Population responses of perch (Perca fluviatilis) and roach (Rutilus rutilus) to recovery from acidification in small Finnish lakes

Decreasing trends in atmospheric emissions and acidic deposition during the 1990s have resulted in chemical recovery from acidification in the sensitive surface water systems of southern Finland. Responses of perch and roach populations to the improved water quality were studied in 30 small lakes with the aid of water chemistry monitoring data gathered in 1987–2002 and the data collected from two consecutive periods of gillnet test fishing, 1985–1988 and 2001–2002. In the most acidified lakes, alkalinity and ANC have increased and sulphate and labile aluminium concentrations decreasæed markedly. The response of perch populations to the improved water quality is seen in improved reproduction success, indicated by a higher CPUE in numbers and a lower mean weight. The growth rate of perch has declined as the population density has increased. Roach populations have not recovered in the same way as perch, there being no major changes in NPUE or mean weight. Lower growth rates were, however, observed in the roach populations of all study lakes. The increased perch population density as an obstacle to the recovery of roach populations is discussed. Despite the chemical and biological recovery of the study lakes, the buffer capacity of many headwater lakes is low and the lakes will be sensitive to any increases in acidic deposition in the future.     

Delayed spawning of perch, Perca fluviatilis L., in acidified lakes

The timing of spawning of perch was examined in four acidified lakes (pH 4.4–4.8) and in one circum-neutral lake (pH 6.3) in southern Finland in spring 1987. In three of the lakes, perch started to spawn soon after the ice melt (4–14 degree days > 5° C) and had spawned by the end of May at about 100 degree days > 5° C. In the two most acidified lakes, fish started to spawn later, at 35 and 60 degree days > 5° C, and had spawned in early June, at about 200 degree days > 5° C. The maturing of gonad products was delayed in both males and females.     

Temperature effects on spawning and egg development in Eurasian perch

Spawning, egg strand development and embryo survival was studied during 1995–1996 in perch Perca fluviatilis exposed to heated effluent from two coastal nuclear power plants on the Baltic. Spawning was earlier and more prolonged in the heated areas. Although the fertilization rate was high, most egg strands sampled in the heated areas did not develop to hatching. The egg strands disintegrated within a few days of spawning, and egg mortality was very high. All samples collected from reference areas developed up to hatching. Exposure to high temperatures during the final maturation of the gonad influenced the follicular mechanism producing the jelly membrane which constitutes the matrix of the egg strand. Effects were observed in fish in open effluent areas as well as in an enclosed research basin. These results suggest that there is a conflict between optimal reproduction and temperature preference in temperate area fish. Fish are attracted to cooling water effluent, at the expense of reduced reproductive performance.     

Long-term studies on the Windermere populations of perch (Perca fluviatilis), pike (Esox lucius) and Arctic charr (Salvelinus alpinus)

SUMMARY. 1, Windermere is a glacial valley lake with two basins separated by shallows and a total area of 14.8 km². Perch, pike and charr are the dominant fish species and have been studied on a continuous basis for over 45 years.
2. The initial slow-growing stocks of numerous small perch and moderate numbers of large pike were subjected to a heavy initial fishing pressure that was subsequently relaxed for perch but not for larger (>550 mm) pike. The number of charr in the lake has increased 6–8-fold during the study, probably due to the sustained reduction in the abundance of the larger pike that heavily predate the autumn spawning race of charr when it enters the shallow littoral areas to spawn.
3. The biomass of perch did not recover from the initial fishery until the late 1950s when strong year-classes occurred in warm summers. However, it was now composed of larger and faster growing individuals. In 1976, disease killed over 98% of all adult perch in the lake and the subsequent recovery has again been slow.
4. Existing models have explained the variation in perch year-class strength for the years 1959–74 in terms of adult perch biomass, pike cohort strength and temperature in the year of hatch. A new. more generally applicable model, incorporating a stock-recruitment expression, gives a better fit to the earlier data, particularly from the North Basin, where there have been periods of extremely low adult biomass.
5. The importance of continuing to extend these long-term data sets and the scope for future modelling work are discussed.     

