Bioenergetics of growth of a cyprinid, Phoxinus phoxinus: the effect of ration, temperature and body size on food consumption, faecal production and nitrogenous excretion

Food consumption, faecal production and nitrogen excretion by minnows, Phoxinus phoxinus , weighing 1–5.5 g were studied at five rations ranging from starvation to ad libitum and four temperatures ranging from 5 to 15°C.
The maximum rate of food consumption (C_max) was related to body weight ( W ) and temperature ( T ) by the relationship: C _max= aW^b1T^b2 . There were significant daily variations in C_max, which tended to decline over time. Absorption efficiency increased with increasing ration size and decreasing temperature. Body weight had no significant effect on the faecal production. The equation F = a C^b1e^b2 T described the relationship between faecal production ( F ), food consumption ( C ) and temperature. Ammonia-N predominated over urea-N in the excreta of most experimental fish. The proportion of urea-N in the total nitrogen excreted was generally higher at lower rations than at higher rations. Rates of nitrogen excretion increased with increased ration size and were, to a lesser extent, influenced by temperature. Body weight had no significant effect on the nitrogen excretion by feeding minnows. The equation N = a+b_lT+b₂C described the effects of food consumption and temperature on nitrogen excretion ( N ) other than urea-N excretion. The relationship between urea-N excretion ( N_u ), food consumption and temperature was described by the equation N_u= ae^b1T ((C+1) ^b 2.
On the average, 11 % of food energy was lost in faecal production and nitrogen excretion by minnows feeding on whiteworms, Enchytraeus spp.     

Consumption, growth and respiration of bleak, Alburnus alburnus (L.), and roach, Rutilus rutilus (L.), during early ontogeny

Three components of the energy budget, consumption ( C ), production ( P ) and respiration ( R ) in juvenile roach and bleak kept under controlled food ( Anemia salina , 2400 ind. l⁻¹) and temperature (20° C) conditions were measured in a study aimed at defining differences between the two species and elucidating the patterns of energy partitioning during ontogeny.
Daily food consumption rates (J day⁻¹ fish⁻¹) increased allometrically ( C =a W ^b) with body size ( W , mg dry weight) in both species. Covariance analysis indicated no differences in slope or intercept for the two regression lines ( P ≤ 0.05, n = 82). However, the two species grew at significantly different rates, roach faster than bleak.
The dependence of the respiration rate (μmol h⁻¹ fish ⁻¹) on body weight ( W ) can be described by an allometric function: R = a W ^b, where a ± 95% C.L. = 0.17 ± 0.15 for roach and 0.18 ± 0.20 for bleak. The slope for roach (b ± 95% C.L. = 0.78 ± 0.01) is slightly higher than that for bleak (0.69 ± 0.03).
Assimilation efficiency [AE = ( P + R ) C ⁻¹] was significantly higher in roach than in bleak. Different levels of AE correlated with differences in relative gut length (gut length as percentage of body length). Due to the shorter relative gut length above a weight of 5 mg, bleak has lower powers of digestion, which may explain lower production rates. These differences in energetic performance between the two species indicate mechanisms leading to niche differentiation in the early life history of the fishes.     

Some aspects of the reproductive biology of Spratelloides gracilis (Schlegel) in the Ysabel Passage, Papua New Guinea

Fecundity, length at first spawning, spawning seasonality and ovarian development of Spratelloides gracilis were determined by examining preserved ovaries from fish captured over a 5-year period. Fecundity ( F ) was estimated from the number of eggs in the most advanced ovarian mode and was related to fish length ( L ) by the function: log_e F =3.764 log_eL-7.308, and to weight ( W ) by: log_eW= 1.210 log_eW+7.337. Spawning was observed in most months of the year without any recognizable pattern between the years. Ova diameter frequencies were bimodal but the smaller mode remained stationary regardless of the position of the larger mode. This was interpreted as evidence that individual fish may spawn only once. From data on egg length versus fish length and gonad index versus fish length it was estimated that fish were first capable of spawning at 45 mm (fork length).     

Growth studies with fish—overcoming the problems of size variation

The relationship between growth rate and fish size is described by the equation log_e G _w= a —0·4 log_e W, where G_w is the specific growth rate and W is fish weight. Since the intercept (a) represents the log_e G_w of a fish unit size, the relationship presents a method allowing comparison of data from experiments involving fish of different sizes. The application of the method is demonstrated by examining the effects of environmental temperature on growth rates of cod, Gadus morhua , and it is suggested that the optimum temperature for growth of cod is 13–15° C.     

