Gastric digestion and evacuation in whiting, Merlangius merlangus (L.)

Three meal sizes of sandeels were fed to whiting in order to monitor the evacuation of food out of the stomach. The stomach contents were sampled at intervals after feeding, using a stomach pump. In such experiments, the proportion of fish with empty stomachs tends to increase with time and, since stomach contents are limited to zero or positive values, the variance of the stomach contents becomes censored at zero. This tends to produce a curved relationship between mean stomach content and time, which gives the impression that evacuation rate slows down at low levels of stomach fullness. By taking account of censoring, it was shown that the evacuation curve generated for whiting was consistent with and could be generated from a linear model in which the rate of gastric evacuation exhibited by the fish was constant and independent of meal size, level of stomach fullness and time after feeding. The parameters of the linear model were estimated by maximum likelihood and then applied in a second model to predict the observed mean stomach content. The average gastric evacuation rate of whiting of mean weight 268 g at 10°C was 0.31 g h⁻¹.     

Gastric evacuation in the sandbar shark, Carcharhinus plumbeus

Sandbar sharks maintained in an enclosure in the natural environment were fed meals of soft blue crab, Callinectes sapidus , or menhaden, Brevoortia lyrannus , and were sacrificed at intervals after feeding to measure rates of gastric emptying. Regression analysis was used to evaluate the adequacy of various models in describing the decrease in stomach contents with time after feeding. A Gompertz growth curve provided the best fit to the data for both food types. This model suggests that gastric emptying is characterized by an initial lag phase during which evacuation rate increases, a period of maximal evacuation, and a decreasing evacuation rate during the later stages of digestion. Passage of food from the stomach was slow relative to other species of fish studied. The time required to evacuate 98% of a meal was 92–3 and 70–7 h for sharks fed menhaden and blue crab respectively. Evacuation rates did not differ between day and night periods.     

Recent developments for making gastric evacuation and daily ration determinations

Synopsis Gastric evacuation rate estimates often suffer from an important bias caused by fitting experimentally-derived data distributions that are inherently constricted by the X-axis (Y = 0). Monte Carlo simulations were used to evaluate the bias. Truncating constricted distributions prior to curve fitting was suggested as a means to circumvent the problem. A food consumption model developed by D.S. Robson was presented. It employs the integral of the function fit to percentage gastric evacuation data, and does not require an a priori assumption of exponential gastric evacuation. The methods were illustrated using experimental gastric evacuation data and stomach contents data for fishery-caught yellowfin tuna, Thunnus albacares.     

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.     

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.     

The Role of gastric evacuation rate in handling time of equal-mass rations of different prey sizes in northern pike

In a laboratory experiment, northern pike Esox lucius gastric evacuation rates did not differ between equal-mass rations of small and large prey. In a comparison between intermediate and large prey, the pike were unable to fit two intermediate prey completely into the stomach at the same time, resulting in two consecutive evacuations, and changes in patterns of gastric evacuation. Thus, total gastric evacuation time was not affected by prey size composition in equal-mass rations, but patterns in evacuation rate may depend on the size ratio between predator and prey. Cumulative manipulation time between strike and complete swallowing of prey differed between equal-mass rations of small, intermediate and large prey, in that small prey took the shortest time to manipulate. Pike had problems striking and redirecting intermediate prey to swallow them head first, and the manipulation times for intermediate prey were as long as for large prey. Since an increased time manipulating prey in the mouth increases risk of predation and intraspecific interactions in pike, it is concluded that risks associated with long manipulation times, and not only energy per total handling time, determine prey value and prey size preference in this piscivore.     

Effect of experimental conditions on the stomach evacuation of Coregonus lavaretus L.

The gastric evacuation of juveniles of Coregonus lavaretus L. fed on living Daphnia pulicaria was investigated. Three successive stages of stomach evacuation were observed when one meal per day was given: (i) a lag phase between the end of food intake and the beginning of stomach evacuation, (ii) a linear reduction of stomach content, (iii) a long residence time for food relics in the stomach. The initial stomach content and the stomach evacuation time are correlated positively. The stomach content increased during feeding when three meals day ⁻¹ were provided and it decreased when no food was available. During the course of an experiment the highest stomach content found increased with increasing daly ration. Excess feeding resulted in a low stomach content similar to that found with rations about 30–50% of the maximum daily food intake. Therefore the daily food intake cannot be determined by the single parameter of stomach content alone. Identical initial stomach contents showed significantly higher stomach evacuation rates under three meals day⁻¹ conditions than under one meal day⁻¹ conditions.     

