichiometric disposal of excess ingested C. Respiration rates of feeding animals were measured after short- (0.5 h), medium- (12 h) and long- (five days) term acclimation to P-limited food. The respiration rates of animals during active feeding were not affected by the acclimation period per se, whereas food quality had a significant effect; respiration rates of feeding animals increased slightly in individuals receiving low-P food under all acclimation regimes. Respiration was also measured on nonfeeding and fasting animals that had been acclimated for five days to P-limited food. Respiration rates of these animals were strongly affected by feeding conditions but not by food quality; feeding individuals had higher respiration rates than those deprived of food, which again had higher respiration than fasting animals. Although animals grown on low-P food had strongly reduced growth and thus were expected to have decreased respiration rates due to reduced growth-related costs, this seems to be canceled out by increased stoichiometric respiration under P-deficiency. These results indicate that D. magna partly releases excess C as CO₂, but other means of stoichiometric regulation most likely add to this.     

Size-dependent flight initiation by a lotic mayfly in response to a predatory fish

1. Laboratory experiments were performed to determine whether flight initiation by lotic mayfly larvae of Baetis tricaudatus exposed to the longnose dace ( Rhinichthys cataractae ) is affected by an individual's size and its physiological state.
2. We used a three factorial ANOVA design to test whether flight initiation distances (FIDs) were affected by Baetis size (small, large), starvation level (low, high), and the length of a predator-free foraging period (short, long).
3. FIDs were significantly affected by the interaction between Baetis size and starvation level, and the main level effects of starvation and Baetis size. FIDs of small larvae were unaffected by starvation level and the length of the predator-free feeding period, whereas starvation reduced FIDs of large larvae 2-fold.
4. Subsequent experiments showed that size-dependent differences in FIDs could not be attributed to size-specific differences in the economics of rates of food intake or gut evacuation. For both small and large larvae, food intake rates declined with increasing time spent foraging and both small and large larvae consumed the majority (70–75%) of food within the first 3 h of the 12-h feeding period. Similarly, gut evacuation declined with increasing starvation time and rates of evacuation did not differ between small and large larvae. These data suggest that size-dependent differences in flight initiation by Baetis larvae do not involve the energetics of food intake or processing.     

The energetics of coprophagy: a theoretical analysis

Lagomorphs, many rodents and some other small mammals eat their faeces during the part of the day when they are not foraging for fresh food. One of the possible benefits of this habit of coprophagy is that it may enable them to extract more energy from their food. A computer model is used to assess the likely benefits and explore their relationships to food, feeding rate and gut morphology. The predicted benefits are much larger for hindgut fermenters than for foregut fermenters, and especially large for hindgut fermenters with relatively small fermentation chambers. They are larger for poor foods (with lower proportions of cell contents) than for richer ones. At low feeding rates the energetic advantage of coprophagy may disappear if the faeces from food eaten during one feeding period emerge largely during the next, but this can be avoided by adjusting the rate of passage of gut contents during the intervening rest period.     

Feeding rates of Balloniscus sellowii (Crustacea, Isopoda, Oniscidea): the effect of leaf litter decomposition and its relation to the phenolic and flavonoid content

