Generalization of learned predator recognition: an experimental test and framework for future studies

While some prey species possess an innate recognition of their predators, others require learning to recognize their predators. The specific characteristics of the predators that prey learn and whether prey can generalize this learning to similar predatory threats have been virtually ignored. Here, we investigated whether fathead minnows that learned to chemically recognize a specific predator species as a threat has the ability to generalize their recognition to closely related predators. We found that minnows trained to recognize the odour of a lake trout as a threat (the reference predator) generalized their responses to brook trout (same genus as lake trout) and rainbow trout (same family), but did not generalize to a distantly related predatory pike or non-predatory suckers. We also found that the intensity of antipredator responses to the other species was correlated with the phylogenetic distance to the reference predator; minnows responded with a higher intensity response to brook trout than rainbow trout. This is the first study showing that prey have the ability to exhibit generalization of predator odour recognition. We discuss these results and provide a theoretical framework for future studies of generalization of predator recognition.     

The effect of prey type and density on the foraging efficiency of juvenile perch, (Perca fluviatilis)

The foraging efficiency of juvenile perch (Perca fluviatilis), feeding on two types of prey, was studied in laboratory experiments. Waterfleas (Daphnia magna) and phantom midge larvae (Chaoborus flavicans) were offered in a range of densities, either separately or combined. Perch fed more efficiently on each prey type separately than when both were mixed. Foraging efficiency decreased with an increase of mixed prey density with both prey types present in equal numbers, but also when the proportion of Chaoborus increased. This could be caused by the existence of different hunting techniques, each of which is fully efficient in the presence of one prey type only. In the presence of two prey types, the predator constantly has to switch from one hunting technique to another.     

Distribution of perch (Perca fluviatilis L.) during their first year of life in Lake Constance

The distribution and behaviour of larval and juvenile perch (Perca fluviatilis L.) were studied for two years in large, deep Lake Constance. After hatching larvae were transported by water currents to the open water. The majority of larvae remained in the pelagic zone for about one month. In both years, their return to the littoral zone coincided with the decline of pelagic zooplankton abundance. After returning to the littoral zone, juveniles stayed among submerged macrophytes within 5 m depth and lived apart from larger perch which lived at depths of 6–20 m. By late summer, juveniles changed their distribution pattern: during the day they stayed intensively close to piers and ports, but increased their swimming activity at dusk, cruising among shallow and deep waters and feeding on zooplankton, and rested on the bottom at night. This behaviour appears to be related to the decrease of inshore food resources and to the presence of predators in deeper water. 0⁺ perch left the littoral zone and moved into deep waters when autumnal mixing began in late October. They overwintered near the bottom at depths of more than 30 m. During most of the year, juvenile and adult perch were separated from each other. But as soon as they occupied the same habitat, the occurrence of cannibalism increased.     

The predator-prey relationship of perch,Perca fluviatilis,larvae and zooplankton in two Scottish lochs

Synopsis Food consumption of perch larvae and the impact of this on zooplankton were examined in two adjacent shallow Scottish lochs. Maximum annual abundance of zooplankton occurred in mid-May at L. Kinord with minimum values in mid-June. Copepods were prominent in spring but were followed by a multi-species community of cladocerans and rotifers in summer. At L. Davan zooplankton biomass remained high through summer with cladocerans dominating andDaphnia longispina the most frequent species. Availability of food items was a principal factor governing feeding behaviour of larvae. Copepodite stages were initially the most common item in the diet in L. Kinord in 1976 and 1977 and rotifers the principal food in June 1977, reflecting the dominance of these items in the zooplankton. Cladocerans were dominant in the plankton community in L. Davan and constituted the greater part of food intake. Overlying this general pattern there was an increase in the size of food items taken by larvae with time and also a definite pattern of food selection for copepods, with initially selection for smaller copepodite stages and later for larger stages and adults. On most occasions larvae selected forCyclops strenuus abyssorum andPolphemus pediculus and selected againstDaphnia longispina. The reduction in the total zooplankton biomass attributed to perch larvae was minimal, with the exception of mid-June at L. Kinord in 1976. However, predation on particular species and copepodite stages was occasionally intense and may have impacted the zooplankton populations.     

Survival of fathead minnows after injury by predators and its possible role in the evolution of alarm signals

Synopsis Some mechanisms proposed to account for the evolution of animal alarm signals require that the signal sender survive in order to benefit. The ostariophysan alarm pheromone system requires mechanical damage for the release of the alarm pheromone. A natural situation is described in which up to 16% of the fathead minnows in a population have survived damaging encounters with predators. This indicates that post-signal selective benefits can operate in the evolution of the ostariophysan alarm system.     