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

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

Gastric emptying rate in Pleuronectes platessa L.

X-ray studies of gastric emptying of Pleuronectes platessa force-fed a moist, artificial diet give the relationship:
where GET is gastric emptying time (h), S is meal size (g), W is body weight (g) and T is temperature (°C).
The corresponding gastric emptying curve is predicted to be:
where stomach contents (g net weight) are S ₀ at start and S_t at time th. Direct observations of residual stomach contents using serial slaughter are in close agreement with the predicted curve, save for small plaice (below 50 g body weight) which empty at only 50–60% of the rate found in larger fish.
The small effects of body weight on gastric emptying rate may reflect the relatively small role of the stomach [Volume (ml) = 0.024 body weight (g)] in digestion when compared with other flatfish.
The emptying curve established here can be used in field studies to estimate feeding rate on natural food items, provided fish greater than 50 g body weight are used and the energy density of the food is greater than 0.5 kcal/g wet weight. Smaller fish will have emptying rates which are only 30–60% of those found in larger fish; the change appears to be abrupt and corresponds to the size at which young plaice move offshore from their nursery grounds.     

Respiratory metabolism of Utah chub, Gila atraria (Girard) and speckled dace, Rhinichthys osculus (Girard)

Measurements of active, standard and routine oxygen consumption were made for 207 Utah chub at 6°, 9°, 12°, 18° and 22° C; and 123 speckled dace at 4°, 8°, 12° and 18° C. These were expressed as mathematical functions of water temperature and size of the fish. The mean slopes of log_e oxygen consumption (mg O₂/h) and log_e wet weight (g) were 0.771, 0.526, and 0.619 for active, standard, and routine rates respectively for Utah chub, and 0.943, 0.486, and 0.683 respectively for speckled dace.     

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.     

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.     

Food consumption and growth of two piscivorous fishes, the mandarin fish and the Chinese snakehead

Rates of maximum food consumption and growth were determined for immature mandarin fish Siniperca chuatsi (47·2—540·2 g) and Chinese snakehead Channa argus (45·0—546·2 g) at 10, 15, 20, 25, 30 and 35) C. The relationship between maximum rate of food consumption ( C max), body weight ( W ) and temperature ( T ) was described by the multiple regression equations: In C max=−4·880+0·597 In W +0·284 T −0·0048 T ² for the mandarin fish, and In C max=−6·718+ 0·522 In W +0·440 T −0·0077 T ² for the Chinese snakehead. The optimum temperature for consumption was 29·6) C for the mandarin fish and 28·6) C for the Chinese snakehead. The relationship between growth rate ( G ), body weight and temperature was ln( G +0·25)=−0·439−0·500 ln W +0·270 T −0·0046 T ² for the mandarin fish, and ln( G +0·25)=−6·150+ (0·175−0·026 T ) ln W +0·571 T −0·0078 T ² for the Chinese snakehead. The weight exponent in the growth–weight relationship was −0·83 for the mandarin fish, but decreased with increasing temperature for the Chinese snakehead. The optimum temperature for growth was 29·3) C for the mandarin fish, but tended to decrease with increasing weight for the Chinese snakehead, being 30·3) C for a 45-g fish, and 26·1°C for a 550-g fish.     

Gastric emptying in Limanda limanda (L.) and the return of appetite

Limanda limanda (35–225 g) fed during daytime when trained to use demand feeders under natural photoperiods. Under continuous illumination different feeding rhythms developed which were detected by periodogram analysis. Interfeeding periods (P hours) varied with body weight (W grams) and temperature (T° C) over the range 6.5–15° C as: In P=4.4 – 0.167 (In W) – 0.05 T. Daily food intake (D grams increased with both temperature and size (In D = 0.099 T + 0.579 In W – 3.49). Relative daily intake (R as % body weight, b.w) increased with temperature but decreased with size (In R = 0.099T–0.41 In W + 1.05). These changes in daily intake are primarily caused by changes in feeding frequency; the amount eaten at a meal is relatively constant save at low temperatures (6.5° C) when appetite is suppressed. Dilution of the food with kaolin (from 4.8 to 2.2 kcal/g) did not cause a compensatory increase in meal size or frequency. Gastric emptying time (GET hours) measured by X-radiography changed with body weight, temperature and relative meal size (M % b.w.) as In GET = 0.68 In M + 0.39 In W – 0.035T + 1.46. With the exception of the lowest temperature (6.5° C) interfeeding periods closely followed GET (P = 1.09 GET – 2.6) supporting the proposal that stomach fullness is a major factor controlling appetite in the dab.     

