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.     

Gastric evacuation in horse mackerel. I. The effects of meal size, temperature and predator weight

Gastric evacuation experiments were performed on horse mackerel Trachurus trachurus. A nearly full matrix experimental design with respect to the variables predator weight (<10–400 g) meal size (up to 7·8% body weight) and temperature (10–20°) was covered with 0-group smelt Osmerus eperlanus as prey. A general evacuation model without meal size as a variable was fitted to the data on wet weights as well as on dry weights by means of non-linear regression technique. Two methods of data transformation, relative data and square root transformation, were applied to improve variance homogeneity. The most reliable model fit was achieved on dry weight data applying the square root transformation technique: where S_t=stomach content (g wet weight) at time t after ingestion, S₀=the initial meal size, W =predator (g wet weight), and T =temperature. The estimated coefficient of the exponential temperature function, δ=00·032, corresponds to a Q ₁₀ value of 1·4 which is outstandingly low in comparison with results on other species. However additional experiments to determine maximum daily food rations indicated that appetite in contrast to gastric evacuation is strongly temperature dependent.     

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.     

Gastric evacuation rates and daily ration of piscivorous coho salmon, Oncorhynchus kisutch Walbaum

Effects of temperature and meal size on gastric evacuation rates of juvenile coho salmon, Oncorhynchus kisutch , consuming sockeye salmon, O. nerka , fry were examined and used in the estimation of daily meal, daily ration and number of fry consumed by coho in Chignik Lake, Alaska. Evacuation of fry by coho was best described by a negative exponential model (average R² = 0.93). A square root model also provided a good fit (average R² = 0·93), but the y-intercepts deviated more from the expected value than did the y-intercepts of the exponential model. The effect of temperature ( T , 5–13° C) and meal size (MS, 0·166–0·367 g) on the exponential evacuation rate (r_e, h^-1) could be described as
In the lake, coho fed continuously during the 24-h period in early June 1986 and 1987. Estimates of daily meal and ration of coho calculated by the Eggers method and the geometric mean of prey weight ranged from 0·224 to 0·435 g (2.1–4.4% body wt) depending on location and year. The Elliott & Persson method provided similar estimates of food consumption, whereas estimates based on the Pennington method and square root evacuation of prey differed from the exponential models. Sockeye fry represented 93% of the total prey weight. The average number of sockeye fry consumed per coho per 24 h, based on the arithmetic mean of prey weight, was 3·0–3·9 fry.     

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.     

Differential digestion and evacuation rates of prey in a warm-temperate grouper, gag Mycteroperca microlepis (Goode & Bean)

Prey-specific gastric evacuation rates and digestion state indices were modelled for gag Mycteroperca microlepis , a large warm-temperate grouper, consuming meals of either baitfish (scaled sardine Harengula jaguana ) or crab (purple swimmer crab Portunus gibbesii ). Power exponential models best fit the wet and dry mass gastric evacuation data and the average digestion indices over post-prandial time (PPT), regardless of prey type or gag size (Adjusted R² ≥ 0·79). Gag mass ( M ) or total length ( L _T) incorporated into an expanded power exponential model, along with exponential scalars, resulted in highly predictive ( R² ≥ 0·87) gastric evacuation and average digestion state models. The expanded power exponential models fit to the baitfish and crab wet mass gastric evacuation data differed significantly (Kimura's likelihood ratio test (LRT), both P < 0·001). Gag consuming crab showed a digestive lag period of at least 4 h (wet mass) and took a longer time to complete digestion relative to gag consuming baitfish. Gag, as well as many other warm-temperate and tropical groupers, consume a mixture of fish and crab prey and they will therefore require the development of a consumption model that incorporates mixed-prey gastric evacuation models.     

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.     

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.     

Mechanistic modelling of gastric evacuation in predatory gadoids applying the square root model to describe surface‐dependent evacuation

A new model approach to gastric evacuation in predatory fishes was shown to give accurate and reliable estimates of evacuation. It is believed that such a model would prove particularly useful in understanding predator‐prey interactions in natural systems. The model is a simple, geometric abstraction of the square root model predicting that evacuation time is proportional to the square root of meal size in accordance with extensive empirical evidence. Digestive processes are assumed to be restricted to an outer surface of total stomach content that is represented by the curved side of a cylinder. This way, total stomach content is considered a cylinder of constant length the mass of which is gradually reduced by successive peeling off its side leaving the ends unaffected. The fundamental rate δ describes mean thickness of the layer that is peeled off the cylinder per time unit. In the model, δ is constant independent of cylinder thickness but inversely proportional to the square root of cylinder length. Anatomical and dynamic characteristics of the stomach render this formulation of δ plausible. Using mean evacuation rate over time during evacuation of a meal, the model disregards prey heterogeneities that influence instantaneous evacuation rate. Relationships between prey heterogeneities and evacuation patterns were indicated by application of a general power model to evacuation data, and it was suggested how the effects of prey characteristics might be incorporated into the square root model to produce a generic model of gastric evacuation.     

