Development and validation of a bioenergetics model for juvenile and adult burbot

Oxygen consumption of juvenile and adult burbot Lota lota was measured in an intermittent-flow respirometer to determine the effect of temperature and fish body mass on metabolic rate. These results were combined with data from earlier experiments and the 'Wisconsin bioenergetics' model was constructed. The model was validated under laboratory conditions by comparing observed and predicted food consumption and growth of burbot fed on dead vendace Coregonus albula . There was a good correspondence between observed and estimated growth and food consumption under experimental conditions: the mean absolute per cent errors of growth and food consumption were 4·8 and 24·0%. Estimated values with the new model were an improvement over the Atlantic cod Gadus morhua model previously used for burbot. In the field, the reliability of food consumption estimates was verified by using polychlorinated biphenyls (PCB) accumulation as an indirect indicator of the food consumption rate. The total PCB concentration of nine out of 13 burbot was estimated accurately. Thus, the burbot model produced good estimates of food consumption, even under field conditions.     

Prey consumption rates and growth of piscivorous brown trout in a subarctic watercourse

Prey consumption rates of piscivorous brown trout Salmo trutta were studied in the Pasvik watercourse, which forms the border between Norway and Russia. Estimates of food consumption in the field were similar to or slightly less than maximum values from a bioenergetic model. The piscivore diet consisted mainly of vendace Coregonus albula with a smaller number of whitefish Coregonus lavaretus . Individual brown trout had an estimated mean daily intake of c . 1·5 vendace and 0·4 whitefish, and a rapid annual growth increment of 7–8 cm year⁻¹. The total population of brown trout >25 cm total length was estimated as 8445 individuals (0·6 individuals ha⁻¹), giving a mean ±  s . e . annual consumption of 1553880 ± 405360 vendace and 439140 ± 287130 whitefish for the whole watercourse. The rapid growth in summer of brown trout >25 cm indicated a high prey consumption rate.     

Morphological differentiation and resource polymorphism in three sympatric whitefish Coregonus lavaretus(L.) forms in a subarctic lake

Three field‐identified whitefish Coregonus lavaretus forms in Lake Muddusjärvi, Finland, were compared in morphology, diet and prey size. First, these forms were studied with univariate and multivariate analysis to assess morphological divergence at a higher resolution level than in the field. Second, stomach contents were analysed to estimate diet‐overlap among forms. Finally, the relationship between prey size and morphology was examined. The whitefish were assigned to the initial field‐classification with 99·2% and 98·8% accuracy for morphologic and meristic traits, respectively. The small sparsely‐rakered form (SSR) had the shortest rakers and largest gillraker space, followed by the large sparsely‐rakered form (LSR) with intermediate gillraker length and gillraker space, while densely‐rakered whitefish (DR) had the longest rakers and smallest gillraker space. The two sparsely‐rakered whitefish forms (LSR and SSR), consumed mainly benthic macroinvertebrates, while densely‐rakered whitefish (DR), utilized pelagic food items. Average diet‐overlaps between whitefish forms were low in June‐September (Schoener's α = 0·02 − 0·23). Gillraker number and length were negatively correlated to prey length in the diet ( r  = −0·73, and r  = −0·60), while gillraker space was positively correlated with prey length ( r  = 0·81). The fact that these whitefish forms were morphologically and ecologically segregated, and that gillraker traits probably have a functional value in food selection, further suggests that natural selection has been important in structuring life‐history trajectories into divergent niche use.     

Abundance and distribution of Diphyllobothrium ditremum Creplin (Cestoda: Pseudophyllidea) plerocercoids in benthic whitefish, in northern Finnish Lapland

The prevalence of Diphyllobothrium ditremum plerocercoids in whitefish Coregonus lavaretus ranged between 70 and 100% in Lake Kilpisjärvi and in three other lakes in Northern Lapland, Finland. The mean abundance in Lake Kilpisjärvi (age groups 1+-10+ years), ranged between 103·5± 71·3 in 1992–1993 to 110·9± 80·0 plerocercoids per fish in 1997. The asymptotic value of the infection levelled at 113 plerocercoids per host after age 3. No significant difference in abundance was detected between study years ( P >0·10). Abundances in other lakes ranged between 4·8±9·7 and 91·1±115·1. Two seasonal peaks of plerocercoid recruitment were observed in Lake Kilpisjärvi; between March and April ( P <0·002) and between September and October ( P =0·042). In autumn the numbers of larvae increased particularly in female fish. The invasion rate of the parasite was lower in other lakes studied, and the infection rate in whitefish was closely related to the copepod food eaten.     

