Feeding Responses of Hatchery-Reared Gilthead Sea Bream (Sparus aurata L.) to a Commercial Diet and Natural Prey Items

Many fish species have evolved feeding mechanisms and behaviours enabling them to feed on specific prey. However, such mechanisms may not be optimal for feeding on commercial-pelleted diets in aquaculture. Gilthead sea bream chew and occasionally eject pellets or parts of pellets from the mouth when feeding on commercial diets. This may result in an increase in nutritional waste from the intensive culture of this species. In this study we examined the prevalence of this food processing behaviour in two sizes of sea bream, feeding on three types of natural prey items in comparison to a commercial pellet, to give an insight into the circumstances in which excess chewing and ejection of food items from the mouth occurred. These included two hard-textured food items (commercial pellet and hard-shelled prey) and two soft-textured food items (larvae and small crustacean). Both sizes of sea bream frequently consumed the soft-textured food types, however large sea bream also frequently consumed hard-textured pellets. Hard-textured food required longer handling times and elicited more chewing and the ejection of food items from the mouth. These results suggest that future investigations on the food processing behaviour and consequent waste when fed commercial diets differing in texture could give an insight into improving diets and feeding efficiency for intensively cultivated gilthead sea bream.     

Feeding Responses of Hatchery-Reared Gilthead Sea Bream (Sparus aurata L.) to a Commercial Diet and Natural Prey Items

Many fish species have evolved feeding mechanisms and behaviours enabling them to feed on specific prey. However, such mechanisms may not be optimal for feeding on commercial-pelleted diets in aquaculture. Gilthead sea bream chew and occasionally eject pellets or parts of pellets from the mouth when feeding on commercial diets. This may result in an increase in nutritional waste from the intensive culture of this species. In this study we examined the prevalence of this food processing behaviour in two sizes of sea bream, feeding on three types of natural prey items in comparison to a commercial pellet, to give an insight into the circumstances in which excess chewing and ejection of food items from the mouth occurred. These included two hard-textured food items (commercial pellet and hard-shelled prey) and two soft-textured food items (larvae and small crustacean). Both sizes of sea bream frequently consumed the soft-textured food types, however large sea bream also frequently consumed hard-textured pellets. Hard-textured food required longer handling times and elicited more chewing and the ejection of food items from the mouth. These results suggest that future investigations on the food processing behaviour and consequent waste when fed commercial diets differing in texture could give an insight into improving diets and feeding efficiency for intensively cultivated gilthead sea bream.     

Experimental Predictions of The Functional Response of A Freshwater Fish

The functional response is the relationship between the feeding rate of an animal and its food density. It is reliant on two basic parameters; the volume searched for prey per unit time (searching rate) and the time taken to consume each prey item (handling time). As fish functional responses can be difficult to determine directly, it may be more feasible to measure their underlying behavioural parameters in controlled conditions and use these to predict the functional response. Here, we tested how accurately a Type II functional response model predicted the observed functional response of roach Rutilus rutilus, a visually foraging fish, and compared it with Type I functional response. Foraging experiments were performed by exposing fish in tank aquaria to a range of food densities, with their response captured using a two‐camera videography system. This system was validated and was able to accurately measure fish behaviour in the aquaria, and enabled estimates of fish reaction distance, swimming speed (from which searching rate was calculated) and handling time to be measured. The parameterised Type II functional response model accurately predicted the observed functional response and was superior to the Type I model. These outputs suggest it will be possible to accurately measure behavioural parameters in other animal species and use these to predict the functional response in situations where it cannot be observed directly.     

Gastric emptying and the return of appetite in juvenile turbot, Scophthalmus maximus L., fed on artificial diets