A desk study of the relationship between temperature and hatching time for the eggs of five species of salmonid fishes

SUMMARY. Information on temperature (T°C) and time from fertilization to 50% hatch ( D days) for five species of salmonid fishes has been used to assess several mathematical models relating D and T . No single equation gave the best fit to all five data sets. The power law with temperature correction (α), log₁₀₁ D = log₁₀ a + b log₁₀ ( T - α) and the quadratic, log₁₀ D = log₁₀ a + bT + b ₁ T ² (where a, b, b ₁, and α are constants), each accounted for over 97 % of the variance of D and were good fits to the observed data points for all five species. There was little difference between the predictions obtained from these two equations within the range of observed temperatures. Therefore, the simpler power-law model is preferred. However, there were substantial within-species differences between values of D predicted from extrapolations of the two models from 2 or 3°C down to 0°C. When more data for low temperatures become available it will be possible to make a more objective choice of model.     

The link between respiratory capacity and changing metabolic demands during growth of northern pike, Esox lucius L.

The relationship between metabolic rate of pike (Y, mgO2) and body weight (X, g) over the range 40–1291 gat 15° C is of the form: Y=aX^b. For resting metabolic rate (V_o2, rest), the scaling coefficient, b , is 0.80 and for maximum metabolic rate measured after exhaustive swimming (V₀₂, max), b is 0.99. Factorial metabolic scope (V₀₂, max/ V₀₂, rest) increases with body weight. Peak postprandial oxygen consumption (V₀₂, ASDA) is a constant multiple of V₀₂ rest for any discrete meal (expressed as % of body weight) up to 10% body weight. V₀₂ASDA after a single meal can utilize the entire metabolic scope (V₀₂, max—V₀₂, rest) of juvenile but not adult pike.     

The influence of body size and temperature on oxygen consumption of the European eel, Anguilla anguilla

The effect of temperature and body size on oxygen uptake of European eels, Anguilla anguilla , was studied. The mass specific oxygen uptake of large eels was lower at all temperatures than that of small eels. The effect of temperature on metabolism was greater on small eels than on large eels. The relationship between oxygen consumption and body size is described by the equation M = a W^b , where M is metabolism (O₂ h^-1 ), W is body weight (g), b is the slope of the function which describes the relationship between body weight change and metabolism, and a is the temperature-dependent constant of the equation. In this study it was found that increased temperature caused an increase in a and a decrease in b .     

Growth rates of Lutjanidae (snappers) in tropical Australian waters

Growth rates of three commercial species of Lutjanidae were estimated from sections of vertebrae cross-checked with scales. The von Bertalanffy growth constants for Lutjanus malabaricus were K =0.168, L_∞ =70.7 cm standard length (S.L.). For Pristipomoides multidens these were K = 0.219, L _∞= 59.1cm, and for Pristipomoides typus K =0.254, L _∞= 51.5 cm. Length/weight was analysed in the form W= aL^b where W was fresh weight (g) and L = S.L. (cm). The constants were: L. malabaricus, a = 0.041, b = 2.842; P. multidens, a =0.032, b = 2.897; P. typus, a = 0.038, b = 2.822. The growth rates were considered low in comparison with demersal fish counterparts from temperate seas.     

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.     

Response of beech (Fagus sylvatica) to elevated CO2 and N: Influence on larval performance of the gypsy moth Lymantria dispar (Lep., Lymantriidae)