Short-term effects of liming on perch Perca fluviatilis populations in acidified lakes in South-West Sweden

Gill net catches of perch Perca fluviatilis in eight acidified lakes immediately before liming are compared] with catches in the same lakes two years after liming. Simultaneously, six control lakes, not limed, were investigated with the same procedures. A significant increase in the average catches of perch was found in the limed lakes, but not in the control lakes. This result confirms earlier reports, although seldom supported with data from controls, on successful recovery after liming.     

Individual variation in the foraging strategies of young yellow perch (Perca flavescens) from the Ottawa River

Differences in the foraging strategies among young individuals of the yellow perch (Perca flavescens) were observed in the laboratory by using two kinds of food (Daphnia and brine shrimp) separately and together. Individuals differed significantly in their ability for feeding attempts, time interval between two consecutive feeding attempts, feeding angles, regurgitation rate, and number of unsuccessful attempts and in their food preference. It is concluded that there are individuals with different foraging strategies. Variability in foraging strategies within single species populations is important because it may explain how the conspecific individuals may differ in their overall feeding behaviour.     

Contrasting migration behaviour of Daphnia pulicaria and D. galeata x hyalina, in avoidance of predation by 0+perch (Perca fluviatilis)

In spring and early summer, a small population of the large-bodied Daphnia pulicaria coexists with a much larger population of the medium-sized hybrid Daphnia galeata × hyalina in the epilimnion of Lake Maarsseveen (The Netherlands). When large shoals of juvenile perch (Perca fluviatilis) appear in the open water, both species start to migrate vertically. Since D. pulicaria has a larger body-size than D. galeata × hyalina, and is therefore competitive dominant over the hybrid, it is unlikely that both species interact via their common food resource, but they react both to fish predation. However, since they differ in size, and therefore in vulnerability for fish predation, both species adopt different strategies. The smaller bodied, and less vulnerable D. galeata × hyalina exhibits diel vertical migration ascending to the surface at dusk, and staying there during the night. In this way, benefiting from the higher temperatures of the surface layers. In contrast, the large-bodied, and more vulnerable D. pulicaria selects the deep cold hypolimnion water layers as refuge against fish predation. In this way it benefits from a safe habitat, free from fish predators, but on the other hand suffers from low water temperatures, which decrease its fitness. It is likely that the relatively higher temperature in the upper water layers serves as a proximate factor for the downward migration of D. pulicaria.     

Ontogenetic changes in the bathypelagic distribution of European perch fry Perca fluviatilis monitored by hydroacoustic methods

The behaviour of bathypelagic perch fry (BPF) was monitored during 21 May, 24 June, 8 August and 5 September 1996 along the longitudinal profile of Římov reservoir (Czech Republic), using acoustic methods (SIMRAD EY 500 echosounder, elliptical split-beam transducer, Sonar5 postprocessing software). In May, the BPF layer comprised a mass of solitary perch larvae at an abundance of over 31,000 ind. ha⁻¹. In contrast, in June, the BPF layer consisted almost exclusively of shoaling individuals. The abundance of BPF increased up to 166,000 ind. ha⁻¹. Similary, in August, fish in shoals prevailed in the BPF layer but the abundance dropped to less than 3,000 ind. ha⁻¹. In September, the BPF layer consisted of both individuals and small groups of fish. At this time the abundance of BPF slightly increased to almost 5,000 ind. ha⁻¹. Individual parameters describing the BPF layer, depth of the main layer, abundance of BPF in the main layer and thickness of the whole layer, differed significantly among months. The negligible abundance of potential predators below the thermocline, both in June and in August (exclusively adult perch), suggests that the BPF shoals were more likely to be created as a result of a social need rather than as a defense against predators. During similar surveys carried out prior to (25 April, 5 May, 7 May, 12 May, 15 May) and after this period (3 October, 7 November, 2 December), the BPF layer was not recorded.     

The predation on Asellus aquaticus (L.) by perch,Perca fluviatilis (L.), in a small forest lake

Asellus aquaticus (L.) was the most important benthic food item for perch, Perca fluviatilis (L.), in a small, extremely humic forest lake in southern Finland. The proportion of A. aquaticus in the diet of perch varied according to the former's availability, which, in turn, depended on its life cycle. Perch 11.0–12.9 cm (total length) most frequently fed on A. aquaticus; smaller and larger perch fed more frequently on crustacean zooplankton and aquatic insect larvae, respectively. The high density of the perch population, the importance of A. aquaticus in the benthos of the study lake and the general high activity level of the prey resulted in a high predation (0.1–1.8% per day). The population of A. aquaticus was also limited by the scarcity of macrophytic vegetation and the small area of oxygenated littoral in the lake.     