The contribution of gastric emptying scintigraphy to the treatment of obesity with gastric bandage--preliminary results

Introduction: Gastric banding for morbid obesity is among the least mutilating of procedures used in bariatric surgery and is classified as a restrictive surgical method. Although it is widespread, so far, however the mechanism responsible has not been fully explained. Methods and results: The authors present the preliminary results from scintigraphic examination of the evacuation ability of the stomach using food labeled with (99m)Tc-colloid in six obese patients with a gastric bandage. This initial study showed that the functionality of the bandage demonstrated as a significant drop in body weight, is connected with slower evacuation of the stomach. However, the use of adjustable bandages would have significantly slowed and restricted the passage of food through the cardia of the stomach. Conclusions: Since it can be extremely difficult to adequately objectively determine the functionality of gastric bandages, evaluating the gastric emptying scintigraphy may be useful in fulfilling this purpose.     

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.     

A surface-dependent gastric evacuation model for fish

A gastric evacuation curve expresses how fast prey disappear from the stomach, and empirical models are used generally for the relationship between weight of prey remaining ( W_t ) and time ( t ) after a meal. Unfortunately, empirical models are likely to have restricted applicability because their parameters often represent limited biological mechanisms. This paper develops a simple digestion model. The simplest form of the model has four parameters; the digestion velocity (expressing enzymatic breakdown of prey), prey length, initial prey radius and the density of the prey. Two more parameters are included in an extended version; a time-delay before digestion starts and environmental temperature. The approach is based on the assumption that prey digestion is a surface process in that digestive enzymes attack progressively deeper into a prey of known size and shape so that the average digestion rate is proportional to the prey radius r (m). This process is characterized by the digestion velocity d_s (m s ^–1). Unknown parameters are estimated with uncertainty using the maximum likelihood technique. Model evaluation using published data sets demonstrated that the new model is flexible. Prey geometry is incorporated into the model and temperature effects upon gastric evacuation are linked directly to the digestion velocity.     

Prey exoskeletons influence the course of gastric evacuation in Atlantic cod Gadus morhua

This study examined the effects of prey exoskeleton characteristics on gastric evacuation patterns in Atlantic cod Gadus morhua. Three distinct stages were highlighted in the gastric evacuation of crustacean prey characterized by a robust exoskeleton. The experiments confirmed that the three shrimp species, Pandalus borealis, Pandalus montagui and Eualus macilentus, and the crab Chionoecetes opilio, were evacuated from the stomach at different rates. The duration of all stages increased with increasing ash (and carbonate) content of the fresh prey. Thickness, chemical composition and morphology of the prey exoskeleton all affected gastric evacuation: duration of initial delay, overall evacuation rate and a decreased evacuation rate at the end of the process. The power exponential function (PEF), with its shape parameter, described the course of evacuation for these prey types well, especially the initial delay. The PEF does not, however, allow describing evacuation by the current stomach content mass independent of meal size, which limits its usefulness in estimating consumption rates of wild G. morhua. To predict and describe gastric evacuation of prey with a robust exoskeleton, it is therefore suggested that the square‐root function be expanded with an initial lag phase, coupled to the mechanistically based cylinder model of gastric evacuation.     

Gastric evacuation of mixed stomach contents in predatory gadoids: an expanded application of the square root model to estimate food rations

A mechanistic gastric evacuation model of mixed stomach contents was established. Its simple generic nature should prove useful for studying predator‐prey interactions in fish communities. Only prey lengths were required as additional input. Surface dependent digestive processes were considered to act on equal fractions of individual prey‐cylinder surfaces. Primary and interactive effects of size, energy density and resistance to digestion of individual prey in a stomach were described by the model. Model predictions of results from experiments on gastric evacuation of meals composed of different prey types demonstrated the capability of this model, as opposed to previously applied model principles, to predict evacuation of mixed stomach contents involving the three above‐mentioned prey characteristics. The study furthermore illustrated that estimates of food ration might be severely biased by using improperly formulated effects of prey characters on gastric evacuation, and demonstrated that the new model holds the potential to predict food ration and diet composition for wild populations of predatory fishes.     

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.     