Gastric emptying time in Scophthalmus maximus , when fed friable artificial pellets based on fishmeal, is composed of two phases:
(a) a delay time (t_d) during which the meal forms a bolus and which shortens with temperature, and
(b) an emptying phase (duration t_end ) which varies with meal size ( S ), body weight ( W ) and temperature (71 according to:
(where t _end is in h, S is in g, W is in g and T is °C). During the emptying phase, stomach contents decrease curvilinearly according to:
(where S_t , & S_o is in g and t is in h) in which the instantaneous digestion rate, K , varies with fish weight and temperature as:
Food pellets were prepared which remained separate and did not form a bolus in the stomach; K increased if a given meal size was subdivided to increase surface area. If meal size was increased by ingestion of identical pellets, K decreased. After a satiation meal, appetite in young turbot returns in direct relation to the degree of stomach emptiness. When food is regularly available, young turbot feed steadily at a rate which maintains their stomachs at c. 85% maximum fullness. When trained to use demand feeders, the fish interact as a group to feed rhythmically, but feeding rate falls 33% to only two-thirds of the previous rate since stomach fullness, and hence digestion rate (g h⁻¹), is maintained at a lower level. Reduction in dietary energy density below 1 kCal g⁻¹ increases gastric emptying rate and the turbot demonstrate partial compensation by increasing food intake. On energy-rich diets, protein nitrogen and energy assimilation efficiencies remain high (97 5% and 91% respectively) irrespective of feeding rate and frequency.     

Patterns of habitat and invertebrate diet overlap between tiger salamanders and ducks in prairie potholes

During the breeding season, migratory waterfowl are attracted to wetlands characterized by high macroinvertebrate availability. Many of these prairie potholes are fishless and this apparent void is filled, at least partially, by tiger salamanders. Based on gut contents from 98 tiger salamanders and published diet data from over 1500 ducks, we show that there is general overlap in diet between both larval and adult tiger salamanders and 10 duck species. Furthermore, when the ducks were split into foraging guilds and compared with tiger salamanders, prey type overlap was 1.7 times higher and prey size was 1.8 times higher with dabbling ducks than diving ducks. Field surveys show that tiger salamander density is more highly correlated with diving duck density across potholes than dabbling duck density. Tiger salamanders have higher diet overlap with dabbling ducks than diving ducks whereas tiger salamanders have higher spatial overlap with diving ducks than dabbling ducks suggesting that these consumers coarsely partition diet and habitat resources. It has been reported that tiger salamanders have specialized diets that are associated with foraging preferences for benthic habitats. This view is too narrow: in southwestern Manitoba, Canada, tiger salamanders are more general consumers with diets more like dabbling ducks that forage mostly in planktonic and littoral habitats. Our results suggest that dabbling and diving ducks are, to different extents, liable to the effects of indirect interactions, specifically competition for common prey, with tiger salamanders.     

Diet segregation between two sympatric 'small'Barbus spp: an experimental study of mechanisms

Gut contents of two co‐occurring species of 'small' diploid barbs (<10  L _F cm) in Lake Tana revealed that zooplankton is the major diet component for B. tanapelagius (75% based on volume), but less prominent in B. humilis (40%). Functional response experiments in the laboratory were conducted to elucidate the mechanisms causing this difference. The type of functional response by the two 'small' barbs under different microcrustacean zooplankton densities (10, 20, 40, 60 and 80 ind.l⁻¹) was examined. The functional response of B. tanapelagius to increasing prey densities corroborates with Holling Type II model, whereas B. humilis exhibits a Type III functional response. Predation rate is higher for B. tanapelagius at low zooplankton density (<40 ind.l⁻¹) and equals the level of B. humilis at higher densities (>40 ind.l⁻¹). This suggests that at lower zooplankton densities B. humilis is a less efficient forager on zooplankton prey items than B. tanapelagius . In Lake Tana average zooplankton density is relatively low (<35 ind.l⁻¹). Under these food conditions, B. humilis is forced to feed on other food items (e.g. benthic invertebrates), whereas B. tanapelagius primarily feeds on zooplankton. The feeding potentials of the two 'small' barbs, as deduced from their morphology explain their different performances and their segregation in space and food resources.     

Effects of acute and chronic temperature changes on the functional responses of the dogfish <Emphasis Type="Italic">Scyliorhinus canicula</Emphasis> (Linnaeus, 1758) towards amphipod prey <Emphasis Type="Italic">Echinogammarus marinus</Emphasis> (Leach, 1815)