Feeding range of age 1+ year Eurasian perch Perca fluviatilis in the Baltic Sea

Using the widespread Eurasian perch Perca fluviatilis as a model organism, feeding ranges were investigated using stable‐isotope ratios (δ¹⁵N and δ¹³C) and body condition. Differences were found between closely located sampling sites in a littoral area without obvious migration barriers, indicating that individual fish had small feeding ranges. Body condition differences between sampled stations were consistent over 4 years. Such sedentary behaviour is important to consider in, e.g. fisheries management and environmental monitoring, as local catch regulations may be meaningful or geographic stability in sampling locations may reduce noise in data.     

Temporal distribution of perch (Perca fluviatilis L.) in a lake-reservoir (Moselle,France): analysis of catches with vertical gill nets

Four fishing periods within a yearly cycle between 1990 and 1991 were carried out in the lake-reservoir of Mirgenbach, lake of the nuclear power station in Cattenom (Moselle, France). The catches of perch (Perca fluviatilis L.) were analyzed.In all, about 630 perch were caught in 8 sampling stations. The analysis of seasonal catches of this carnivorous fish was made according to size and mesh; a simple correspondence factor analysis enabled us to establish that this species lived in different areas depending on the season.     

Ovarian fluid plays an essential role in attachment of Eurasian perch, Perca fluviatilis eggs

Eggs of the Eurasian perch, Perca fluviatilis, are strongly attached to each other to form 0.5-5.5 m long and 1-6 cm wide floating jelly strands or ribbons. The processes involved in the initiation of egg attachment were investigated in the present study. Eggs are released together with small amounts of ovarian fluid. When the eggs come in contact with water, a fibrous jelly-like structure is formed by interactions between proteins of the ovarian fluid and proteins of the zona radiata externa (ZRE) leading to the egg attachment with each other. This mechanism is unique in teleost fish and is described for the first time in the present paper.     

Cascading life-history interactions: alternative density-dependent pathways drive recruitment dynamics in a freshwater fish

Although density-dependent mechanisms in early life-history are important regulators of recruitment in many taxa, consequences of such mechanisms on other life-history stages are poorly understood. To examine interacting and cascading effects of mechanisms acting on different life-history stages, we stocked experimental ponds with fathead minnow (Pimephales promelas) at two different densities. We quantified growth and survival of the stocked fish, the eggs they produced, and the resulting offspring during their first season of life. Per-capita production and survival of eggs were inversely related to density of stocked fish; significant egg cannibalism by stocked minnows resulted in initial young-of-the-year (YOY) densities that were inversely related to adult densities. Subsequent growth and survival of YOY were then inversely related to these initial YOY densities, and survival of YOY was selective for larger fish. Because of these compensatory processes in the egg and YOY stages, treatments did not differ in YOY abundance and mean size at the end of the growing season. Because of differences in the intensity of size-selective mortality, however, variation in end-of season sizes of YOY was strongly (and inversely) related to densities of stocked fish. When mortality was severe in the egg stage (high densities of stocked fish), final YOY size distributions were more variable than when the dominant mortality was size-selective in the YOY stage (low stocked fish densities). These differences in size variation could have subsequent recruitment consequences, as overwinter survival is typically selective for YOY fish larger than a critical threshold size. Density-dependent effects on a given life stage are not independent, but will be influenced by earlier stages; alternative recruitment pathways can result when processes at earlier stages differ in magnitude or selectivity. Appreciation of these cascading effects should enhance our overall understanding of the dynamics of stage-structured populations.     

Smooth muscles in relation to the gill skeleton of Perca fluviatilis: organization and innervation

Summary Two laminae composed of smooth muscles, elastic tissue and collagen have been described in relation with the gill skeleton in Perca fluviatilis. A transverse smoothmuscle lamina joins the base of the cartilage rods of the two opposite hemibranchs. A longitudinal smooth-muscle lamina runs parallel to the afferent branchial artery and joins the cartilage rods from one filament to the other. In both laminae, the formaldehyde-induced fluorescence technique (Falck-Hillarp) reveals a network of nerve fibers displaying a green fluorescence characteristic of catecholamines. At the ultrastructural level, the presence of nerve endings containing clear and granular vesicles, and the degeneration of these endings after 6-hydroxydopamine treatment confirm the aminergic nature and the sympathetic origin of this innervation. Surgical denervation brings evidence that the innervation of both laminae is supplied by the metatrematic branches of the branchial nerves. The role of these smooth-muscle laminae remains speculative.     