The population dynamics of pike,Esox lucius,and perch,Perca fluviatilis,in a simple predator-prey system

Synopsis The population dynamics and predator-prey relationship of pike, Esox lucius, and perch, Perca fluviatilis, were examined in simple fish communities in two adjacent shallow lakes, Lochs Kinord and Davan, Deeside, Scotland. Few perch survive to age 3 but Z is low for fish > 3 years and perch live up to 17 years. Population fecundity remained relatively high and constant in perch because of the multi-age spawning stock and the presence of older more fecund perch. Growth rates of perch in both lochs are relatively high as a consequence of low stock abundance. The N, B, and P of adult perch were unusually low. The age range of pike, and N, B, P, and growth were in the range of values reported elsewhere. There was little variation in the strength of pike year classes and the importance of cannibalism and low occurrence of alternative prey in the lochs suggest that the populations were self-regulating. Cannibalism by adults was responsible for most of mortality in perch larvae, and predation by pike and adult perch was responsible for the majority of juvenile losses. This conclusion is supported by the high biomass ratios of pike:juvenile perch of 1.0–30.8. While the number of adult fish was almost equal, the biomass of adult pike was 2–3 × that of perch in Kinord and 6 × in Davan. In L. Kinord, where year class strength was stable, high predation pressure from perch and pike reduced perch abundance rather than eliminated year classes. Perch year classes fluctuated in abundance in L. Davan and were eliminated in the first summer in two sampling years. The pike, and particularly the perch populations, have features characteristic of fish communities in unperturbed ecosystems: namely, a wide range of age classes, stability in biomass with variation dampened by longevity, and low production.     

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.     

Ergasilid copepods as parasites of perch Perca fluviatilis and roach Rutilus rutilus in Central Finland: seasonality, maturity and environmental influence

A total of 1255 roach Rutilus rutilus (L.) and 866 perch Perca fluviatilis (L.) from four interconnected lakes in Central Finland differing in trophic status and pollution level were studied for parasitic ergasilid copepods between August 1985 and December 1988. In addition, 109 whitefish ( Coregonus sp.) were studied from one of the lakes.
Four ergasilid species were found: (the prevalence and intensity/fish, respectively, for the whole material are given in parentheses) Ergasilus briani (16·9%, 0·5), Neoergasilus japonicus (15·6%, 0·4) and Paraergasilus longidigitus (2·1%, 0·02) on the roach and Ergasilus sieboldi (9·9%, 0·1) and P. longidigitus (4·9%, 0·05) on the perch. Logit analysis was used to assess the significance of the year, the lake and the season for the prevalence of infection. Generally, the prevalence of E. briani on roach depended on the season only, with higher prevalences in summer. That of E. sieboldi on perch was mainly dependent on the lake, being most frequent in the oligotrophic lake. Both lake, year and season affected the prevalence of N. japonicus in roach. The more complicated system for N. japonicus was explained by the long duration of its free-living stage, its loose attachment and different location on the host, being found on the fins whereas E. briani and E. sieboldi were on the gills.
All species were found throughout the year, and age had no clear influence on infection. The occurrence of egg-sacs and length distributions of the copepods suggest that there are two generations a year, e.g. an overwintering E. briani generation which produces offspring in June to attack the fish in July, and a second generation which attacks the fish from August onwards. Whitefish were not infected by these parasites, only one fish being found which carried a single individual of E. sieboldi.     

Loss of genetic variability in reintroduced roach (Rutilus rutilus) populations

Genetic variation at five microsatellite loci and in mtDNA was surveyed in reintroduced and 'control' populations of roach ( Rutilus rutilus ) in Sweden. Microsatellite allelic richness and allele size ranges were significantly reduced in reintroduced populations, and mtDNA diversity was nearly significantly reduced in reintroduced populations. These measures of genetic variability were strongly correlated with lake characteristics that influence population size and food availability.     

Predator-induced respiratory responses in juveniles of vendace Coregonus albula, whitefish C. lavaretus, perch Perca fluviatilis and roach Rutilus rutilus

Predator-induced respiratory responses of juvenile vendace, whitefish, perch and roach were studied in an intermittent-flow respirometer by creating visual contact between test fishes and predators (juvenile northern pike). Vendace and whitefish always responded to the presence of pike with increased oxygen consumption. The response of perch was the opposite: oxygen consumption decreased to near the standard respiratory level. Roach responded more variably: some individuals increased their oxygen uptake while others decreased it. Changes in the rate of respiration during exposure to predation are supposed to be caused by the differences in locomotory activity due to induced antipredator behaviour. According to their responses, these species could be grouped into escapers (coregonids), hiders (perch) or indeterminates (roach). The species-specific differences in the antipredator behaviour originate in morphological and physiological characteristics which together determine the effectiveness of a particular antipredator style (i.e. either hiding or escaping).     

