The effects of temperature and meal size on the rate of gastric evacuation in perch (Perca fluviatilis) fed on fish larvae

SUMMARY. The rate of gastric evacuation in perch (Perca fluviatilis) (89–170 mm length) fed on fish larvae was studied at temperatures between 12.0 and 21.7°C. Gastric evacuation rates were usually described by an exponential function. The instantaneous rate of gastric evacuation ( R ) was constant for a large number of different meal sizes. At higher food rations, a lag phase in digestion was found during the first part of digestion, and this ration size was smaller for smaller perch (89–110 mm) than for bigger perch (120–170 mm). Below these larger meal sizes, gastric evacuation was similar for the different size classes studied. The relation between R and temperature was described by an exponential function. The effects of meal size, number of food items, fish size and temperature on the rate of gastric evacuation are discussed.     

Rate of food evacuation in roach (Rutilus rutilus) in relation to temperature, and the application of evacuation rate estimates for studies on the rate of food consumption

SUMMARY. Food evacuation in roach (total body length, range 75–150 mm) was studied at temperatures between 4.2 and 24.0 °C. The rate of food evacuation was described by an exponential function and was similar for two different food items ( Daphnia and Chaoborus ). The relation between the instantaneous rate of food evacuation ( R ) and temperature was also described by an exponential function. A model developed for estimating food consumption rates in fish was tested in a simulation experiment and there was good agreement between estimated and actual values for food consumption.     

An in situ estimate of daily food consumption and alimentary canal evacuation rates of common carp, Cyprinus carpio L.

Daily food consumption and alimentary canal evacuation rates for common carp, Cyprinus carpio L., were estimated at 4-week intervals from April to October 1979. Both the rate of food evacuation and its relationship to temperature were described by an exponential function. Gut evacuation rates ranged from 6.0% of gut contents per hour at 9°C to 32.7% h⁻¹ at 26.5°C. Temperature accounted for 72–91% of the variation in gut evacuation rate among months. Daily food consumption varied from a low of 3.87 mg g⁻¹ of fish wet wt per day in April at 14°C (0.39% of wet body wt) to a high of 40.75 mg g⁻¹ (4.08%) in August at 26.5°C. Although a greater quantity of food appeared to be consumed during the morning in spring and early summer and a greater amount in the evening after July, there was not a statistically significant time of day for peak feeding activity. The field method described here seemed to provide reasonable estimates of food consumption compared with other field and laboratory studies.     

Rates of gastric evacuation in brown trout, Salmo trutta L.

Brown trout of similar length and weight were fed a standard meal which contained a known number of food organisms of the same size-group and taxon (seven taxa were used). The weight of digestible organic matter in a trout stomach decreased exponentially with time. i.e. at a constant relative rate. At a particular water temperature, the food organisms were either evacuated from the stomach at similar rates (Group 1: Gammarus pulex, Baetis rhodani, Chironomidae, Oligochaetes) or at progressively slower rates (Group 2: Protonemura meyeri, Hydropsyche spp., Tenebrio molitor). Rates of gastric evacuation were not significantly different for food organisms of different size groups of the same taxon, or for different sized meals, or for different sizes of trout (range 20–30 cm), or for mixed and multiple meals (three meals over 16 h). Times are given for the gastric evacuation of 50%, 75%, 90% and 99% of the digestible organic matter in a meal. Starvation periods of 1, 2, 3,4 and 5 days prior to feeding did not affect evacuation rates. The rates were slightly, but not significantly, slower for starvation periods of 6 and 7 days, and were often significantly slower for starvation periods of 10, 15 and 20 days. Evacuation rates increased exponentially with increasing water temperature. It was possible to estimate both the rate and time for the gastric evacuation of different meals at water temperatures between 3–8°C and 19·1°C.     