Activity and food choice of piscivorous perch (Perca fluviatilis) in a eutrophic shallow lake: a radio-telemetry study

SUMMARY 1. Radio transmitters were implanted in large perch (27–37 cm) in a shallow lake in Denmark. Between 6 and 13 perch were tracked every 3 h for 24‐h periods twice (summer) or once a month (winter) from August 1997 to July 1998. Activity levels were recorded as minimum distance moved per hour. 2. No significant differences in activity levels of individual fish were observed. 3. Highest activities were observed at daytime with peaks at dawn and dusk or midday. This diel pattern was most pronounced from October to April, whereas diel variations were less in the summer months, with no peaks occurring in midsummer. The general lack of activity at night supports the idea that perch is a visually oriented forager. 4. There was no significant relationship between daytime activity during the year and temperature or day length, but nighttime activity was correlated with temperature. In contrast with previous findings, activity levels varied little seasonally, except for high activity levels that occurred concomitantly with high temperatures in August. Instead, we found a significant relationship between the total distances moved per day and temperature, indicating that perch moved at the same average speed in the wintertime, but did so for shorter periods than in summer because of shorter day lengths. 5. Diet of the tagged perch shifted from fish dominance between August and January to invertebrates from February to June. There was no correlation between the diet shift and activity levels, indicating that feeding on invertebrate requires similar activity levels as predation on fish. 6. The results of this telemetry study throughout a year suggest that perch are more active during the winter than previously inferred from gill‐net catches. This observation underscores the importance of perch as a predator of 0+ planktivorous fish in lakes and has potential implications for pelagic food web structure and lake management by biomanipulation.     

Acidification affects the perch,Perca fluviatilis,populations in small lakes,of southern Finland

Synopsis Characteristics of perch were studied in 4 recently acidified lakes in southern Finland. The densities of the populations were 0–250 ha^–1 compared with 1400–3300 ha^–1 in two circumneutral reference lakes. The perch were close to extinction in one acidic lake and dominated by 7 year old fish in two other acidic lakes, suggesting decreased reproduction. Individual fecundity was not reduced by the acidity, but high mortality (close to 100%) of eggs in two acidic lakes indicated recruitment failure. In one of the acidified lakes reproduction had continued resulting in a population dominated by younger fish. However, increased egg mortality, low density, and the resulting high growth rate of perch in this lake were considered to be consequences of the acidity. These findings are the first documentation on the possible decrease of fish populations in Finland due to acid precipitation.     

Competition between perch (Perca fluviatilis) and ruffe (Gymnocephalus cernuus): the advantage of turning night into day

1. The outcome of interspecific competition for food resources depends both on the competitors’ sensory abilities and on environmental conditions. In laboratory experiments we tested the influence of daylight and darkness on feeding behaviour and specific growth rate (SGR) of two species with different sensory abilities. 2. We used perch (Perca fluviatilis) as a visually orientated, and ruffe (Gymnocephalus cernuus) as a mechano‐sensory oriented predator and tested their growth rates and behaviour under conditions of interspecific and intraspecific competition. Three different foraging conditions were used: food supplied (i) only during the day, (ii) only during the night or (iii) during both day and night. 3. In perch neither SGR nor feeding behaviour were influenced substantially by interspecific competition during daylight. During darkness their foraging behaviour changed markedly and their access to the food source as well as their SGR were negatively affected by the presence of ruffe. 4. Ruffe's foraging behaviour did not change during either day or night with interspecific competition. During the night ruffe's SGR was higher with interspecific competition, probably because of a release from intraspecific competition and the competitive inferiority of perch during the night. 5. Because of its seonsory abilities ruffe feeds predominantly at night, thereby reducing competitive interference from perch.     

Physiological correlates of growth and condition in the yellow perch (Perca flavescens)

This study on yellow perch (Perca flavescens) examines a series of enzymatic markers and the relative weights of pyloric caeca and visceral lipids, their response to changes in feeding regime and their potential use to infer recent changes in growth rate and fish condition. Fish were exposed to four different feeding regimes for 12 weeks resulting in specific growth rates ranging from 0.3% to 3.5% (%/day). Growth and condition responded rapidly to changes in ration and the weight of pyloric caeca and visceral lipids reflected increased feed intake. Growth rate was correlated with muscle citrate synthase and caecal nucleoside-diphosphate kinase activities, whereas condition was correlated with muscle citrate synthase and lactate dehydrogenase activities and with caecal glucose-6-phosphate dehydrogenase activity. Results showed that enzyme activities and biometric parameters responded rapidly to increased feed intake, but the response was slower when food intake decreased. Plateaus were attained for both condition and visceral lipid index, but the relative weight of pyloric caeca continued to increase throughout the experimental period. Results from this study could, in principle, be used to infer recent growth and energy status in wild yellow perch and thus provide an indicator of food availability in their environment.