A validated bioenergetics model for ruffe Gymnocephalus cernuus and its application to a northern lake

A bioenergetics model was constructed for ruffe Gymnocephalus cernuus , and the model was used to estimate the food consumption of a native G. cernuus stock in Pyhäjärvi, a large mesotrophic lake in south-west Finland. The parameters for the bioenergetics model were determined through laboratory experiments conducted at several temperatures between 5 and 22° C for different sized G. cernuus (from 6 to 27 g). The model was verified and validated with independent laboratory growth experiments. The validated model was applied to field data from Pyhäjärvi to estimate the food consumption of G. cernuus and its effects on the zoobenthos community during the years 2002–2004. The total annual food consumption of the G. cernuus stock ranged from 11·6 to 15·7 g m⁻² (wet mass). Chironomidae was the most abundant prey, 81% of the total consumption. The annual total production of zoobenthos in Pyhäjärvi was 15·2 g m⁻² (wet mass). Therefore, it can be concluded that G. cernuus stock is likely to exert a significant influence on the zoobenthos community in this lake, indicating a potential role of G. cernuus as an influential invasive species in many freshwater ecosystems in Europe and North America.     

The effects of food consumption rate, body size and temperature on net food conversion efficiency in saithe and whiting

Net food conversion efficiency κ was estimated from growth experiments on saithe Pollachius virens and whiting Merlangius merlangus that were fed natural prey at a range of ration levels including satiation rations. The conversion efficiency, which specifies the net energy fraction of ingested energy C , was described appropriately by a simple power function of food consumption rate as κ  = 0·272 C ^0·18 and κ  = 0·426 C ^0·11 for saithe and whiting. This functional relationship was supported by the patterns of accretion of lipids and proteins in saithe. No significant effects of temperature and body size on κ was, however, demonstrated in this study except for the indirect influence using feeding levels (rations expressed relatively to satiation rations) in bioenergetics models.     

Does specific dynamic action (SDA) regulate return of appetite in the lesser spotted dogfish,Scyliorhinus caniculcla?

Specific dynamic action (SDA) and the return of appetite (RA) were measured in juvenile (weight range 10·9–16·3 g) and adult (weight range 588·8–802·2 g) Scyliorhinus canicula by means of respirometry and re-feeding trials respectively. Juvenile S. canicula consumed relatively more food than the adults at all times indicating faster rates of food processing in juvenile dogfish. At no time during the SDA effect was there any evidence that elevated levels of oxygen consumption completely prevented further food consumption in either adult or juvenile dogfish. The coefficient (Duration_SDA/Duration_RA) was similar for both adult (0·52) and juvenile (0·49) S. canicula suggesting that SDA metabolism was closely associated with the processes involved in appetite return.     

Temporal repeatability of relative standard metabolic rate in juvenile Atlantic salmon and its relation to life history variation

There was a strong correlation between the relative standard metabolic rate (rSMR) values of individual Atlantic salmon Salmo salar L. measured 5 and 22 weeks after first feeding in June and October respectively (Pearson's r =0·68, 26 d.f., P <0·001). However, this is a conservative estimate of repeatability as two separate regressions were used to calculate SMR in October due to the separation of the population into an upper modal group made up of early migrants and a lower modal group comprised of delayed migrants. SMR values in June and October were similar (paired t test, t =-0·85, 27 d.f., P >0·05) when expressed as the percentage deviation from those predicted for a fish of that size based on the body mass/SMR relations in June or October indicating that relative standard metabolic rates were stable over time when food was not limiting. rSMR status was maintained in 19 of the 28 fish (i.e. 68%) between the two measures. rSMR status was correlated with life history strategy: salmon fry with a high SMR in June were more likely to become smolts during the autumn than those with a low SMR.     

