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.     

Direct measurements of metabolism, activity and feeding behaviour of pike, Esox Zucius L., in the wild, by the use of heart rate telemetry

Heart rate telemetry records of up to 5 days duration were obtained from pike living in Lochs Kinord and Davan, Scotland. Applying metabolic rate correlations it was found that mean metabolic rate (R) was 1.5 times standard metabolic rate (R_a), The fish rarely worked near their metabolic limits. Activity metabolism (R_a) was much higher than estimates based on mean swimming speed and comprised up to 10% of R. Most activity metabolism was the result of localized bursts of activity. Less than 10% of activity showed evidence of oxygen debt. Specific Dynamic Action or feeding metabolism (R_f) comprised 15–25% of R. Food intake estimated from heart rate was 1.5% wet body weight day⁻¹, consumed in the form of small items captured during the day and digested during the afternoon and night.     

The limitations of heart rate as a predictor of metabolic rate in fish

Although telemetered heart rate (f_H) has been used as a physiological correlate to predict the metabolic rate (as oxygen consumption, V?O₂) of fish in the field, it is our contention that the method has not been validated adequately for fish. If f_H in fish is to be used to estimate V?O₂, a single linear (or log-linear) relationship must be established for each species between the two variables which allows V?O₂ to be predicted accurately under all environmentally relevant conditions. Our analyses of existing data indicate that while a good linear (or log-linear) relationship can be established between f_H and V?O₂, the conditions under which the relationship applies may be quite restricted. Physiological states and environmental factors affect the relationship between f_H and V?O₂ significantly such that several curves can exist for a single species. In addition, there are situations in which f_H and V?O₂ do not covary in a significant manner. In some situations f_H can vary over much of its physiological range while V?O₂ remains constant; in others V?O₂ may vary while f_H is invariate. The theoretical basis for this variability is examined to explain why the use of telemetered f_H in predicting V?O₂ of fish may be limited to certain specified applications.     

Foraging among cannibals and kleptoparasites: effects of prey size on pike behavior

The northern pike (Esox lucius) is an important and selective piscivorethat chooses smaller prey than predicted from energy / timebudgets. In a laboratory experiment, we investigated pike predatorybehavior to explain this selectivity. Northern pike feedingon different prey sizes in aquaria were observed when foragingalone, when in the presence of chemical cues from similar-sizedor larger conspecifics, and when in the presence of conspecifics thatwere allowed to interact with the focal pike. The results showthat prey handling time increases with prey size and that theduration of manipulating and handling prey inflicts a risk ofexposure to cannibals and kleptoparasites on the pike. Therefore,the risk of falling victim to cannibals or kleptoparasites increaseswith prey size. Attracting and experiencing intraspecific interactorscan be regarded as major fitness costs. Chemical cues from foragingconspecifics have only minor effects on pike foraging behavior.Furthermore, the ability to strike and swallow prey head first improvespike predatory performance because failing to do so increases handlingtime. Our findings emphasize the increasing potential costswith large prey and explain previous contradictory suggestionson the underlying mechanisms of behavior, selectivity, and trophiceffects of northern pike predation.     

Early life characteristics of pike, Esox lucius, in rearing ponds: temporal survival pattern and ontogenetic diet shifts

Survival, biomass and diet of pike, Esox lucius , larvae and juveniles were studied over 3 years by stocking free embryos at a density of six fish m⁻² in 12 drainable outdoor ponds. The ponds were sequentially drained at six larval/juvenile developmental stages, up to a total length (T.L.) of 139 mm. The mean rate of survival at harvest decreased irregularly over time and the highest mortality rates were recorded during the early larval period (13 to 27 mm t.l .) and two intervals of the juvenile period (46 to 99 mm T.L. and 121 to 139 mm t.l .). Mean biomass increased dramatically between 46 mm T.L. (19.8 kg ha⁻¹) and 121 mm T.L. (181.8 kg ha⁻¹) and stabilized between 121 and 139 mm t.l . Sharp increases in the mean weight coefficient of variation were recorded during the early larval period (13 to 27 mm t.l .) and between 74 and 121 mm t.l . (development of cannibalism). Diet breadths were relatively narrow in pike larvae and reached maximum levels in 99 mm t.l . juveniles. Average-sized pike exhibited a sequence of size-dependent shifts from a diet composed primarily (in terms of weight) of micro-crustaceans (at 13 mm t.l .), to chironomid larvae (at 74 to 99 mm t.l .), and finally macrocrustaceans (at 121 to 139 mm t.l .). Cannibalism was detected first among the largest fish at the 74 mm t.l . stage. Between-year diet similarity at various developmental stages was consistently high. In 70 mm t.l . fish harvested from different ponds, we found significant among-pond differences in diet composition; however, similar trends of diet changes in relation to fish size were observed from pond to pond. Our results are discussed in light of existing knowledge of young pike trophic ecology and current aquaculture practices.     

