A note on the age composition, growth rate and food of perch Perca fluviatilis (L.) in four eutrophic lakes, England

The perch populations of all lakes sampled are characterised by high growth rates (l± 320–410 mm) and year-class instability. The concurrence of four year-classes (1970,1973,1975and 1976) in three of the lakes is good evidence for climatic control of recruitment. Poor recruitment in Rostherne Mere during 1973 is attributed to the effects of cannibalism. An increase in the growth rate of young fish (0+) in all lakes during 1975 and 1976 is noted. The relatively poor growth of perch in Rostherne Mere is a consequence of the low specific growth rate there during the first year of life. An outbreak of the 'perch-disease' killed most of the adult perch present in 1975 in Rostherne Mere.     

A note on the growth-rate and year-class strength of bream, Abramis brama (L.), in three eutrophic lakes, England

In this study some comparative observations on the growth-rate and year-class strength of bream, Abramis brama , in three Shropshire-Cheshire meres are given. All populations are exposed to variable recruitment. The occurrence of four strong year-classes (1959, 1966, 1969,1973) suggests that climate is an important factor influencing spawning success. Water temperatures ≥16°C in July and August may be particularly important. All other intervening year-classes are absent. The growth-rate of bream in Tatton Mere and Ellesmere Mere is good ( L ∞= 541–561 mm) but comparatively poor ( L ∞= 448 mm) in Cole Mere. The significance of population density, food supply and parasitic infestation by Ligula intestinalis is discussed.     

Consequences of reduced nutrient loading on a lake system in a lowland catchment: deviations from the norm?

1. Lake restoration from eutrophication often rests on a simple paradigm that restriction of phosphorus sources will result in recovery of former relatively clear‐water states. This view has apparently arisen from early successful restorations of deep lakes in catchments of poorly weathered rocks. Lakes in the lowlands, however, particularly shallow ones, have proved less tractable to restoration. This study of three lowland lakes provides insights that illuminate a more complex picture. 2. The lakes lie in a sequence along a single stream in a mixed urban and rural landscape. Severely deoxygenating effluent from an overloaded sewage treatment works was diverted from the catchment in 1991. Effects on two lakes, Little Mere (z_max <2 m) and Rostherne Mere (z_max 31 m) were followed until 2002. Mere Mere (z_max = 8 m), upstream of the former works, acted as a comparison for changes in water chemistry. Mere Mere showed no change in total phosphorus (TP), total inorganic nitrogen, or planktonic chlorophyll a concentrations. Increased winter rainfall was associated with higher winter soluble reactive phosphorus (SRP) and ammonium concentrations in its water. 3. Little Mere changed from a deoxygenated, highly enriched, fishless system, with large populations of Daphnia magna Straus, clear water and about 40% aquatic plant cover, to a slightly less clear system following diversion. Daphnia magna was replaced by D. hyalina Leydig as fish recolonised. Spring peaks of chlorophyll a declined but summer concentrations increased significantly. Annual mean chlorophyll a concentrations thus showed no change. Submerged plants became more abundant (up to 100% cover), with fluctuating community composition from year to year. Summer release of SRP from the sediment was substantial and has not decreased since 1993. The summer phytoplankton was apparently controlled by nitrogen availability perhaps with some influence of zooplankton grazing. SRP was always very abundant. The lake appeared to have reached a quasi‐stable state by 2002. 5. Rostherne Mere showed a steady decline in TP and SRP concentrations following effluent diversion apparently as a result of steady dilution by water with lower phosphorus concentration. Decline in phosphorus concentrations was much less rapid than expected because of internal remobilisation from the hypolimnion and sediments. There have been no changes in chlorophyll a concentration or of nitrogen availability and by 2002 the phytoplankton probably remained limited by a combination of mixing, grazing and nitrogen. 6. A seeming paradox is, thus, that immense changes in phosphorus budgets have shown no consequences for phytoplankton chlorophyll concentrations in either of the lakes, although the seasonal distribution has altered in Little Mere. Although these case studies deviate from others, for both shallow and deep lakes, they represent distinctive situations rather than undermining conventional models.     

The growth of whitefish in relation to water quality and fish species composition

Water quality in 16 Finnish lakes did not affect directly densely gill-rakered whitefish growth, except possibly in an acid (pH 4·9) lake, Iso Lehmaälampi, where acidity may have retarded the growth of whitefish. The density of roach affected whitefish growth in the second year of life: highest growth rates were in lakes without a roach population and lowest growth rates in lakes having strong roach populations. Competition by vendace also retarded the growth of young whitefish. The efficient mass removal of roach from a eutrophic lake was considered to have increased the growth rate of young whitefish. It is suggested that an examination of the fish species composition and relative abundance, as well as the growth of whitefish, can be used as an aid in predicting the success of stocking with whitefish.     

