Cannibalism in pike fry, Esox lucius L.: some experiments with fry densities

Pike fry were kept in 0.054-m³ tanks at densities of 50, 100 and 150 (277, 555, 833 fry m⁻²) for seven weeks during which the development of social and feeding behaviour was observed. Zooplankton, macro-invertebrates and perch fry were provided sequentially as food; the pike fry were allowed to feed ad libitum.
Zooplanktivorous fry stopped growing at 22 mm whilst, in the presence of abundant suitable alternative prey, 1–4% turned cannibalistic at 5 weeks of age; cannibals subsequently grew rapidly (mean 1.88 mm day⁻¹). Cannibalism ensued in all tanks when the ratio of predator size: prey size was c .2:1.
Fry tended to space-out evenly in the tanks with no overt aggression or territoriality. Behaviour was typified by remaining still for long periods, particularly subsequent to the onset of cannibalism. Cannibals were attracted by fry movements which often initiated attacks.
Daily per capita mortality rates showed no crowding effects before cannibalism but significant density-dependent mortality due to cannibals. Consumption rates of cannibals varied between 0.63 and 6.0 fry per cannibal per day. Cannibals accounted for 54–96% of daily mortality in the experimental tanks. These results are discussed in relation to proposed mechanisms of pike population density regulation.     

Hydrocephalus and red-disease in pike fry Esox lucius L.

Two diseases of pike fry have been investigated, both causing serious losses in Dutch pike culture. In one of these diseases the lump developing on the head of the fry was associated with an internal hydrocephalus.
The other one, 'red disease' after the haemorrhagic areas in the trunk, may be identical with red-sore disease of pike.
The etiologies of both diseases are unknown.     

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 biology of pike, Esox lucius L., in a southern eutrophic lake

The biology of pike, Esox lucius L., in a southern productive lowland lake is described. Scales and opercular bones were difficult to read and interpret, and growth estimated had perforce to be based on scale readings. Growth of Slapton pike is average and intermediate between recorded extremes. Both male and female pike reached maximum condition in February, just prior to spawning. The population of pike of over 450 mm fork length in the lake was estimated at 870·6 ± 389·3 in 1975, and 950·4 ± 143.3 in 1976/77. This leads to the conclusion that the pike population has one of the highest biomasses per surface area of water ever recorded. Number of pike per unit area of surface was also high, despite the fact that a significant proportion of the population (fish under 450 mm) could not be adequately sampled. Mean instantaneous mortality rate was 0·53, and mean instantaneous survival rate was 0·59. The roach provided the main item of diet of the pike, with perch taken less readily. Immature pike ate a significant proportion of invertebrates, but roach was again a common feature of the diet of even small fish. Two spawning migrations were identified; at other times of the year, pike were non territorial. The majority of pike spawned in March. Fecundity of a sample of females was assessed. Overall sex ratio was 1:1.     

The annual reproductive cycle of pike, Esox lucius L., in two Scottish lakes

Histological changes in the ovaries of pike from two shallow, adjacent, Scottish lakes are described and related to a macroscopic scale of maturity stages and to changes in gonad weight. There was no evidence of pre-ovulatory degeneration although occasional residual oocytes were present in spent ovaries. The gonadosomatic index (GSI) was high in males from October to March, and in females it increased through the winter months to a maximum immediately prior to spawning. Maximum ovarian weight was 8.3–9.0% of total body weight compared with only 0.9–1.1% for testes. These GSIs were low compared with other reports for the British Isles, suggesting lower reproductive output. Spawning occurred in the first two weeks of April at a water temperature of 6.0–7.5° C. Fecundity was described from a regression of egg number (F) on fork length (f.l.): log₁₀F= -1.51 + 3.33 log₁₀ f.l. (cm). Mean egg diameter after preservation varied from 1.1 to 2.4 mm over the size range examined. Males were mature at age 2 at a length of ≥ 27 cm, while females first spawned at age 3 at a length of ≤ 28 cm, although a minority were mature at age 2. The sex ratio was 1.2 males: 1 female at Kinord and 1.4 : 1 at Davan. Greater egg production at Davan, 50 678 ha^?1, compared with Kinord, 33 557 ha^?1, was due to higher individual fecundity-at-age resulting from better growth. Fecundity (19.29 × 10³ calculated for a pike of 55 cm F.L.), relative fecundity of only 10–24 eggs per gram wet weight of adult, and GSI values were low compared with pike in North America and Eurasia. Food supply may restrict reproductive output of pike in Lochs Kinord and Davan. The abundance ratio of predator: prey is high with the main prey species, perch, present in low numbers, and the only alternative species is eel in these simple fish communities.     