The growth of young pike in small Finnish lakes with different acidity-related water properties and fish species composition

Pike growth was generally slower in small Finnish lakes than has been reported from large lakes or brackish coastal waters. Growth was unrelated to pH, alkalinity, COD, or conductivity of the water, but length of 1–2-year-old pike correlated positively with water colour (i.e. humic content), nitrogen content, and phosphorus content. These water properties also correlated with each other. Growth of 1–2-year-old pike was negatively correlated with lake area in lakes smaller than 20 ha.
In lakes with a strong roach population or without ruffe the 0 group pike grew faster than in lakes without roach or with ruffe. The growth of pike aged 1–4 years in two very acid and practically fishless lakes did not differ clearly from the rest of the lakes, although the pike were forced to forage for abundant acid resistant invertebrates like dragonfly nymphs (Odonata) or Asellus aquatlcus .
The results of this work support earlier studies where the growth rate above all depends on water temperature and food availability. Acidity was not shown to affect the growth rate of pike.     

Individual specialization and trophic adaptability of northern pike (Esox lucius): an isotope and dietary analysis

Northern pike (Esox lucius) are often considered to be specialist piscivores, but under some circumstances will continue to eat invertebrates as adults. To examine effects of fish assemblage composition on the trophic ecology of pike, we combined stable isotope analysis (SIA) of carbon and nitrogen and stomach content analysis (SCA) on pike from five lakes in northern Alberta, three of which contain only pike (“pike-only”) and two that also contain yellow perch (Perca flavescens) or white sucker (Catostomus commersoni) (“pike-other”). Fish were more important as prey and empty stomachs, which often characterize piscivores, were significantly more frequent in pike-other than in pike-only lakes. However, even though invertebrates were more important for pike in pike-only lakes, SIA and SCA indicated that invertebrates were also an important component of pike diets in pike-other lakes. SIA and SCA also revealed considerable intrapopulation variation in trophic ecology, with individuals in some populations differing by as much as two trophic levels. Comparisons of stomach contents and isotope signatures of the same fish suggested that within these variable populations, specialization on invertebrates or fish was a long-term trait of some individuals. SIA indicated that trophic position increased and diets shifted to a greater importance of littoral prey as pike grew in pike-only lakes, but not in lakes with other fish present. Trophic adaptability in northern pike is expressed at both the population level, where the trophic ecology is sensitive to differences in prey regimes, and at the organismal level, in the form of intrapopulation variation and individual specialization. Received: 1 July 1998 / Accepted: 3 February 1999     

The response of bleak to predator odour of unfed and recently fed pike

Feeding and growth responses of bleak Alburnus alburnus to the odour of unfed and recently fed pike Esox lucius were quantified in laboratory experiments. Bleak reacted more strongly to the latter than to the former and feeding rates were lowered by 35 and 20%, respectively. The decreases in feeding rates were not an effect of the absence of fish from the feeding place, but of a lowered feeding efficiency due mainly to diminished time spent foraging. The lowered consumption rate of bleak exposed to the odour of recently fed pike caused a decrease in their individual growth rate. The stronger reaction to the recently fed pike stimulus supports the view that alarm substance is a marker for predator odour. In the absence of any further signal from the predator, the bleak became habituated to the predator odour and showed the reaction for only 5 days.     