Gastric evacuation in plaice, Pleuronectes platessa L.: effects of temperature and fish size

Using a volume dependent model of gastric evacuation, the effects of temperature and fish size were examined. Rates of gastric evacuation were unaffected by fish size but increased with increasing temperature. The relationship between maximum stomach volume and fish weight was found to be a linear one. From information of gastric evacuation rates and stomach volume, the amount of food evacuated from the stomach per day was calculated for different size classes of fish. Daily food evacuation increased in proportion to body weight to the power 0·68. Assuming these methods give a crude estimate of daily food intake, the results are discussed in relation to published work on food intake in fishes.     

Rates of gastric evacuation in piscivorous brown trout, Salmo trutta

SUMMARY. 1. The dry weight of food remaining in the stomachs of piscivorous trout decreased exponentially with time. Gastric evacuation rates increased exponentially with increasing temperature but were unaffected by predator size, meal size or type of fish prey.
2. Mathematical models were developed to estimate both the rate and time for the gastric evacuation of different meal sizes (expressed as dry weight), and were applicable to piscivorous trout of different sizes (length range 10–32 cm) feeding on trout fry or sticklebacks at different temperatures (range 5–18°C).
3. The wet weight of food in the stomachs also decreased exponentially with time, but evacuation rates both increased with temperature and decreased with increasing meal size; the latter relationship occurred because relative rates of water loss from a meal also decreased with increasing meal size. Use of wet or dry weights can therefore lead to different conclusions about the effect of meal size on evacuation rates.
4. When piscivorous trout were fed three consecutive meals of varying size, the models predicted the total dry weight of food left in the stomach, but not the weight remaining for each individual meal. Interactions between meals led to an increase in evacuation rates for meals consumed early in the series and a decrease in evacuation rates for later meals.
5. Evacuation rates for piscivorous trout were compared with those for trout feeding on invertebrates in an earlier study, and were close to those for caddis larvae as prey, higher than those for mealworms and lower than those for a variety of invertebrate prey. Although a great deal is now known about the daily food intake and growth rates of trout feeding on invertebrates, there is little comparable information for piscivorous trout.     

A study of the food and feeding of perch, Perca fluviatilis L., in Windermere

SUMMARY. Perch were sampled for their stomach contents at regular intervals throughout 24 h from June until October in 1973 and from February 1975 until January 1976. They were found to feed on benthic organisms from November until April, on benthos and plankton during May and June and on perch fry and zooplankton from July until October. Perch over a wide size range feed on similar prey. Fish showed great variability in the weight of their stomach contents. A method based on a points system was developed to estimate the weight of food in the stomach for a given weight of fish at a known time. A diel feeding pattern which varied with the season was apparent from these data. Rates of gastric evacuation were assumed to be exponential and were calculated from the drop in night-time stomach content weights when food intake was assumed to be zero. The rates ranged from 0.18 mg h⁻¹ at a mean water temperature of 11°C in May to 0.35 mg h⁻¹ at a mean water temperature of 17°C in July. Assuming that food consumption followed a linear rate of intake, the standard Bajkov method was considered an adequate model to calculate daily food consumption. Daily food consumption (mg dry weight) was calculated for 150 g perch for all months of the year (November to April and September and October being combined). A second series of values was calculated making corrections for the time spent in the fishing gears when food intake was zero but gastric evacuation continued. Daily food consumption figures for 150g male perch based on Winberg's hypotheses (1956) and growth data showed no significant differences from this second series of values, when both were expressed in energy terms. When certain assumptions have been tested, growth data and Winberg's equations together may be a suitable method for calculating an energy budget for the Windermere perch population.     

Temperature and state‐dependence of feeding and gastric evacuation in juvenile Pacific halibut

Relationships between nutritional state, behavioural response to prey and gastric evacuation rates were examined in juvenile Pacific halibut Hippoglossus stenolepis feeding on squid. Pacific halibut reared at 2, 6 and 10° C were fasted for 1 or 7 days to generate variation in energetic state. The 7 day fast resulted in measurable declines in condition indices at 10 and 6° C but not at 2° C. At 10° C, all Pacific halibut consumed the first meal offered, but fish previously fasted for 7 days took significantly longer to locate and consume the meal than fish fasted for only 1 day. At 2° C, Pacific halibut fasted for 7 days did not generally consume the first meal offered, but resumed feeding 2·1 days sooner, on average, than fish fasted for only 1 day. The gastric evacuation rate of the squid meal was best described by a power model with near‐exponential curvature ( a  = 1·011). The evacuation rate was strongly temperature‐dependent ( Q ₁₀ = 3·65) but displayed the same degree of variability at each temperature. The evacuation rate in Pacific halibut was not affected by feeding history, body size or energetic state. Furthermore, individual variation in gastric evacuation rate was not correlated with feeding responsiveness at any temperature. These results indicate a general plasticity in the behavioural but not physiological aspects of energy acquisition.     