The goal of this study was to compare the feeding rates of Balloniscus sellowii on leaves of different decomposition stages according to their phenolic and flavonoid content. Leaves from the visually most abundant plants were offered to isopods collected from the same source site. Schinus terebinthifolius,the plant species consumed at the highest rate, was used to verify feeding rates at different decomposition stages. Green leaves were left to decompose for one, two, or three months, and then were offered to isopods. The total phenolic and flavonoid contents were determined for all decomposition stages. Consumption and egestion rates increased throughout decomposition, were highest for two-month-old leaves, and decreased again in the third month. The assimilation rate was highest for green leaves. The mode time of passage through the gut was two hours for all treatments. Ingestion of leaves occurred after two or three days for green leaves, and on the same day for one-, two- and three-month-old leaves. The speed of passage of leaves with different decomposition stages through the gut does not differ significantly when animals are fed continuously. However, it is possible that the amount retained in the gut during starvation differs depending on food quality. The digestibility value was corrected using a second food source to empty the gut of previously ingested food, so that all of the food from the experiment was egested. The digestibility value was highest for green leaves, whereas it was approximately 20% for all other stages. This was expected given that digestibility declines during decomposition as the metabolite content of the leaves decreases. The phenolic content was highest in the green leaves and lowest in three-month-old leaves. The flavonoid content was highest in green leaves and lowest after two months of decomposition. Animals ingested more phenolics when consumption was highest. The estimated amount of ingested flavonoids followed the same trend as assimilation rate. Flavonoids accounted for a large portion of total phenolics, and the estimated amount of flavonoids consumed was similar for one-, two- and three-month-old leaves. Our results suggest that the high phenolic and flavonoid concentrations in green leaves are feeding deterrents. Isopods may discriminate among concentrations of flavonoids and modify their consumption rates to maintain their intake of flavonoids when ingesting leaves with lower flavonoid content.     

Filtration and digestion responses of an elementally homeostatic consumer to changes in food quality: a predictive model

In the study of the stoichiometric relationship between autotrophs and herbivores, attention has been largely focused on effects of the encountered mismatch between needs and supplies of an element on herbivore growth and ecosystem processes. Herbivore adaptation to poor food quality has rarely been investigated. This study presents a predictive model of feeding, assimilation, digestion and excretion of Daphnia facing a dietary deficiency in phosphorus. Biochemical compounds in the food were divided into phosphorous and non-phosphorus compounds. It was assumed that Daphnia is able to differently assimilate both types of compounds by regulation of target specific digestive enzymes. Feeding rate was regulated by optimal gut residence time of food particles, and assimilation efficiency by gut residence time and optimal secretion of both classes of gut enzymes. The model predicted the optimal strategy for a consumer facing an elementally imbalanced diet: (1) increase the ingestion rate, and (2) increase the secretion rate of both classes of gut enzymes. It resulted in decreased C and nutrient assimilation efficiencies, increased C feeding costs, and reduced growth rate. Sensitivity analysis showed that these predictions were qualitatively not influenced by parameter values. An alternative model was tested that includes an additive term allowing the direct excretion of C assimilated in excess. Results showed that this strategy is not optimal for the consumer growth rate. In conclusion, the model supports the hypothesis that carbon ingested in excess may generate energy that can be used to obtain more nutrients by increased feeding rate.     

Food resource partitioning among Arctic sublittoral lysianassoid amphipods in summer

Five species of lysianassoid amphipods (Anonyx nugax, Anonyx sarsi, Onisimus caricus, Onisimus edwardsii, Orchomenella minuta) co-occur in the shallow sublittoral of Kongsfjorden (Svalbard, Arctic), share similar functional attributes, and exploit a common food base. All species are known to be necrophagous; however, this study revealed through combination of gut-content analysis with the morphological characteristics of their mandibles and available information on feeding ecology, the complexity of their trophic strategies. Only mature individuals of A. nugax are true scavengers; immature individuals of this species feed on both animal and non-animal food. A. sarsi and O. caricus are scavengers and predators; the former prefers polychaetes, while the latter relies mostly on zooplankton. O. edwardsii is an omnivorous feeder, and the set of its primary food consists of carrion, crustaceans, and algae. O. minuta is a detrivorous–carnivorous species. These results indicate that niche overlap among these closely related species is reduced, not only by spatial segregation, but also by the exploitation of different components of food resources.     