Predation is a strong driver of population dynamics and community structure and it is essential to reliably quantify and predict predation impacts on prey populations in a changing thermal landscape. Here, we used comparative functional response analyses to assess how predator-prey interactions between dogfish and invertebrate prey change under different warming scenarios. The Functional Response Type, attack rate, handling time and maximum feeding rate estimates were calculated for Scyliorhinus canicula preying upon Echinogammarus marinus under temperatures of 11.3 °C and 16.3 °C, which represent both the potential daily variation and predicted higher summer temperatures within Strangford Lough, N. Ireland. A two x two design of “Predator Acclimated”, “Prey Acclimated”, “Both Acclimated”, and “Both Unacclimated” was implemented to test functional responses to temperature rise. Attack rate was higher at 11.3 °C than at 16.3 °C, but handling time was lower and maximum feeding rates were higher at 16.3 °C. Non-acclimated predators had similar maximum feeding rate towards non-acclimated and acclimated prey, whereas acclimated predators had significantly higher maximum feeding rates towards acclimated prey as compared to non-acclimated prey. Results suggests that the predator attack rate is decreased by increasing temperature but when both predator and prey are acclimated the shorter handling times considerably increase predator impact. The functional response of the fish changed from Type II to Type III with an increase in temperature, except when only the prey were acclimated. This change from population destabilizing Type II to more stabilizing Type III could confer protection to prey at low densities but increase the maximum feeding rate by Scyliorhinus canicula in the future. However, predator movement between different thermal regimes may maintain a Type II response, albeit with a lower maximum feeding rate. This has implications for the way the increasing population Scyliorhinus canicula in the Irish Sea may exploit valuable fisheries stocks in the future.     

Feeding dynamics of sturgeon fingerlings (Acipenseridae) depending on food concentration and stocking density

Sturgeon fingerlings of 20.0 g at a water temperature of 18.20° C, fed for 130 min and consumed 6.5% of their body weight when the food concentration was 2 g m⁻². At 38 g m⁻² the fingerlings stopped feeding after 45 min but consumed 9.0% of their body weight. As satiation approached the intervals between successive food intakes increased and feeding rate decreased. The rate of feeding was inversely proportional to the amount of food already consumed, and an equation describing this relationship is given. During the first minutes of feeding the ration size was directly proportional to food concentration but with further feeding the relationship changed. The equations of Ivlev and Rashevsky were close approximations to the relationship between ration size and food concentration. At a density of 12 individuals/m² the feeding duration and ration size was low in comparison to single fingerlings and to fish fed at higher densitites.     

Optimal foraging: food patch depletion by ruddy ducks

Summary I studied the foraging behavior of ruddy ducks (Oxyura jamaicensis) feeding on patchily distributed prey in a large (5-m long, 2-m wide, and up to 2-m deep) aquarium. The substrate consisted of a 4x4 array of wooden trays (1.0-m long, 0.5-m wide, and 0.1-m deep) which contained 6 cm of sand. Any tray could be removed from the aquarium and loaded with a known number of prey. One bird foraged in the aquarium at a time; thus, by removing a food tray after a trial ended and counting the remaining prey, I calculated the number of prey consumed by the bird. I designed several experiments to determine if ruddy ducks abandoned a food patch in a manner consistent with the predictions of a simple, deterministic, patch depletion model. This model is based on the premise that a predator should maximize its rate of net energy intake while foraging. To accomplish this, a predator should only remain in a food patch as long as its rate of energy intake from that patch exceeds the average rate of intake from the environment. In the majority of comparisons, the number of food items consumed by the ruddy ducks in these experiments was consistent with the predictions of the foraging model. When the birds did not forage as predicted by the model, they stayed in the patch longer and consumed more prey than predicted by the model. An examination of the relation between rate of net energy intake and time spent foraging in the food patch indicated that by staying in a patch longer than predicted, the ruddy ducks experienced only a small deviation from maximum rate of net energy intake. These results provided quantitative support for the prediction that ruddy ducks maximize their rate of net energy intake while foraging.     