Differences in growth of perch (Perca fluviatilis L.) in two small forest lakes

Growth patterns and food composition of perch, Perca fluviatilis L., was studied in two small forest lake populations in southern Finland. Size and morphometry of the lakes and physical and chemical properties of water are similar. There is a clear difference in the growth rates of perch between the two lakes. The difference in growth is highly significant in all age groups. In the first lake there is a perch population of 2 000 (1750 ind · ha^–1) adult fishes. In the second lake there is a small population of pike, that keeps the perch population down: 200 adult perch (530 ind · ha^-1). The main food items of perch are crustacean zooplankton, Asellus aquaticus L. and Trichoptera larvae in the first lake and zooplankton, Odonata larvae, Ephemeroptera larvae and Heteroptera in the second.It is concluded that the main reason for the growth difference of studied perch populations is the different population density. There are also differences in species composition of bottom fauna of the lakes, maybe owing to the floating Sphagnum peat moss vegetation in the second lake. This can also affect the growth difference between the two populations of perch.     

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.     

Genetic characterization of 18 novel microsatellite loci in northern pike (Esox lucius L.)

The northern pike (Esox lucius L.), an important predatory freshwater species, is undergoing significant population decline. In this study, 18 novel polymorphic microsatellite loci were isolated and used for assessing genetic variation in the Chinese Ulungur and Hungarian Balaton populations of the species. The number of alleles ranged from 2 to 13, observed heterozygosity from 0.154 to 0.920 and expected heterozygosity from 0.145 to 0.921, thereby indicating the specific usefulness of these suites of markers for investigating genetic variability.     

Reproductive disturbance in a roach (Rutilus rutilus) population affected by cooling water discharge

To analyse a recruitment failure in a roach ( Rutilus rutilus ) population exposed to cooling water from a Swedish nuclear power plant, histopathological examinations were made on female gonads. Several disorders were documented, corresponding to effects previously seen in Lithuanian, Moldavian and Russian heated waters, The relative gonad size (GSI) was generally lower in the exposed population. Oocyte degeneration was observed in about 50% of the females, potentially seriously reducing their reproductive capacity. The gameto- and gonadogeneses also were clearly affected and generally occurred at a faster rate. Sexual maturity was earlier and was accompanied by an increased occurrence of asynchionic gonad and oocyte growth. A normal gonad development should pass through different stages, coupled to the seasons, and the difference between individuals in a population should be small. However, in some exposed fish, the sequence of seasonal development was disturbed and a simultaneous, intensive, growth of sex cells in different stages of development could be seen. An increase of variation in gonad development both within, as well as between, fish thus was documented. Conclusively, the histopathological analysis provided evidence that the observed recruitment deficit could be related to gonad failure.     

Consumption of cyanobacteria by roach (Rutilus rutilus): useful or harmful to the fish?

1. The ability of roach to use cyanobacterial food is generally believed to be one reason for the dominance of roach over perch in eutrophic European lakes. The aim of this study was to test whether cyanobacteria really are a suitable food for juvenile roach. Special attention was paid to differences between the two cyanobacteria species Aphanizomenon and Microcystis which are common in eutrophic lakes and are ingested by roach there.
2. We performed growth and behaviour experiments with juvenile roach fed with zooplankton and the different cyanobacteria. Growth rate with Aphanizomenon was lower than with Daphnia but significantly higher than without food, whereas growth rate with Microcystis was as low as without food.
3. In cultivation experiments of roach faeces, Microcystis was found not to have been digested and grew exponentially after passing through the gut whereas Aphanizomenon stayed at low biomass. Differences in growth were not related to the toxin content of cyanobacteria. Investigations of roach motility showed no differences whether fed with Aphanizomenon or Microcystis .
4. In contrast to Microcystis , Aphanizomenon can be regarded as a suitable food source for juvenile roach probably because of its better digestability. We conclude that the ability to feed on cyanobacteria is not a general competitive advantage for roach, but the outcome depends on the species composition of the cyanobacteria.     

Myxosporea parasites in roach, Rutilus rutilus (Linnaeus), from four lakes in central Finland

Ten myxosporean species belonging to three families were found in roach, Rutilus rutilus (Linnaeus), obtained in 1985 and 1986 from four lakes in central Finland which are connected to each other, but differ in water quality. One of the lakes is polluted by paper and pulp mill effluent, two are eutrophic and one is oligotrophic and still in its natural state. Eight species were found in all the lakes. The most common species were Myxidium rhodei Léger, 1905, Myxobolus muelleri Bütschli, 1882 and Myxobolus pseudodispar Gorbunova, 1936 with prevalences varying between 66–80, 16–31 and 32–59%, respectively, in the four lakes. The largest difference in myxosporean prevalence between lakes was found in the case of M. pseudodispar infection, which was highly significantly lower in the polluted lake. The locations of the myxosporean species in the tissues of the fish were found to be species-specific. M. rhodei and M. muelleri being prevalent in the kidney and M. pseudodispar in the muscles.
No clear seasonal variation was found but a tendency for a decrease in infection with increasing age was recorded in the case of M. rhodei and M. pseudodispar .     