Changes in a deep lake following sewage diversion – a challenge to the orthodoxy of external phosphorus control as a restoration strategy?

1. The restoration of deep lakes has traditionally focused on reducing the external phosphorus loading. 2. Following the diversion of sewage effluent, that led to marked reductions in nutrient concentrations in its main inflow, Rostherne Mere has shown no reduction in phosphorus or chlorophyll a concentrations. A shallow lake upstream (Little Mere), however, has shown a marked response to effluent diversion. 3. Nutrient budgets for Rostherne Mere reveal that sewage effluent was by far the most significant external source of total phosphorus and that diffuse drainage from the catchment was the most significant external source of dissolved inorganic nitrogen. Phosphorus loads from groundwater and a bird roost were insignificant. Internal sources of phosphorus were, however, considerable and were largely responsible for the observed delay in recovery. 4. Phosphorus limitation of phytoplankton biomass may never be attainable because of substantial internal and diffuse sources of phosphorus, combined with a long retention time. Nitrogen is likely to be more important in limiting phytoplankton biomass. Control of diffuse nitrogen sources may therefore be more effective in the restoration of the deeper lakes of this region.     

Trophic dynamics in a whole lake experiment: size-structured interactions and recruitment variation

Horizontal and vertical heterogeneity as a result of size‐structured processes are important factors influencing indirect effects in food webs. In a whole‐lake experiment covering 5 years, we added the intermediate consumer roach (Rutilus rutilus) to two out of four lakes previously inhabited by the omnivorous top predator perch (Perca fluviatilis). We focused our study on the direct consumption effect of roach presence on zooplankton (and indirectly phytoplankton) versus the indirect effect of roach on zooplankton (and phytoplankton) mediated via effects on perch reproductive performance. The patterns in zooplankton and phytoplankton abundances were examined in relation to population density of roach and perch including young‐of‐the‐year (YOY) perch in the light of non‐equilibrium dynamics. The presence of roach resulted in changed seasonal dynamics of zooplankton with generally lower biomasses in May–June and higher biomasses in July–August in roach lakes compared to control lakes. Roach presence affected perch recruitment negatively and densities of YOY perch were on average higher in control lakes than in treatment lakes. In years when perch recruitment did not differ between lakes as a result of experimental addition of perch eggs, total zooplankton biomass was lower in treatment lakes than in control lakes. Phytoplankton biomass showed a tendency to increase in roach lakes compared to control lakes. Within treatment variation in response variables was related to differences in lake morphometry in treatment lakes. Analyses of the trophic dynamics of each lake separately showed strong cascading effects of both roach and YOY perch abundance on zooplankton and phytoplankton dynamics. Consideration of the long transients in the dynamics of top predators (fish) in aquatic systems that are related to their long life span involving ontogenetic niche shifts is essential for making relevant interpretations of experimental perturbations. This conclusion is further reinforced by the circumstance that the intrinsic dynamics of fish populations may in many cases involve high amplitude dynamics with long time lags.     

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.     

The viability of akinetes of blue-green algae recovered from the sediments of Rostherne Mere

Akinetes from species of blue-green algae were recovered from the recent sediments of Rostherne Mere. Gas vacuolate populations of Aphanizomenon and Anabaena were cultured from sediment up to 18 and 64 years old respectively. Akinetes of blue-green algae may not only have a temporary, or overwintering, function but may also ensure long term survival.     

Some mineral nutrient studies of a lowland mere in Cheschire,England

Summary Analyses of nine ionic constituents in Rostherne Mere water were made at regular intervals and five depths over a two year period. Only nitrate-nitrogen showed a regular seasonal pattern. Phosphate-phosphorus and ammonium-nitrogen increased with depth in summer. The results are discussed in relation to the origin of the nutrients, stratification and the phytoplankton of the mere. The levels of nutrients present did not appear to limit algal growth with the possible exception of nitrogen.     