Testing the validity of functional response models using molecular gut content analysis for prey choice in soil predators

Analysis of predator–prey interactions is a core concept of animal ecology, explaining structure and dynamics of animal food webs. Measuring the functional response, i.e. the intake rate of a consumer as a function of prey density, is a powerful method to predict the strength of trophic links and assess motives of prey choice, particularly in arthropod communities. However, due to their reductionist set‐up, functional responses, which are based on laboratory feeding experiments, may not display field conditions, possibly leading to skewed results. Here, we tested the validity of functional responses of centipede predators and their prey by comparing them with empirical gut content data from field‐collected predators. Our predator–prey system included lithobiid and geophilomorph centipedes, abundant and widespread predators of forest soils and their soil‐dwelling prey. First, we calculated the body size‐dependent functional responses of centipedes using a published functional response model in which we included natural prey abundances and animal body masses. This allowed us to calculate relative proportions of specific prey taxa in the centipede diet. In a second step, we screened field‐collected centipedes for DNA of eight abundant soil‐living prey taxa and estimated their body size‐dependent proportion of feeding events. We subsequently compared empirical data for each of the eight prey taxa, on proportional feeding events with functional response‐derived data on prey proportions expected in the gut, showing that both approaches significantly correlate in five out of eight predator–prey links for lithobiid centipedes but only in one case for geophilomorph centipedes. Our findings suggest that purely allometric functional response models, which are based on predator–prey body size ratios are too simple to explain predator–prey interactions in a complex system such as soil. We therefore stress that specific prey traits, such as defence mechanisms, must be considered for accurate predictions.     

Increased searching and handling effort in tall swards lead to a Type IV functional response in small grazing herbivores

Understanding the functional response of species is important in comprehending the species’ population dynamics and the functioning of multi-species assemblages. A Type II functional response, where instantaneous intake rate increases asymptotically with sward biomass, is thought to be common in grazers. However, at tall, dense swards, food intake might decline due to mechanical limitations or if animals selectively forage on the most nutritious parts of a sward, leading to a Type IV functional response, especially for smaller herbivores. We tested the predictions that bite mass, cropping time, swallowing time and searching time increase, and bite rate decreases with increasing grass biomass for different-sized Canada geese (Branta canadensis) foraging on grass swards. Bite mass indeed showed an increasing asymptotic relationship with grass biomass. At high biomass, difficulties in handling long leaves and in locating bites were responsible for increasing cropping, swallowing, and searching times. Constant bite mass and decreasing bite rate caused the intake rate to decrease at high sward biomass after reaching an optimum, leading to a Type IV functional response. Grazer body mass affected maximum bite mass and intake rate, but did not change the shape of the functional response. As grass nutrient contents are usually highest in short swards, this Type IV functional response in geese leads to an intake rate that is maximised in these swards. The lower grass biomass at which intake rate was maximised allows resource partitioning between different-sized grazers. We argue that this Type IV functional response is of more importance than previously thought.     

Comparisons of the diets and reproductive performances of two sympatric rotifers, Asplanchna girodi and Asplanchna priodonta

SUMMARY. 1. Diet composition and the production of embryos were measured in Asplanchna girodi and Asplanchna priodonta collected through 3 years in the San Joaquin-Sacramento delta of California. The two species are sympatric in low to moderate densities for 5 months of the year.
2. Despite marked differences in the structure of the trophi, both species consumed the same array of prey species. Percentage composition of the diet varied both intra- and interspecifically. The food niches of the two species appear to differ little from one another.
3. Individuals of Asplanchna girodi had a larger mean number of prey in the gut per individual than did those of A. priodonta , due to their larger size. Per unit volume, each species prey gathering performance was the same. Asplanchna priodonta produced more embryos per calculated unit volume of prey ingested than did A. girodi .
4. During spring, when both species were present in the plankton, A. priodonta did not capture Size Class 2 and 5 animals (volumes 0.09–1.0 μm³×10⁶ and 3.0–3.5 μm³X10⁶ respectively) nor large Synchaeta . Both these prey classes were in the diet during the summer and autumn months. Several hypotheses are examined as possible explanations for this phenomenon.     

Experimental evidence of cannibalism and prey specialization in Arctic charr,Salvelinus alpinus

Synopsis Use of a radiographic technique enabled the study of prey selection and individual specialization in Arctic charr, Salvelinus alpinus, fed with small charr and dry pellets under laboratory conditions. Both naive and experienced fish (mean weight 475 g, mean length 34.9 cm), selected the smallest individuals when offered juvenile charr (6–16 cm) as prey. The selected prey were, on average, 22% of the predator length. Cannibalism appeared to involve individual specialization, since when groups of large charr were given the opportunity to feed on juveniles one day every two weeks, the same individuals were cannibalistic throughout the sampling period of two months. When large charr were presented with alternate cycles of different food types consisting of dry pellets and fish prey, the charr exhibited a strong and consistent feeding specialization with three distinct groups being recognised: cannibals, pellet eaters and non-feeding fish.     