Survival and sublethal responses of early life stages of pike exposed to low pH in artificial post-mining lake water

The threshold pH for survival of early life stages of pike Esox lucius in post-mining lakes was determined in a laboratory experiment using artificial water characterized by high concentrations of dissolved Ca, Al, Fe, Mn and SO2/4⁻. At pH 3·50–4·00, high mortality was observed before hatching. At pH 4·00 and 4·25, hatching rates were reduced compared with the controls (pH 7·40), and many embryos died in a partly hatched state. Hatching success of embryos exposed to pH levels of 4·50 or higher was not affected. However, at pH 5·00 many newly hatched embryos were deformed. Furthermore, pike exposed to pH 4·00–5·00 did not start feeding. At pH 4·75 and below, mortality increased to 100%, and at pH 5·00, only few eleutheroembryos which were in extremely poor condition survived to the end of the experiment. At pH 5·50, survival was in the same range as in the control group, but growth was reduced. Therefore, early life stages of pike are expected to survive in Lustian post-mining lakes when a pH of 5·50 and above is reached and maintained.     

Seasonal changes in some blood chemistry parameters and in relative liver and gonad weights of pike (Esox lucius L.) from the River Danube

Samples from a natural population of pike (Esox lucius L.) from the River Danube were used in a 12-month study to determine seasonal variations in biochemical parameters of pike blood sera, hepatosomatic index (HSI) and gonadosomatic index (GSI). The ranges of enzyme activities for sample means were: aspartate aminotransferase (ASAT) 252.0–583.8 U 1⁻¹, alanine aminotransferase (ALAT) 4.9–11 -2 U1 and alkaline phosphatase (AP) 39.5–91.8 U1⁻¹. The ranges of other parameters analysed in serum were total protein 27.7–40.1 g 1 ¹, urea 0.57–l.52 mmol 1 ¹ and creatinine 21.2–118.6 μmol 1 ¹. The range of sample means for HSI and GSI were 1.28–4.16 and 0.07–20.2 respectively. Temperature ranged from 4.5 to 23.5°C. The activity of serum AP was positively correlated to water temperature in males only, while urea and creatinine showed a positive correlation to water temperature in individuals of both sexes. GSI was correlated significantly with HSI in females. Total protein reached the lowest values during the spawning period, while creatinine levels depended on both the sex and season.     

Effects of northern pike on patterns of nest use and resproductive behavior of male fathead minnows in a boreal lake

We conducted a two-part study to assets predator avoidance byreproductive male fathead minnows (Pimephales promelos) subjectedto predation threat from northern pike (Esox lucius). First,we determined if patterns of nest use by egg-guarding male minnowsin a boreal lake were related to pike densities. We samplednorthern pike and identified four areas of "high pike-density"and three areas of "low pike-density." We censused natural nestsand placed nest boards in these areas. We found eggs on naturalnests more frequently in areas with low densities of pike thanin areas with high densities of pike. However, we could notfully explain the distribution of nests by predation risk. Second,we evaluated the behavioral response of egg-guarding males toa control stimulus (a piece of wood) or a live pike in a wirecage. We used time to return to the nest after a stimulus asa measure of risk taking. Males took different amounts of riskbased on predation threat; males in the predator treatment tooklonger to return to their nests than control males. Risk takingwas not related to the number or age of the eggs but to distanceto nearest egg-guarding neighbor; males with close neighborsreturned sooner than more isolated males. Males in the predatortreatment had lower total activity and egg rubbing than controlmales after they returned to their nests. We conclude that malefathead minnows altered their reproductive behavior in waysthat reduced predation risk, but the cost of predator avoidancemight include egg predation, lost mating opportunities, or usurpationof nests     

Field metabolic rates of rainbow trout estimated using electromyogram telemetry

Field metabolic rates (FMR) of five rainbow trout were estimated using electromyogram (EMG) telemetry of the axial muscle. A series of laboratory experiments indicated that the EMG transmitter output was related strongly to total oxygen consumption of the fish over a wide range of swimming speeds and temperatures. No differences were evident when the oxygen consumption v . EMG relationship for routine swimming was compared with that for forced swimming. FMR was assessed on two time scales, revealing diel patterns and seasonal patterns. On the diel scale, the FMR pattern could be classified as crepuscular. At the upper and lower limits of temperature tolerance, the diel pattern was less distinct. On the seasonal scale, mean daily FMR was strongly dependent on mean environmental temperature. Comparisons between FMR and laboratory derived estimates of standard and maximally active metabolism indicate that the rainbow trout in the field utilize <20% of the available scope for activity.     