Seasonal reproductive cycle of pike, Esox lucius L., from the River Danube

Seasonal changes in ovaries and testes of the pike, sampled in the Danube (at 1163 and 1175 km) were analysed in this research. According to the presence of developing and resting oocytes in histological sections of pike ovaries sampled during August, it is possible to determine which female will be sexually mature the following spring. Active vitellogenesis starts in September (with oocyte diameters of 514 ± 54 μm and 453 ± 68 μm, at the upstream and downstream locality, respectively), at which time the differences between the developing and resting oocytes are even more obvious. Active spermatogenesis starts in September and lasts for 2 months, and by November primarily spermatozoa can be seen in histological sections. The reproductive score points to differences in samples from the two localities that can be explained by the differing characteristics of the habitat. Values of absolute fecundity (AF) ranged from 524 to 123 896 eggs, and the mean value of relative fecundity (RF) was 40.4 ± 12.5.     

Cost of prey capture, growth rate and ration size in pike, Esox lucius L., as functions of prey wheight

Prey intake, amount of time feeding, pursuit and manipulation times and growth rate were studied in the pike, Esox lucius , in relation to prey weight. Predator and prey were observed in tanks at 15°C and with a 14:10 h LD photoperiod. A total of 444 captures were recorded from five 1 g prey weight groups varying from 0 to 4.9 g. Ten pike were used weighing between 50 and 149 g.
As prey weight increased, the number and weight of prey consumed decreased as did the time spent feeding, which became concentrated into the first trial of the day. Pursuit time did not vary with prey weight, but manipulation time and the weight of prey gained per unit pursuit plus manipulation time increased with prey weight.
The growth rate of pike was a positive function of ration size and a negative function of the amount of time spent capturing each gramme of prey (capture cost). Growth rate was also correlated with the mean length and weight of prey eaten.
The most profitable prey, as estimated from optimal foraging theory, need only be available once every 84 min for pike to maximize their net energy gain. The data show that feeding behaviour can have a direct influence on the reproductive success of pike through its effect on growth rate.     

The reproductive success of pike, Esox lucius: aspects of fecundity, egg density and survival

Power-law relationships have been estimated between fecundity and fish length, weight and age for pike from two gravel pit lakes (Main Lake and St Peter's Lake) in Buckinghamshire, England. Relative fecundity, estimated at 17.6 eggs g⁻¹ for Main Lake pike and 19.4 g⁻¹ for St Peter's Lake pike, did not differ significantly between the lakes. Population fecundity for the Main Lake was estimated at 10.6 million eggs and 6.4 million eggs in 1986 and 1987, respectively.
Experimental work involving samples of eggs from 18 artificially fertilized Main Lake female pike showed a significant relationship between egg diameter and female length, and a significant difference between mean egg diameters for 41-cm and 101-cm females. No significant relationships were found between mean egg size and mean fry size, nor between fry length and adult length. Mean size differences between newly hatched, 18-day-old and 41-day-old fry from the 41-cm and 101-cm females were not significant.
Samples of eggs taken at 2-day intervals from 6 April to 5 May in a pike spawning area of the Main Lake revealed clumped distributions of eggs, with average egg densities ranging from 1 to 47 eggs per 0.071 m² (14–671 m ²) on sand and silt substrata. The maximum egg density of 51 per 0.071 m² (729 m⁻²) was found on flooded grass. The stages of development of the eggs were identified over the sampling period. Survival from stage 1 to fry was estimated at 3–6%. Egg losses were estimated at 9–10% day⁻¹.
Egg survival experiments gave overall hatching success rates of 11 % on sand/silt compared with 2.5% on aquatic plants (1986), and 18.9% on undisturbed sand/silt compared with 7.9% on disturbed sand/silt (1987). The effect of siltation on egg hatching is discussed.     

The ultrastructure of the blood cells of the pike Esox lucius L.

The ultrastructure of pike peripheral blood cells, lymphocytes, thrombocytes, granulocytes, and monocytes is described. At present there are no reliable criteria for differentiating between round thrombocytes and small lymphocytes of fish on a routine basis. At the ultrastructural level thrombocytes could be clearly differentiated from lymphocytes by cytoplasmic canals and vesicles, marginal microtubules, and large glycogen deposits. Electron microscopic identification of thrombocytes was confirmed by examining the ultrastructural features of a purified thrombocyte fraction. In addition, a preliminary investigation of the structure of the haemopoietic cells in the thymus, anterior kidney, and spleen was carried out.     

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.     

Seasonal anatomical variations in the testes of European pike, Esox lucius L.

The seasonal development of the testes in European pike was examined using wild fish and biopsies from pike housed in tanks. The size of the tubules and the different cell types were measured and their histological appearance described. Four stages of development can be distinguished during the annual cycle. They are: (I) the stage of rest from June to August; (II) the stage of development from September to November with intensive spermatogenesis; (III) the stage of maturity from December to March/April with spermatogenesis completed; and (IV) the post-spawning stage from March to May.     

Experimental studies on some trophic relationships in juvenile pike, Esox lucius L.