Effect of sampling interval and temperature on the accuracy of food consumption estimates from stomach contents

The effect of temperature and sampling interval on the accuracy of food consumption estimates based on stomach contents was studied using simulation. Three temporal patterns of feeding were considered (scattered throughout the day, one 5 h period or two 5 h periods) and gastric evacuation was modelled according to published values. Sampling intervals of 3 h gave reasonable food consumption estimates (2 to 19% error) at all temperatures. Comparably, sampling intervals as large as 12 h gave reasonable estimates of food consumption (1 to 20% error) when temperature was set to ≤10° C. At temperatures <5° C, even 24 h intervals (equivalent to one daily sampling) provided reasonable estimates of daily food consumption (2 to 19% error) for all but the highest gastric evacuation rate combined with one daily feeding period (47% error). The temperature effect on estimation error resulted from diminishing temporal fluctuations in stomach contents with slower gastric evacuation rates. It follows that sampling effort may be considerably minimized when estimating food consumption from stomach contents during periods with low temperatures such as the winter time experienced by temperate fishes.     

Are the dynamics of gastric evacuation of natural prey in a piscivorous flatfish different from what is going on in a gadoid?

Despite the fact that the stomach of turbot Psetta maxima is a curved tube that forms a half circle, it was demonstrated that gastric evacuation in this predatory flatfish fed natural prey closely followed the surface‐dependent cylinder model developed from studies on gadoids with a straight stomach. Evacuation experiments were performed on two size groups of P. maxima fed sandeel Ammodytes tobianus as well as on P. maxima fed brown shrimp Crangon crangon at three different temperatures. This enabled the provision of a gastric evacuation model for studies on P. maxima, which takes into account the effects of the explanatory variables predator size, temperature, prey energy density and resistance of prey to the digestive processes in the stomach. Basically, the cylinder model predicts that a square‐root function accurately describes gastric evacuation, which is inconsistent with the conclusion of a previous study on P. maxima that evacuation of A. tobianus is essentially linear with time. Use of the cylinder model to the values of the explanatory variables presented in the latter study, however, gave accurate predictions of the actually acquired evacuation data, which points to the generic value of the model.     

Food limitation in a nursery area: estimates of daily ration in juvenile scalloped hammerheads, Sphyrna lewini (Griffith and Smith, 1834) in Kāne'ohe Bay, ō'ahu, Hawai'i

Gastric evacuation and daily ration were studied in juvenile scalloped hammerhead sharks in Kāne'ohe Bay in order to better understand their ecology and role as apex predators in the marine environment. Three major variables known to affect the rate of gastric evacuation were manipulated: meal size, prey species, and temperature. Rates of gastric evacuation were faster than have previously been measured for sharks. The time for 80% of the meal weight (dry) to be evacuated ranged from 5.4 to 22.1 h. Daily ration was estimated using two different methods based on gastric evacuation rates and stomach content data. Estimates ranged from 2.12% to 3.54% of the body weight, which is relatively high compared to other elasmobranchs, but lower than estimates of maintenance ration for juvenile scalloped hammerheads. Data available for diet and growth of juvenile scalloped hammerhead sharks in Kāne'ohe Bay as well as data for their prey species suggest that these sharks may be surviving much of the time at consumption levels below maintenance ration.     

How are prey fishes of multiple meals evacuated from the stomach of a piscivorous fish?

The observations that the gastric evacuation of a meal was accelerated after ingestion of an additional meal, whereas the evacuation of the latter meal was slowed down were compared to the predictions of mass-dependent models derived from experiments on evacuation of single meals. In contrast, the present study on cod Gadus morhua fed identical double meals of sandeel Ammodytes tobianus demonstrated that the evacuation of each meal was slowed down relative to the evacuation of a single meal. This fact could be explained by the exposed meal surface area being reduced while the meals were together in the stomach as it was expected by a surface-dependent cylinder interpretation of the square-root model. The cylinder model forecasted accurately the observed evacuation of each meal in the double-meal situation. The simple surface considerations of the model further explained the discrepancies between observed and expected evacuation rates using mass-dependent models, which suggested that the above finding of the previous studies was an artefact. The ability of the cylinder model to predict properly also the evacuation of multiple meals in juvenile coho salmon Oncorhynchus kisutch indicated that the model would apply to gastric evacuation in small-sized predatory fishes as well. This should increase considerably its value as a generic tool for studies on the feeding biology of fishes in the wild.     

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 in the young lemon shark,Negaprion brevirostris,under field conditions

Synopsis Gastric evacuation in young lemon sharks, Negaprion brevirostris, was studied in a field enclosure. Regression analysis was used to evaluate the adequacy of linear, exponential, and square root models in describing the decrease in stomach contents with time after feeding. The linear model produced the best fit and was thus used to compare gastric evacuation at the three temperatures. Gastric evacuation in young lemon sharks is considerably longer than for carnivorous teleosts but shorter than for other elasmobranchs. These differences are the result of differing energy requirements determined by the physiology and behavior of the species.