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.     

Comparison of field-based and indirect estimates of daily food consumption in larval perch and zander

Since bioenergetics models for 0+ fish have seldom been validated by field consumption estimates, field-based and indirectly estimated daily food rations were compared in larval perch Perca fluviatilis and zander Stizostedion lucioperca. Field-based estimates were calculated with linear and exponential evacuation rates based on gut fullness data during a 24-h cycle, with hourly field samplings instead of the normally recommended 3-h intervals. Indirect calculations used bioenergetics modelling with variable activity multipliers ( A ). Field-based estimates of daily rations ranged between 0·21 and 0·27 g g⁻¹ day⁻¹ in perch (mean L _T 13·1 mm) and 0·31–0·40 g g⁻¹ day⁻¹ in zander (mean L _T 10·6 mm). The higher values were calculated by using the exponential model. Daily rations calculated by bioenergetics modelling with A = 1 were only slightly higher than direct estimates in both species. However, if A values >1 were used, calculated daily rations were substantially higher than direct estimates. Estimates of daily ration based only on every third value ranged between 41 and 72% compared with 1-h intervals, mainly because of lower estimates of evacuation rate.     

Effects of temperature, body size and feeding on rates of metabolism in young‐of‐the‐year haddock

The mean rate of oxygen consumption (routine respiration rate, R _R, mg O₂ fish⁻¹ h⁻¹), measured for individual or small groups of haddock Melanogrammus aeglefinus (3–12 cm standard length, L _S) maintained for 5 days within flow‐through respiratory chambers at four different temperatures, increased with increasing dry mass ( M _D). The relationship between R _R and M _D was allometric ( R _R = α  M ^b ) with b values of 0·631, 0·606, 0·655 and 0·650 at 5·0, 8·0, 12·0 and 15·0° C, respectively. The effect of temperature ( T ) and M _D on mean R _R was described by     indicating a Q ₁₀ of 2·27 between 5 and 15° C. Juvenile haddock routine metabolic scope, calculated as the ratio of the mean of highest and lowest deciles of R _R measured in each chamber, significantly decreased with temperature such that the routine scope at 15° C was half that at 5° C. The cost of feeding ( R _SDA) was c . 3% of consumed food energy, a value half that found for larger gadoid juveniles and adults.     

Metechinorhynchus salmonis infection and diet in the river-spawning whitefish of the Bothnian Bay

A total of 206 river-spawning whitefish Coregonus lavaretus (L.) from the Bothnian Bay was investigated for food objects and acanthocephalans in the years 1975–78. The only acanthocephalan species found was Metechinorhynchus salmonis (Müller 1780), the total number of which was 1295.
The whitefish is shown to be highly susceptible to infection by M. salmonis , with as many as 88·9% of specimens infected in a sample in June 1978, although the mean intensity was only 8·9 parasites per host. The river-spawning whitefish does not favour the intermediate host, Pontoporeia affinis , in its summer diet in the Bothnian Bay preferring to feed on eggs and molluscs, so that the incidence of M. salmonis infection decreases to 33·3% by the middle of summer. This is further reduced in the late summer by the fast associated with the migration to their upriver spawning grounds. The fact that the other acanthocephalan inhabiting the river-spawning whitefish in the southern Gulf of Bothnia, Echinorhynchus gadi Müller, 1776 is not to be found in the Bothnian Bay population suggests that the fish captured in spring have remained in the Bothnian Bay throughout the winter.     