Diel feeding habits of juveniles of Mullus surmuletus (Linneo 1758) in the lagoon of the Stagnone di Marsala (Western Sicily, Italy)

Diet composition, feeding rhythm, gastric evacuation rate and daily ration were investigated in juvenile Mullus surmuletus (Linneo 1758). Fish were collected in the lagoon of the Stagnone di Marsala in western Sicily, in July 1995, during a 24 h sampling period. Copepoda, Polychaeta, Amphipoda and Tanaidacea were shown to be the most frequent prey items. The feeding index values showed two different daily feeding times. A unimodal trend in the daily rhythm of food consumption was derived, with a peak in feeding between 1200 and 2000 h. Gastric evacuation in juvenile M. surmuletus is best described by an exponential model, with a gastric evacuation rate R  = 0.66 g h⁻¹ ( r = 0.88) ( T  = 24.45 ± 0.64°C). The amount of food consumed daily, calculated according to the Elliott and Persson model, was 0.079 g, equal to about 8% of the average dry weight of the fish.     

Gastric evacuation rates and daily rations of Arctic cod (Boreogadus saida) at low temperatures

Gastric evacuation rates were determined for different sizes of Arctic cod (Boreogadus saida) at sub-zero temperatures (−1.4 and −0.5°C). These temperatures represent ambient conditions for Arctic cod in the Canadian high Arctic. Evacuation half-times, the time required for half of the content of the stomach to be evacuated, were longer (36–70 h; mean=51 h) than those reported in studies carried out on other fish species. Gastric evacuation rates at low temperatures were equal to, or below, those predicted by extrapolation from experiments conducted at higher temperatures. There were no significant differences in evacuation rates among fish size-groups or diets, but evacuation rates were slower for fish that had been starved prior to experiments. Estimated daily rations for Arctic cod in Resolute Bay, N.W.T., were 0.51% body weight for small fish (4.5 g) and 1.13% body weight for large fish (51 g). Slow stomach evacuation rates at low temperatures may limit daily food intake when food is seasonally abundant. This may contribute to slow growth rates and limited maximum size of Arctic cod in Canadian high Arctic waters. Received: 14 July 1997 / Accepted: 15 November 1997     

Effect of temperature and salinity on the gastric evacuation of juvenile sole Solea solea and Solea senegalensis

The juveniles of Senegal sole, Solea senegalensis, Kaup 1858, and common sole, Solea solea (Linnaeus 1758) concentrate in estuarine and coastal nurseries of widely differing temperatures and salinities. Yet, little is known about the effect of these physiologically important variables on the gastric evacuation rates of these species. Gastric evacuation experiments were performed on juveniles of S. senegalensis and S. solea. Three temperatures were tested, 26, 20 and 14°C at a salinity of 35‰. A low salinity experiment was also carried out at 15‰, at 26°C. Experimental conditions intended to reflect conditions in estuarine and coastal nurseries where juveniles of these species spend their first years of life. The relation between stomach contents and time was best described by exponential regression models for both species. An analysis of covariance (ancova ) was performed in order to test differences in evacuation rate due to temperature and salinity (slope of evacuation time against stomach contents) for each species. While increasing temperature increased evacuation rates in both species (although not at 26°C in S. solea), the effect of low salinity differed among species, leading to a decrease in gastric evacuation rate in that of S. senegalensis and an increase in S. solea. Differences in gastric evacuation rate between species were related to its metabolic optimums and to its distribution in the nursery area where fish were captured. Implications for the habitat use of estuarine and coastal nurseries are discussed.     