Feeding and digestion by the mussel Mytilus edulis L. (Bivalvia: Mollusca) in mixtures of silt and algal cells at low concentrations

Mussels Mytilus edulis L. from two populations were exposed for 2 days or 2 wk to mixtures of silt and the diatom Phaeodactylum tricomutum Bohlin before measuring rates of feeding, the passage time for food in the gut, absorption efficiency and metabolic rate. Experimental diets were set up to span the range of organic content in seston from the natural habitats (% organic matter by weight: 7–55%), but were less than the total levels of natural seston. Absorption efficiencies were adequately modelled in an exponential relationship to food quality and to gut passage time, although at high proportional silt concentrations metabolic faecal losses led to negative net rates of absorption. Over short-term exposures the scope for growth was a simple function of food quality, and the nutritional quality of the diet was best expressed as organic content per unit volume of particles. Over a period of 2 wk physiological acclimation occurred, across all levels of experimental food quality (which were as low as 10% organic matter by weight), resulting in positive growth potential. Relevant mechanisms of compensation include increased rates of ingestion, increased absorption efficiency and an apparent increase in digestive capacity, estimated here as gut fullness. In experiments in which natural diets are simulated by adding silt to phytoplankton cells, the consequences for net rates of absorption depend on the balance between mean particle size and organic content per unit volume. Calculations show how, in some circumstances, growth may be enhanced by the addition of small silt particles to living phytoplankton cells.     

Food intake, conversion efficiency, and feeding behaviour of tobacco hornworm caterpillars given artificial diet of varying nutrient and water content

ABSTRACT Fifth stadium tobacco hornworm caterpillars, Manduca sexta (L.), given artificial diet diluted to varying extents with either cellulose or water compensated for the food's reduced nutrient content by eating more of it. This compensation was, however, in most cases not sufficient to maintain normal growth rates. When the water content of the diet was reduced, the insects ate less than the usual fresh weight of food but maintained their intake of nutrients. Nevertheless, growth rate was impaired. The insects were better able to compensate for dilution of their food with water than with cellulose. The efficiency of conversion of ingested food (ECI) was decreased when the diet was adulterated with cellulose. At moderate dilution (50% nutrient) this was due mostly to decreased approximate digestibility (AD), but at greater dilution (25% and 10% nutrient content) the efficiency of conversion of digested food (ECD) was decreased. ECI was maintained when the water content of the diet was increased to give 50% nutrient concentration, but was decreased when water content was changed more radically (200%, 25% and 10% nutrient diets). This was due mostly to increased metabolic costs (decreased ECD) in all cases. The retention time of food in the gut was progressively decreased (i.e. speed of passage was increased) as nutrients were replaced by cellulose. By contrast, dilution of the diet with water resulted in only slight changes in retention time, except at extreme dilution (10% nutrient content) when retention time was reduced. Compensation of food intake was achieved by spending more (or less) time eating. Video analysis of feeding behaviour showed that there were significant changes in the length of feeding bouts and of interfeed gaps when caterpillars fed on diets of altered composition. For diets diluted with cellulose, changes in bout length and bout frequency contributed substantially to the increased time spent feeding on the adulterated food. For diets diluted with water, however, almost all of the compensatory change in behaviour was due to increased bout length, with bout frequency affected only slightly. This suggests that volumetric feedback contributes principally to the termination of feeding bouts in caterpillars, while nutrient flow may affect both the initiation and termination of feeding.     

Field estimates of feeding rate for Gammarus pseudolimnaeus (Crustacea: Amphipoda) in the Credit River, Ontario

SUMMARY. The feeding rate of G. pseudolimnaeus was measured monthly for 7 months in the field by monitoring the decline in weight of gut contents when the amphipod was starved. This decline was modelled by an exponential regression of weight on time. As the amphipods appeared to be continuous feeders, feeding rate was calculated by multiplying the dry weight of a full gut by the specific rate of emptying, i.e., the slope of the exponential regression. Specific rate of emptying was independent of animal size, but increased with temperature. Therefore, food has a longer period in which to be digested at low temperatures, which suggests that assimilation efficiency may increase.
However, the assimilation efficiency of amphipods feeding on decaying maple leaves in the laboratory was only 10% and did not vary with temperature. Ingestion and egestion rates were measured in the laboratory by weighing amounts eaten and defecated. The turnover time of the contents of a full gut in the laboratory often agreed very well with turnover time measured in the field, i.e., the reciprocal of the specific rate of emptying, thus confirming the use of an exponential regression.