Two-year-old beech seedlings were kept from germination to bioassays with Lymantria dispar under the following conditions: ambient CO₂/low N, elevated CO₂/low N, ambient CO₂/elevated N, and elevated CO₂/elevated N. The effect of these growing conditions of the trees on the performance of the defoliator L. dispar was studied 2 years after initiating the tree cultivation. The developmental success of third-instar larvae of L. dispar was characterized by the weight gained, percentage of weight gain, relative growth rate (RGR), relative consumption rate (RCR), and efficiency of conversion of ingested food into body substance (ECI). Contrary to our expectations, additional N-fertilization did not increase and elevated CO₂ did not delay larval growth rate. However, the environmental treatments of the beech seedlings were found to affect the larval performance. Larvae consumed significantly higher amounts of foliage (RCR) on beech trees under controlled conditions (ambient CO₂ and low N) compared to those under elevated CO₂ and enhanced N. The opposite was true for ECI. The lowest efficacy to convert consumed food to body substance was observed under control conditions and the highest when the larvae were kept on beech trees grown under elevated CO₂ and additional N-fertilization. These opposite effects resulted in the weight gain-based parameters (absolute growth, percentage of growth, and RGR) of the gypsy moth larvae remaining unaffected. The results indicate that the gypsy moth larvae are able to change their ECI and RCR to obtain a specific growth rate. This is discussed as an adaptation to specific food qualities.     

Roles of endothermy in niche differentiation for ball-rolling dung beetles (Coleoptera: Scarabaeidae) along an altitudinal gradient

Abstract.  1. An analysis of whether niche differentiation in ball-rolling dung beetles can be explained by the way in which they regulate their body temperature was conducted.
2.  A priori assumptions were: (i) if thermoregulation affects niche partitioning, sympatric species must have different endothermic strategies that minimise encounters; or, alternatively (ii) if two co-occurring species show the same thermoregulation pattern and their flight periods overlap, they might be avoiding competition by exhibiting different resource preferences or different food relocation behaviour.
3. The ball-rolling dung beetles studied showed a hierarchical structure based on the species' endothermic capacity, measured as temperature excess [ T _ex= difference between body temperature ( T _b) and ambient temperature ( T _a)]. Those with a high T _ex (10–15 °C) were located exclusively at altitudes >1000 m a.s.l. On the coastal plains, species with a high T _ex were restricted to flying at night when the T _a was lower. Species with a lower T _ex (less than 10 °C higher than T _a) were found in the coastal plains zone.
4. Where there was sympatry with similar trophic habits, the species involved showed very different thermal niches, and where there was significant overlap of thermal niches between sympatric species, trophic habits of species were very different.
5. The results suggest that it is possible to use the concept of the thermal niche as a tool to explain interspecific interactions and the spatial distribution of species.     

Feeding and growth of early juvenile Japanese sardines in the Pacific waters off central Japan

Larval and early juvenile growth was backcalculated for individual Japanese sardines Sardinops melanostictus using the biological intercept method based on the allometric relationship between otolith radii and fish lengths. Sardines grew at 0·81 mm day⁻¹ during the larval stage. In the early juvenile stage, they grew from 32·3 to 45·4 mm fork length ( L ) over a 20-day period (0·64mm day⁻¹). Using the observed relationship between L and wet body weight ( W ), W = 0·00942 L ^2.99, W of the sardine juveniles was calculated to increase from 306 to 832 mg during the 20-day period. The carbon (C) requirement to achieve this growth in weight was estimated to increase from 5·7 to 9·6 mg day⁻¹. Stomach contents of the sardines were composed mostly of copepods (73%) and larvaceans (25%). Wet stomach content weight ( W_s ) was expressed by a power function of the W , W_s=0·731 W ^0·658. Carbon and nitrogen constituted 41·7 ± 1·5 and 10·0 ± 0·4% of the dry W_s , respectively. Stomach C content increased from 2·0 to 3·9 mg during the 20-day period. Three to four cycles of the daily turnover of stomach contents during the 16 h of daytime, corresponding to a gastric evacuation rate of 0·2–0·3 h⁻¹ under continuous feeding, met the C requirement to achieve the backcalculated growth in early juvenile sardines. The Kuroshio frontal waters seem to provide Japanese sardine juveniles with favourable growth conditions.     

Routine metabolism of juvenile spot, Leiostomus xanthums (Lacépède), as a function of temperature, salinity and weight

Routine oxygen consumption rates of juvenile spot, Leiostomus xanthums , were measured over a range of temperatures, salinities and fish weights. As predicted, Q O₂ increased with temperature and decreased with body weight. However, Q O₂ decreased with decreasing salinity and did not show the expected minimum at isosmotic concentrations. The data are best described by the relationship: log₁₀ Q O₂ (mg O₂ g⁻¹ h⁻¹) = 0.129 log_lo salinity (%0) + 1.604 log₁₀ temperature (°C)-0.1401og₁₀(g)-2.767.