A postprandial increase in ammonia nitrogen excretion and oxygen consumption rates was observed in juvenile pike fed a natural diet or an artificial dry diet. Specific growth rate of natural diet fed pike (2.4%) was lower than that of pike fed the artificial diet (3.1%). Fifty per cent of ingesta was evacuated within 5–6 h in pike of 25 mg body weight and 9–10 h in those weighing 150mg. Daily nitrogen excretion rates were related to body weight. Respiratory quotient and energy retention efficiency were affected by the nature of the diet ingested by pike. Parameters of the energy balance (losses, retention, increment due to feeding) were related to energy intake.     

Growth of pike Esox lucius L. in Lake Rubikiai (Lithuania)

The aim of the study was to determine the growth rate and growth parameters of pike Esox lucius L. in the mesotrophic Lake Rubikiai and to evaluate species growth in light of environmental conditions in Lithuanian lakes of different limnologic types. A total of 204 pike specimens were caught in two subsequent years, April 2005 and April 2006, in Lake Rubikiai. Six age classes (2–7 years of age) for males and 10 age classes (3–12 years of age) for females were identified by scale readings; 2‐ to 4‐year‐old males and 4‐ to 7‐year‐old females prevailed. Total length of pike ranged from 26.5 to 107 cm and weight from 106 to 7600 g. There were no statistically significant differences (Student’s t‐test, P > 0.1) between mean lengths and weights of males and females of the same age. Von Bertalanffy growth parameters for the entire population were: L_∞ = 131.7 cm, k = 0.1530 year^?1, t_o = ?0.0398 year. Calculated overall growth performance index ?′ was 3.42. The length‐weight relationship was described by the equation . The growth rate of pike in Lake Rubikiai was higher than in most other investigated Lithuanian lakes. Pike from Lake Rubikiai are characterized by average growth, compared to other populations from various geographical locations. No significant differences were established between the growth of pike in mesotrophic and eutrophic lakes. Pike growth rate did not correlate with lake surface area, maximum or mean depth, and negatively correlated with bogginess in mesotrophic lakes and limpidity and bogginess in eutrophic lakes (P < 0.05).     

The effects of insulin on plasma amino acid levels in the Northern pike, Esox lucius L

Cod and bovine insulin (2 i.u. kg⁻¹) were injected intra-arterially into cannulated Northern pike, Esox lucius L. and the levels of 13 plasma amino acids analysed by gas-liquid chroma-tography after 9 and 48 h. Both insulins caused an equally significant reduction in the level of each amino acid after 9 h which was followed by a recovery to near-control levels after 48 h. The data support the hypothesis that insulin plays a major role in the regulation of protein metabolism in this species.     

Reproduction effort versus the environment; case histories of Windermere perch, Perca fluviatilis L., and pike, Esox lucius L.

The theoretical limits of net reproduction rates for perch, Perca fluviatilis L. and pike, Esox lucius L, have been estimated from the upper and lower limits of fecundity, growth, mortality, age of maturity and biomass of parental stock, observed in Windermere over 40 years. Differences in egg numbers due to changes in fecundity were for perch × 1.5, for pike × 2. Assuming the same mortality, a fast growing cohort would produce more eggs during its lifespan than a slow growing, for perch × 2, for pike × 9. Changes in mortality from high to low resulted in more eggs, for perch × 7, and for pike (where high mortality includes natural and fishing mortality) × 10. Change in age at first maturity from 3 to 2 years was unimportant. Biomass of parental stock varied by × 6 for perch and × 3 for pike, and number of eggs laid varied by x2 for perch and × 5 for pike. The number of recruits at age 2 years varied by × 300 for perch and × 7 for pike. It was concluded that temperature in the first summer of life and predation were of major importance in regulating the number of recruits. Ricker recruitment curves and a simple model showed that perch, but not pike, could produce maximum recruitment from minimum parental stock. For perch, but not pike, compensatory density dependent mortality increased with increased parental stock. It was concluded that responses by the perch and pike populations to a changing environment can not be of sufficient magnitude to prevent wide fluctuations in recruitment.     

A biochemical genetic analysis of populations of the northern pike, Esox lucius L., from Europe and North America

Genetic variation as identified by starch gel electrophoresis and enzyme staining was studied in seven populations of the northern pike, Esox lucius L. Samples derived from populations in northern U.S., Canada, Swedish fresh water, Swedish Baltic, the Netherlands, England and Ireland were analysed for variation at between 10 and 26 enzyme-coding loci. Polymorphism was found to be restricted to three loci: Sod-1 (mitochondrial superoxide dismutase), Ldh-2 (heart-type lactate dehydrogenase) and Got-1 (cytoplasmic glutamateoxalate transaminase). Mean heterozygosity ( H ) over all seven population samples was estimated to be 0·019. This extremely low variability, relative to reported data for other fish species, is discussed in terms of the structure and possible origins of pike populations.