Impact of temperature on food intake and growth in juvenile burbot

The effect of temperature on food consumption, food conversion and somatic growth was investigated with juvenile burbot Lota lota (age 0 years). Juvenile burbot showed a significant dome shaped relationship between relative daily food consumption ( C _R) and temperature ( T ) with C _R = − 0·00044 T ² + 0·01583 T  − 0·06010; ( n  = 90, r ² = 0·61). Maximum C _R was at 17·9° C (95% CL 17·2–18·6° C). The temperature related instantaneous growth rate ( G ) also followed a dome shaped function with G  = − 0·000063 T ² + 0·002010 T  − 0·007462; ( n  = 95, r ² = 0·57), with maximum growth rate at 16·0° C (95% CL 15·3–16·6° C). A significant linear relationship was found between the water temperature and the conversion coefficient ( C _C) with C _C = − 1·63 T  + 59·04; ( n  = 80, r ² = 0·74). The results indicate that juvenile burbot in large lakes benefit from higher water temperatures in the littoral zone, by increased food uptake and growth, especially during the warm summer months. Because profundal water temperatures do not reflect the optimal temperature for food consumption in large burbot, temperature is unlikely to be the main proximate factor for the obligate littoral‐profundal migration of juvenile burbot observed in many lake populations.     

Growth and food consumption of 0+ Arctic charr fed pelleted or natural food at six different temperatures

The effects of temperature and diet on the specific growth rate and food consumption of 1-summer-old Arctic charr Salvelinus alpinus were studied. Fish were reared singly in aquaria at six different constant temperatures (5, 9, 13, 16, 18 and 20°C). They were fed Neomysis integer or commercial pelleted food for 2 weeks and growth and food consumption were measured. In both experiments, growth rate increased to an optimum at 15°C. Growth rates were high in the range 13–18°C, with no significant ( P >0·05) differences between temperatures. No significant ( P> 0·05) differences in growth were found between fish at 9 and 20°C. There were no effects of diet on size-adjusted growth rates. The growth efficiency decreased with increasing temperature in both treatments, but the decrease was faster in the Neomysis treatment. Charr seemed to compensate for the high water content (79·5%) of Neomysis by having a higher food intake.     

Influence of different rations and water temperatures on the growth rates of shortfinned eels and longfinned eels

Juvenile (12–152 g) shortfinned eels Anguilla australis and longfinned eels A. dieffenbachia caught in New Zealand streams were fed squid mantle Nototodarus spp. 4 days per week in laboratory experiments. A linear multiple regression equation showed the amount of food eaten (0–2·7% w day⁻¹) explained 77·7% of the variation in specific growth rates (–0·60 to +1·07% w day⁻¹) among individual eels, while previous growth rates, water temperature (10·0–20·6°C), and eel weight (12–152 g) explained a further 5·6, 1·4 and 0·8%, respectively. Growth in length ranged from –0·3 to +0·9 mm day⁻¹. Eels which were starved and then given high rations grew substantially faster than expected. Once growth rates were adjusted for differences in ration and other factors, there were no significant differences in growth rates between species or individual fish. Growth of shortfinned eels fed maximum rations of commercial eel food depended on fish size and water temperatures and ceased below 9·0°C. Growth rates in the wild were substantially less than the maximum possible, after seasonal changes in water temperatures were taken into account, indicating that food supplies and not low water temperatures were controlling growth rates in the wild.     

Cry (preservation of spermatozoa of the whitefish Coregonus muksun Pallas

The cryopreservation methods for whitefish, Coregonus muksun , spermatozoa were studied using six different extenders with -40°C and -196°C as storing temperatures. In 1980, the spermatozoa of individual fishes were frozen in dry ice, in vials containing three different media based on glycerol as cryoprotectant. The pure glycerol media resulted in a 20% average rate of fertilization when samples were stored at — 40°C. The thawing procedure did not affect fertilization. In 1981, the sperm of individual fishes was frozen in pellets and stored in liquid nitrogen. Cryopreserved semen thawed in 0·119M NaHCO₃ produced 29·6–87·7% eyed eggs. The differences were caused mainly by the type of extender. The highest rate of fertility was obtained using an extender consisting of 0·3 M glucose with 20% glycerol (87·7% eyed eggs; 98·6% eyed eggs when compared to the corresponding control). The same values, using the modified extender of Stoss (Stoss & Refstie, in press), were 38·7–82·4% and, on the average, 90% eyed eggs, respectively.     