Gastric evacuation of three food types in the black and yellow rockfish Sebastes chrysomelas (Jordan and Gilbert)

The evacuation patterns of shrimp, crab and fish from the stomachs of black and yellow rockfish, Sebastes chrysomelas , were examined by feeding meals of known size and measuring the amount remaining after various post-prandial intervals. Linear, square-root, exponential, power exponential, logistic and Gompertz models (the latter two with unrestricted lower asymptotes, or with lower asymptotes restricted to 0% food remaining) were fitted to the wet weight, dry weight and volume of food remaining in the stomach as a function of post-prandial time. Evacuation patterns ranged from steeply concave (fish wet weight, dry weight, volume, shrimp dry weight), to linear (shrimp wet weight, crab dry weight), to highly convex with lag phases of up to 30 h (crab wet weight, crab and shrimp volume). Friability, the ease with which a food item is fragmented in the stomach, may be an important factor in determining evacuation patterns. The evacuation of a crab meal by tagged, free-ranging S. chrysomelas in the field was not significantly different from that of S. chrysomelas held in the laboratory.     

Factors affecting gastric evacuation in cod, Gadus morhua L., fed single-meals of natural prey

Functional relationships were derived in order to describe the effects of fish weight (262–2066 g), temperature (1–14° C), prey size and ration size (0.4–8.4% body weight) on gastric evacuation in cod fed four natural prey types. Interindividual (unexplained) differences among voluntary fed cod were relatively large. Power functions were appropriate for describing the effects of fish weight and ration size on the half-time of evacuation. Effects of meal and/or prey size appeared to counterbalance the effects of cod size. It is suggested that when cod of different sizes are fed fixed proportions of their body weight evacuation time is constant and independent of body size. There was found to be an exponential relationship between temperature and gastric evacuation time but, observations made at similar temperatures at different times of the year suggested that seasonal effects were of minor importance.     

Effects of temperature, prey type and prey size on gastric evacuation in small cod and whiting

Gastric lavage was used to investigate the effects of temperature, prey type and prey size on gastric evacuation in small cod Gadus morhua and whiting Merlangius merlangus . The fish were fed to satiation and subsequently the stomach contents were sampled to determine the rate at which food was evacuated. Satiation meal size was positively related to temperature and differed between prey types. The gastric evacuation rate (GER) also tended to increase with temperature and varied with prey type. GER at temperatures of 11.3–12.7° C averaged 1.5–1.7 times higher than at 6–9.5° C. There was no significant difference (P>0.05) in the evacuation of lugworm Arenicola marina , sandeel Ammodytes spp., and herring Clupea harengus , but the GER of brown shrimp Crangon vulgaris was much slower (P<0.05). No significant changes in GER were observed when fish were fed on three different size groups of either herring or brown shrimp. In whiting, there was no significant difference in the GER of individual herring or brown shrimp when they were fed as single species meals or incorporated in meals containing a mixture of prey species.     

Growth, daily ration, and gastric evacuation rates of milkfish (Chanos chanos) fed supplemental diet and natural food

Growth, daily ration, and gastric evacuation rates of milkfish ( Chanos chanos ) that fed on natural food and supplement diet were evaluated. Milkfish fingerlings (5.5g) were stocked at 1.5 fish/m² in ten 12 m² concrete tanks layered with 15-cm thick earthen bottoms. All tanks were regularly fertilized (16–20–0 and chicken manure) to maintain natural food production; 4 of the tanks additionally received a supplemental diet containing 34.3% protein and 4290 kcal/kg gross energy. Estimates or daily ration (based on dry weight of stomach contents) were calculated using the E lliot and P erson (1978) and E ggers 1977) models. Gastric evacuation rate was lower in fish that fed on natural food (1.57) compared to fish fed a supplemental diet (1.79). Consequently, the lower rate resulted in lower food intake and slower fish growth. When fish were provided a high quality supplemental diet, daily rations for fingerlings (35 g) to marketable size (116 g) ranged approximately from 0.60 to 19.68 kcal/fish/day. The deviation in daily ration (kcal/fish/day) from the above estimates may indicate the insufficient quantity of dietary energy taken by fish from natural food alone, which could be provided by supplemental diet.     