Factors affecting species number and density of dabbling duck guilds in North Europe

We addressed how species number and pair density in guilds of co-existing species is related to habitat structure, and to the abundance and diversity of food resources using the assemblage of seven species of dabbling ducks (genus Anas ) breeding in 60 lakes distributed over six regions in temperate north Europe
Partial correlation and multiple regression revealed that species richness was best predicted by habitat structural diversity as indexed by a principal component analysis based on 18 vegetation and lake characteristics, and by the abundance of aquatic and emergent prey We found no effect of lake size or prey size diversity on species richness Pair density was correlated with the percentage of shoreline with horsetails ( Equisetum ), by habitat structural diversity and by the abundance of emergent invertebrate prey Neither prey size diversity nor abundance of aquatic prey correlated with pair density Species richness and pair density in North European duck guilds vary both with habitat structure and prey availability     

Nucleic acid contents and growth of first-feeding walleye pollock larvae in response to prey densities typical of sub-Arctic ecosystems

The growth, nucleic acid and protein contents of walleye pollock Theragra chalcogramma larvae reared at prey densities of 10, 30, 50, and 500 prey 1^-1 were measured for the first 9 days after the feeding initiation at 6° C. Incremental growth rates of larvae (mm day^-1) were low and variable for the first 7 days after feeding initiation. Growth rates and rates of RNA, DNA, and protein accumulation by larvae reared at 500 prey 1^-1 were positive while those of larvae reared at the lower prey levels did not differ significantly from zero. The RNA/DNA ratio was variable and exhibited no significant trend among food treatments. Estimates of instantaneous protein growth rates ranged from - 6·7 to 13·2% day^-1 at food densities of 10 and 500 prey 1^-1, respectively, and were moderately correlated with larval RNA/DNA ratios ( r = 0·628). The results suggest that in situ protein growth rates of first-feeding pollock larvae may be influenced by prey fields within the range of ambient food densities reported for sub-Arctic ecosystems.     

Effects of food density and temperature on feeding and growth of young inland silversides (Menidia beryllina)

Food consumption and growth rates of 7–28-day-old Menidia beryllina were measured in response to natural ranges of temperature and prey availability. Feeding level, temperature and age all had significant effects on growth rate, although the effect of feeding level explained most of the variance. Feeding level also had a significant effect on gross growth efficiency, but temperature and age did not. Absolute growth rates (mg per day) increased dramatically with temperature, feeding level, and age. Variability in growth was greatest for fish feeding at the lowest feeding level. For a given fish weight, temperature had a positive effect on consumption rate, and maximum consumption ( C _max) of any treatment combination reached 75% body weight per day. Maximum growth rate was estimated at 24.6% body weight per day, and gross growth efficiency reached an estimated maximum of 0.375 at an ingestion rate of 25% body weight per day. Starved larvae lost on average 5.4% body weight per day and larvae required 6.4% body weight food consumption per day for maintenance. Multiple regressions of feeding level, temperature, and age/size on instantaneous growth rates indicated that increases in temperature increased maintenance requirements and required that fish consume a greater proportion of C _max to attain maximum growth. Growth rates decreased with increases in temperature for fish eating a specific weight of food.     

Influence of intra‐ and interspecific variation in predator–prey body size ratios on trophic interaction strengths

1. Predation is a pervasive force that structures food webs and directly influences ecosystem functioning. The relative body sizes of predators and prey may be an important determinant of interaction strengths. However, studies quantifying the combined influence of intra‐ and interspecific variation in predator–prey body size ratios are lacking.
2. We use a comparative functional response approach to examine interaction strengths between three size classes of invasive bluegill and largemouth bass toward three scaled size classes of their tilapia prey. We then quantify the influence of intra‐ and interspecific predator–prey body mass ratios on the scaling of attack rates and handling times.
3. Type II functional responses were displayed by both predators across all predator and prey size classes. Largemouth bass consumed more than bluegill at small and intermediate predator size classes, while large predators of both species were more similar. Small prey were most vulnerable overall; however, differential attack rates among prey were emergent across predator sizes. For both bluegill and largemouth bass, small predators exhibited higher attack rates toward small and intermediate prey sizes, while larger predators exhibited greater attack rates toward large prey. Conversely, handling times increased with prey size, with small bluegill exhibiting particularly low feeding rates toward medium–large prey types. Attack rates for both predators peaked unimodally at intermediate predator–prey body mass ratios, while handling times generally shortened across increasing body mass ratios.
4. We thus demonstrate effects of body size ratios on predator–prey interaction strengths between key fish species, with attack rates and handling times dependent on the relative sizes of predator–prey participants.
5. Considerations for intra‐ and interspecific body size ratio effects are critical for predicting the strengths of interactions within ecosystems and may drive differential ecological impacts among invasive species as size ratios shift.