黑鲷的胃排空率

在实验室内,１８℃条件下测量了体重在２０－４０ｇ的黑鲷的胃排空率,并用３种数学模型拟合了黑鲷的排空率,结果表明,３种模型都能很好地拟合实验数据,但通过统计学分析发现不论是用湿重还是用于重表示胃含物,线性模型都拟合得最好。用湿重表示时模型为Ｙ＝１．１４－０．０６ｔ（ｒ＾２＝０．８２,ｐ〈０．００１）;而用干重表示时模型为Ｙ＝１．３６－０．０７Ｔ（ｒ＾２＝０．６２,ｐ〈０．００１）,实验还得出体重２０     

黑鲷的胃排空率

在实验室内,18℃条件下测量了体重在 2 0～40g的黑鲷的胃排空率,并用 3种数学模型拟合了黑鲷的排空率.结果表明,3种模型都能很好地拟合实验数据,但通过统计学分析发现不论是用湿重还是用干重表示胃含物,线性模型都拟合得最好.用湿重表示时模型为Y=1.14-0.06t(r²=0 82,p<0.001);而用干重表示时模型为Y=1.36-0.07t(r²=0.62,p<0.001).实验还得出体重20～40g的黑鲷在 18℃的胃排空率为每小时排出摄入食物量的 6%～7%.     

A comparison of gut evacuation models for larval mackerel (Scomber scombrus) using serial photography

A novel technique is described, using serial photography of the gut contents of transparent living larval fishes, to generate individual gut evacuation time series. This technique was applied to Atlantic mackerel Scomber scombrus larvae to compare three widely used models of gut evacuation: linear, exponential and square-root. Regression r ² for the exponential model exceeded those for the linear and square root models in 20 of 21 time series, strongly supporting the exponential model. At the initial gut fullness for each time series, total gut evacuation rates calculated with the exponential model averaged 2·2 and 1·3 times greater than those calculated with the linear and square-root models, respectively, and would produce correspondingly higher estimates of feeding rates for field-collected larvae with similar levels of gut fullness. The results highlight the importance of choosing the appropriate evacuation model in feeding studies, particularly those intended to examine short-term changes in larval fish feeding rates, a contributing factor to the highly variable yearly recruitment of many marine fish species.     

有氧运动训练对中华倒刺鲃幼鱼肠道排空及其数学模型选择的影响

为了考察有氧运动训练对中华倒刺鲃幼鱼肠道排空及其数学模型选择产生的影响, 在(250.5)℃条件下, 将108 尾大小相当的实验鱼(24.470.03) g, (10.900.02) cm随机分为对照组(CG), 1 BL/s 训练组(1TG)和2 BL/s 训练组(2TG), 在相应流速下运动训练8w 后, 轻度麻醉灌喂体重1.45%的饵料, 随后在不同时间点分别测定肠道内容物重量及其百分比, 用3 个常见数学模型对实验数据进行拟合并比较。结果发现, 1TG 和2TG 训练组的体重特定生长率(WSGR)显著高于CG 对照组 (P0.05);各实验组肠道内容物干重及其百分比随着摄食后时间的延长而显著降低;所有实验组肠道排空的最优数学模型均为平方根模型;训练组(1TG 和2TG)的肠道排空率(皆为0.49%/h)明显大于CG 对照组(0.41%/h)(P0.05), 1TG 和2TG 训练组的肠道排空时间(分别为 20.39h 和 20.33h)短于 CG 对照组(24.51h)。研究表明, 有氧运动训练没有对中华倒刺鲃幼鱼的肠道排空特征及其最优数学模型选择产生显著影响, 但明显提高了该种鱼的肠道排空率并缩短其肠道排空时间。       

Gastric evacuation rate of burbot fed single-fish meals at different temperatures

The gastric evacuation rates of burbot Lota lota , fed a single meal of vendace, Coregonus albula , were measured in the laboratory at five temperatures (1·3, 2·6, 4·8, 9·4 and 12·6° C). Gastric evacuation rate increased exponentially with increasing temperatrure, but the results suggest that gastric evacuation rates of burbot at low temperatures are lower than those of other freshwater fish species. Temperature and the ratio of meal weight to burbot weight were the most important factors affecting gastric evacuation rate. There was no significant difference in gastric evacuation rate between three different prey species: vendace, perch Perca fluviatilis , and smelt Osmerus eperlanus .     

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.     

Does the cylinder model of gastric evacuation predict observed evacuation of mixed meals of prey of contrasting geometries in a piscivorous fish?

The simple surface abstraction of the cylinder model (each prey as well as the total stomach contents is considered a cylinder that is gradually reduced by successive peeling off its curved side) was challenged by data on evacuation of a meal composed of three sandeels Ammodytes tobianus and a dab Limanda limanda fed to Atlantic cod Gadus morhua. While the body shape of A. tobianus comes close to that of a cylinder, the flatfish L. limanda takes a discoid form. As opposed to a modified form of the cylinder model, where the contrasting geometries of the fish prey were implemented, the simple, original cylinder model held the potential to predict evacuation of the individual prey types as well as the total stomach contents. Thus, the present study adds significantly to the increasing evidence that points to the generic nature of the model and its implicit square root function. Also, the present study corroborated a basic assumption that the variability of evacuation data not accounted for by the cylinder model primarily can be ascribed to the intraspecific variation in gastric performance of the predator.     

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.     

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.