Development and application of a bioenergetics model for juvenile walleye Pollock

A bioenergetics model was parameterized for age-0 walleye pollock, Theragra chalcogramma , based on a synthesis of literature data. The sensitivity of the new parameters was tested by individual parameter perturbation (IPP) analysis. The model was applied to estimate individual and total cohort food consumption of age-0 pollock in two areas of high pollock density in the Gulf of Alaska during the summer of 1990. Total cohort consumption was compared with zooplankton biomass and production estimates for the same areas and times of the year. The model was also used to examine the bioenergetic implications of age-0 pollock diel vertical migration through a thermal gradient. During a 1-month bioenergetics simulation, individual daily consumption decreased from 16·0 to 6·0% of wet body weight. Daily ration estimates corresponded well with independent field estimates of daily ration for the same areas and time of the year. Comparison of total cohort consumption with prey availability (production and biomass) indicated minimal potential for food limitation. Bioenergetic optimization of growth can be a potential benefit of diel vertical migration to age-0 pollock, however more information on prey density and distribution is needed to test this hypothesis thoroughly.     

Application of a bioenergetics model for analysis of growth and food consumption of subarctic whitefish Coregonus lavaretus (L.) in Lake Kilpisjärvi, Finnish Lapland

A bioenergetics model was used to estimate daily food consumption and growth of benthic whitefish Coregonus lavaretus (L.) of age groups 1+ to 10+, in oligotrophic Lake Kilpisjärvi, northern Finnish Lapland. Fish and zooplankton samples and water temperature data were collected twice per month from February to December 1993. Simulation results indicated wide seasonal variations in consumption and growth rates. Growth was in general slow; especially in age-group 6 the net increase in weight was slight. The fit of the model to the weight data was good, and the model was able to follow even rapid seasonal variations in the weight. There were clear changes in the diet during the year. In March, when the food intake was reduced, copepods, mainly adult calanoids, formed the bulk of the food consumed. The most intensive consumption period lasted from June to late September. Chironomid pupae and planktonic cladocerans were the major summer food items. The largest individuals fed mainly on molluscs during summer. Zooplankton survey and consumption estimates did not show directly that the population density of copepodids and adults of Eudiaptomus graciloides were affected by whitefish predation although their population density decreased in spring. Nauplii, showing the maximum population density in April, were not consumed. The role of copepod consumption in the timing of Diphyllobothrium ditremum plerocercoid transmission to whitefish was also discussed. The increased copepod consumption rate during winter results in high energy intake but also an increased risk of food transmitted parasite infection.     

Food consumption and daily feeding periodicity: comparison between pelagic and demersal whiting in the North Sea

Pelagic North Sea whiting Merlagius merlangus fed at night, while demersal whiting fed by day. The estimated specific daily ration ranged from 4·38 to 7·84% in 1992 and from 3·99 to 10·31% in 1993 using the in situ rate of gastric evacuation. Using Andersen's evacuation model the specific daily ration ranged from 0·41 to 1·66% in 1992 and from 0·78 to 1·75% in 1993. The specific daily rations were significantly different where energy density of stomach content by length class of whiting was significantly different between the two layers and years. The fact that daily ration was related to prey composition and energy density of the prey and spatial distribution of the whiting, demonstrates the need for a sampling design that includes both pelagic and demersal layers when quantifying the food consumption of whiting.     

A bioenergetics model for an air-breathing fish, Channa striatus

Food consumption, standard metabolism, and growth of juvenile snakehead, Channa striatus, a cannibalistic and air-breathing fish were measured at 24–26 °C under controlled laboratory condition. Snakehead weighing 3.2–29.5 g were evaluated, and were fed smaller snakehead. Based on our observations, we determined bioenergetics relationships between specific food consumption, metabolic rates, and body weight. These values, along with other published parameter values allowed us to construct a bioenergetics model for snakehead. We then verified our model with growth and food consumption measurements from an independent feeding trial. Predicted fish growth closely matched observed growth. Our model underestimated cumulative food consumption when a constant activity value was used, but consumption estimates improved when we used non-constant activity values (1-5 times of standard metabolism). Predicted fish maintenance ration was 1.7% of body weight per day. Food conversion efficiency was greatest (0.59) when fed 2% body weight daily, but declined when daily consumption exceeded 6% body weight. This model provides a useful approach for assessing food requirements of snakehead under controlled condition. This revised version was published online in July 2006 with corrections to the Cover Date.