Association between heart rate variability and training response in sedentary middle-aged men

The effect of exercise training on heart rate variability (HRV) and improvements in peak oxygen consumption ( _peak) was examined in sedentary middle-aged men. The HRV and absolute and relative _peak of training (n = 19) and control (n = 15) subjects were assessed before and after a 24-session moderate intensity exercise training programme. Results indicated that with exercise training there was a significantly increased absolute and relative _peak (P < 0.005) for the training group (12% and 11% respectively) with no increase for the control group. The training group also displayed a significant reduction in resting heart rate; however, HRV remained unchanged. The trained subjects were further categorized into high (n = 5) and low (n = 5) HRV groups and changes in _peak were compared. Improvements in both absolute and relative _peak were significantly greater (P > 0.005) in the high HRV group (17% and 20% respectively) compared to the low HRV group (6% and 1% respectively). The groups did not differ in mean age, pretraining oxygen consumption, or resting heart rate. These results would seem to suggest that a short aerobic training programme does not alter HRV in middle-aged men. Individual differences in HRV, however, may be associated with _peak response to aerobic training.     

Pike and red swamp crayfish: a new case on predator-prey relationship between aliens in central Spain

Pike in the Ruidera Lakes (central Spain) chiefly ate the recently introduced crayfish Procambarus clarkii . It was the dominant prey in occurrence, number and biomass for every size class and season. Likewise, number (up to 27) and total weight of ingested crayfish were directly related with pike size. The principal prey fish were all introduced species (except the endangered Blennius fluviatilis ), most native species having disappeared since pike were introduced in 1953. (c) 1996 The Fisheries Society of the British Isles     

The rate of metabolism in marine animals: environmental constraints, ecological demands and energetic opportunities

The rates of metabolism in animals vary tremendously throughout the biosphere. The origins of this variation are a matter of active debate with some scientists highlighting the importance of anatomical or environmental constraints, while others emphasize the diversity of ecological roles that organisms play and the associated energy demands. Here, we analyse metabolic rates in diverse marine taxa, with special emphasis on patterns of metabolic rate across a depth gradient, in an effort to understand the extent and underlying causes of variation. The conclusion from this analysis is that low rates of metabolism, in the deep sea and elsewhere, do not result from resource (e.g. food or oxygen) limitation or from temperature or pressure constraint. While metabolic rates do decline strongly with depth in several important animal groups, for others metabolism in abyssal species proceeds as fast as in ecologically similar shallow-water species at equivalent temperatures. Rather, high metabolic demand follows strong selection for locomotory capacity among visual predators inhabiting well-lit oceanic waters. Relaxation of this selection where visual predation is limited provides an opportunity for reduced energy expenditure. Large-scale metabolic variation in the ocean results from interspecific differences in ecological energy demand.     

Acid phosphatases of Esox lucius: tissue distribution and partial characterization

The presence of different acid phosphatase classes was examined in organs and tissues of the fish Esox lucius . The enzymes were separated by Sephadex G-200 column chromatography, and characterized as regards tissue distribution, molecular weight, metal ion activation, inhibition, pH optima, temperature sensitivity, rate of hydrolysis and substrate specificity. High and low molecular weight acid phosphatases as well as high and low molecular weight Zn²⁺-dependent acid phosphatases were present in pike organs and tissues. In addition, their biochemical properties and tissue distribution are very similar to that of respective bird and mammalian enzymes.     

Post-exercise metabolic rate in Atlantic cod and its dependence upon the method of exhaustion

This study tests whether or not post-exercise oxygen consumption rates ( M o₂) in fish are dependent upon how exhaustion is induced. A group of eight Atlantic cod ( Gadus morhua ) were each exercised using (1) a critical swimming speed ( U _crit) protocol, (2) an exercise protocol designed to measure anaerobic capacity of fish ( U _burst), and (3) a protocol in which the fish were chased to exhaustion manually. M o₂ was measured for a 2-h period following exhaustion induced by all three exercise regimes ( U _crit, U _burst and chase). Post-exercise M o₂ following exhaustion from the U _burst and chase protocols were significantly higher than post-exercise M o₂ following the U _crit protocol. Each fish during the U _crit protocol exhibited maximal M o₂ during exercise rather than during recovery, yet 75% of the fish during U _brust recovery and 100% during chase recovery exhibited M o₂ higher than that measured during U _crit exercise. These results, as well as the large interindividual variations in M o₂ among the eight fish, show that post-exhaustion M o₂ is specific to the exercise regime employed, thus, investigators must exercise caution when combining results from different exercise protocols and/or individuals.