Influences of food particle size and dietary energy content on patterns of gastric evacuation in fish: test of a physiological model of gastric emptying

The evacuation of food from the stomach is probably not a continuous smooth process but may occur in a pulse-like (step-wise) fashion. It is hypothesised that the pattern of emptying is influenced by feedback signals from receptors located in the upper intestinal tract and by factors affecting the rate of physical/chemical breakdown of the food particles. Mathematical functions used for describing evacuation patterns imply that emptying is continuous and they are, therefore, at best, only approximations. It was predicted that different mathematical expressions would give best fit to empirical data from gastric evacuation studies dependent upon the experimental conditions employed. An analysis of published data supported the predictions and revealed that an exponential function best described the evacuation of small, easily digested prey items, but a linear expression gave the best fit to data of the emptying of large food items. It is suggested that differences in surface-to-volume ratios between large and small food particles and the friabilities of different food types are important in determining the pattern of emptying. Whilst dietary energy content also appears to be an important factor in the governing of gastric evacuation, it appears that emptying pattern in fish species is rather less sensitive to changes in dietary composition than is gastric emptying in mammals.     

The effects of food type, meal size and body weight on digestion and gastric evacuation in turbot, Scophthalmus maximus L.

Experiments were conducted on a size range of turbot using whole prey organisms. Stomach contents were serially sampled at intervals after feeding on a range of meal sizes of sandeels and sprats. Gastric evacuation was essentially linear with time, though some fluctuations were observed. Control over evacuation appeared to be by regulation of the processes leading to chyme production. Close agreement between evacuation rate and the satiation feeding rate of fish in laboratory experiments was observed. Implications of these findings for techniques of estimating the feeding rates of natural fish populations are discussed.     

A gastric evacuation model for three predatory gadoids and implications of using pooled field data of stomach contents to estimate food rations

The general applicability of the square root model to describe gastric evacuation in predatory gadoids independently of meal size was verified through experiments on whiting Merlangius merlangus , saithe Pollachius virens and cod Gadus morhua fed different fish and crustacean prey. Gastric evacuation rate was related by a negative power function to energy density of fish prey over an extended range from 3·4 to 11·5 kJ g^-1 of the effector variable. Gastric evacuation of crustacean prey seemed to depend on the characteristics of their exoskeleton. Gastric evacuation of mixed meals composed of fish prey with different energy densities could be described as a function of the overall energy density. The evacuation rate of each of the prey could be described directly by their share of the stomach content. A full gastric evacuation model including predator size, temperature and prey energy density was established for whiting, saithe and cod. It was demonstrated that estimates of food rations might be severely biased by use of mean values for prey composition and total mass of stomach contents.     

Evacuation of pelleted feed and the suitability of titanium(IV) oxide as a feed marker for gut kinetics in Nile tilapia

The present study assessed the suitability of titanium(IV) oxide, TiO₂, as a digesta passage marker in Nile tilapia Oreochromis niloticus and studied the shape of the evacuation curve in this species. In three separate trials, fish were given one dose of either 0·5, 0·25 or 0·1% of their body mass (% BME) of feed marked with 1% TiO₂ or 0·5% BME of the same feed without marker. The fish were serially slaughtered at intervals after feeding and the stomach contents analysed for dry mass and marker content. The data for individual trials were analysed with the linear, square root, surface area and exponential evacuation models and parameter comparisons showed that, although the marker interfered slightly with the evacuation process, true meal size could be predicted more accurately from the marker data. The results of an analysis of the combined data sets suggested that stomach evacuation in this species is dependent more on food particle surface area (surface area model) than on stomach content mass (exponential model) as is generally assumed. On the basis of these results, it was concluded that TiO₂ at an inclusion level of 1% is an acceptable marker for quantifying evacuation with a view to predicting food consumption but should be used with caution in digestibility studies.