     

Filter-feeding dabbling ducks (Anas spp.) can actively select particles by size

The relationship between bill morphology, function and prey use among filter-feeding dabbling ducks (Anas spp.) is poorly understood. In particular, it is not clear if interspecific differences in morphology affect the retention of prey to allow prey partitioning. The size of particles retained by ducks may be determined entirely by the distance between adjacent mandibular lamellae (interlamellar distance), possibly allowing interspecific partitioning of prey by size. Alternatively, articulation of the maxilla and mandible allows ducks to increase the distance between the maxillary and mandibular lamellae (lamellar separation) so that it exceeds their interlamellar distance, possibly allowing them to selectively expel unwanted particles larger than their interlamellar distance. In contrast, if interlamellar distance alone determines the size of particles ingested, particles larger than the interlamellar distance will always be retained because lamellar spacing is fixed. When large, preferred millet and wheat seeds were mixed with small, less preferred poppy seeds, all three ducks in this investigation ingested a greater proportion of the millet and wheat seeds than the poppy seeds, even though all three seed types were larger than the ducks' interlamellar distance. The proportion of millet and poppy seeds ingested when seeds were mixed differed from the proportion expected from foraging rates on unmixed prey, indicating the ducks actively avoided poppy seeds. These results are consistent with the lamellar separation hypothesis and clearly reject the singular role of interlamellar distance in prey retention. Because lamellar separation and water filtration rate are both determined by movement of the maxilla and mandible, there may be a trade-off between particle size selection and prey ingestion rate that allows interspecific partitioning of prey by size.     

Scale and the guild functional response: Density-dependent predation varies with plot size

The predator functional response is an important mechanism determining the persistence of prey species; however, little is known about the effects of spatial scale on the functional response. We used a manipulative field experiment to quantify the effects of plot size on the guild functional response on the clam Mercenaria mercenaria, replicating the experiment in the summer in Chesapeake Bay, Virginia, and in the spring and fall in Indian River Lagoon, Florida, to examine the effects of predator and alternative prey abundance. In Virginia, the predation rate increased with both patch size and predator density, and was described by a modified sigmoid Type III functional response model that incorporated the effects of patch size. In Florida in the spring, the predator functional response was a Type III and did not vary with plot size, but in the fall it was a linear Type I at small plot sizes, and a Type III at a larger plot size. We hypothesize that the difference is primarily driven by changes in predator abundance and species between sites. In showing that the functional response can vary with plot size and season, our results indicate that small-scale experiments do not always scale up spatially or temporally. We suggest that the predictive power of such experiments may be limited by the complexity of the food web.     

Growth response of pikeperch larvae in relation to body size and zooplankton abundance

The most critical period at onset of feeding in pikeperch Stizostedion lucioperca is short (<5 days at 20° C). The larvae are sensitive to prey density during the first week of exogenous feeding. First-feeding larvae of 6.5 mm total length ( L_T ) needed prey densities of >585 prey l⁻¹ to maintain mass (C_maint), whereas 5 days older larvae of 7 mm L _T C_maint= 55 prey l⁻¹ and for 11 mm L _T larvae C_maint<10 prey 1⁻¹. Changes in specific growth rates for larvae of 7 and 11 mm were similar to a type-II functional response curve reaching a specific growth rate of 26 and 30% day⁻¹, respectively, at 1000 prey l⁻¹, whereas the 6·5 mm larvae showed a linear growth response reaching a specific growth rate of only 9% day ⁻¹ at 1000 prey l⁻¹. The results suggest that prey density is a limiting factor, which might contribute to the high variation between year-class Strengths.