河鲈胚胎及卵黄囊期仔鱼发育

为探究河鲈(Perca fluviatilis)早期生活史和发育生物学,采用体视解剖镜、显微镜仔细观察、测量、描述、绘图的方法,连续观察了6个批次河鲈胚胎及卵黄囊期仔鱼发育状况,进行比较分析。结果显示:(1)在水温8~13℃时,胚胎期约需265h,有效积温2540~2880℃.h;水温11~13℃时,卵黄囊期约需6d,有效积温1750~2120℃.h;(2)辐射状次级卵膜将受精卵连成长带形单层网片状,每个胚胎周围有6个胚胎,排列很有规则。胚胎卵黄囊表面有一个大圆形油球。出膜前期可见眼球色素、胸鳍突起;(3)胚胎出膜的不同步主要是由于出膜前期长短不一和孵化水温较低所致。     

Changes in feeding niche widths of perch and roach following biomanipulation, revealed by stable isotope analysis

1. We made an empirical test of a recent proposal that feeding niche widths might be determined as variance of stable isotope values. We determined δ ¹³C and δ ¹⁵N values of perch ( Perca fluviatilis ), roach ( Rutilus rutilus ) and their prey from a biomanipulated lake, when the mass removal of fish led to reduced inter- and intra-specific competition and increases in zooplankton abundance and body size.
2. After the first fish removals, both perch and roach mean δ ¹³C values decreased and mean δ ¹⁵N values increased, indicating a greater diet contribution from pelagic sources.
3. Variances of both δ ¹³C and δ ¹⁵N values first increased in both fish populations, indicating a wider food spectrum and expanded feeding niche width following reduced fish abundances. Observed changes were greater for the perch population than for roach.
4. In 2006, the perch population abruptly changed its diet so that most individuals were primarily consuming the abundant young-of-the-year fish, and this was reflected in significantly reduced variances of both δ ¹³C and δ ¹⁵N values.
5. We conclude that isotopic variance can indeed reflect changes in feeding niche width and offers a promising way to study such general ecological concepts.     

Metabolic rate and cost of growth in juvenile pike (Esox lucius L.) and perch (Perca fluviatilis L.): the use of energy budgets as indicators of environmental change

Summary Energy budgets of juvenile pike and perch (weighing approximately 3 g) were determined in experiments lasting up to 4 days, by simultaneously measuring oxygen consumption, food consumption, and growth of individual fish. Although the basic pattern of energy allocation was identical in the two species, perch subjected to constant light (PEL) showed faster growth, higher assimilation and conversion efficiency, and higher oxygen consumption than perch subjected to a short daylength regime (PED). The efficiency with which food energy was converted into body mass was 39±5% in PED but 49±4% in PEL. However, the “work coefficient” (increment of body mass/post-prandial increase of oxygen consumption: mg · μmol O _inf2 ^sup−1 ) differed only insignificantly between the two groups of perch, indicating that the metabolic cost of growth was unaffected by the manipulation of experimental conditions. This identifies the higher assimilation efficiency, i.e. the increased flow of food energy into the tissues as being the cause of accelerated growth of perch under the constant-light regime. In both species the maximum feeding-induced metabolic rate was 4 times higher than the lowest preprandial rate. In perch (which were kept on low rations before the experiments) the post-prandial metabolic rate increased steadily from day to day during the 4-day experiments, so that on the last day the rate of oxygen consumption exceeded the rate on the first day by about 41%. This investigation provides further evidence that the allocation of metabolic energy in fish is based on a flexible strategy which responds sensitively to changes in both internal and external conditions.     

The influence of simulated aquatic macrophytes on the zooplankton consumption rate of juvenile roach, Rutilus rutilus, rudd, Scardinius erythrophthalmus, and perch, Perca fluviatilis

Laboratory experiments showed that, of zooplanktivorous roach, rudd and perch, in the absence of any environmental structure roach were the most efficient feeders, but high densities of simulated submerged marcrophytes elevated perch to this position; rudd matched the performances of perch and roach only at intermediate structure densities. These changes in efficiency rankings with increased structure were the result of overall decreases in the performances of roach and rudd. Simulated water lilies and emergent reeds had their own influences on consumption rate. These effects were largely mediated through relationships between structure density and swimming speed; roach and rudd, not perch, showed reduced swimming speeds in the more structured environments. Observations of the routine behaviour of the fish suggest that their assessment of predation risk may be an important factor in determining their activity level and hence foraging strategy.