The cause of extreme year class variation in a population of roach, Rutilus rutilus L., from a eutrophic lake in southern England

Investigations into the biology of the roach population at Slapton Ley, Devon, U.K. were carried out between October 1982 and December 1984 and data collected from the lake since 1977 have been re-analysed to assist interpretation. The roach population has been characterized in recent years by extreme year class variation and a tendency for year classes to alternate in strength. The survival of the roach was very poor, and the spawning population therefore consisted largely of 2-and some 3-year-old fish. Consequently, the size of the spawning stock was small and dependent on the strengths of the year classes that it contained. Recruitment to many year classes was probably limited by the size of the spawning stock. The occurrence of a poor year class in the population would therefore set up an autonomous cycle of poor recruitment to alternate year classes in a manner similar to many tropical marine fisheries. Observation of other British roach populations showed that a short life-span may be associated with conditions that promote very rapid growth rates. The growth rate of individual roach at Slapton was found to be one of the fastest in Britain, and this was apparently due to the very high productivity of Slapton Ley together with the persistence of the roach population at very low densities.     

Occurrence of cell-associated mucilage and soluble extracellular polysaccharides in Rostherne Mere and their possible significance

The cell-associated mucilage and soluble extracellular polysaccharides (EPS) were investigated in a eutrophic freshwater lake (Rostherne Mere, Cheshire, U.K.) over up to 2 year annual cycles. Five particular lake algae (Anabaena spiroides Klebahn, Anabaena flos-aquae Brébisson ex Bornet & Flahault,Anabaena circinalis Rabenhorst, Microcystis aeruginosa Kützing emend. Elenkin and Eudorina elegans Ehrenberg) were found to be the major contributors to cell-associated mucilage, particularly M. aeruginosa. Calculation of the total amount of cell-associated mucilage in the phytoplankton samples showed that it occupied 0.0001–0.007% (the latter during a bloom of Microcystis) of lake water volume within the epilimnion. Seasonal changes in the total volume of associated mucilage reflected the succession of mucilage-producing algal species in Rostherne Mere, which was closely correlated with the physico-chemical (temperature, oxygen, pH, nutrients) and biological (Secchi depth, phytoplankton) parameters within the lake. High levels of cell-associated mucilage present in the lake may have potential for binding metals or other ions in the aquatic environment. Colourimetric determination of the concentration of soluble EPS revealed concentrations of between 2.5 and 60 mg l^–1, with peak levels during the bacillariophyceaen bloom and late clear water phase. The second phase did not appear to relate directly to changes in algal population, and may result from bacterial activity, algal lysis or zooplankton activity. As soluble EPS forms a major component of the total amount of dissolved carbon in lakes, the study of the soluble EPS is important to understand the carbon cycle in freshwaters. No direct correlation occurred between algal-associated mucilage and soluble EPS over a single annual cycle.     

The effects of a change in the growth rate of roach, Rutilus rutilus (L.), on the biology of the fish tapeworm Ligula intestinalis (L.)

Investigations into the biology of the roach and the pathogenic tapeworm Ligula intestinalis (L.) populations at Slapton Ley, Devon were carried out between October 1982 and December 1984. Data collected from the lake since 1977 have also been re-analysed to determine how the dramatic improvement in the individual growth rate of the roach over this period may have affected the growth, maturation and life-cycle of Ligula. Only the very young roach at this site become infected, so it was possible to follow cohorts of plerocercoids of similar age through each roach year class. Results for the 1978 and 1983 year classes are presented in detail. The roach grew extremely rapidly from May to August in each year, resulting in a pronounced cyclical pattern of changes in the condition of the roach, with the lowest condition occurring in late winter and spring. The yearly increase in the roach growth rate was accompanied by an increase in plerocercoid growth rate in the 0 + roach, but not in the 1 + roach. The growth rate of the plerocercoids was very high compared to that at other sites. It is usual for the parasite index (PI) of Ligula-infected fish to be high and to increase throughout their first few years of life. At Slapton, however, a lack of multiple infections has prevented high PIs from occurring, and in recent years the growth rate of the roach has been so high that the plerocercoids were unable to maintain a high weight relative to the fish, and the highest PIs occurred in the 0 + roach throughout late winter and spring. In recent year classes, therefore, the maximum PIs and highest pathogenicities coincided with the period of lowest condition in the 0+ roach. Observations of both caged and natural populations of 0+ roach over winter showed that a significant loss of roach containing the larger plerocercoids occurred from the population. In vitro cultivation of Ligula plerocercoids showed that they were capable of maturation at weights of 0.5 g, and only 6 months after having infected the roach. The increase of the growth rate of the plerocercoids in the 0+ roach has therefore resulted in a greater proportion of these plerocercoids being capable of infecting the definitive host. As a result of the increase in individual growth rate of the roach at Slapton, the potential for Ligula transmission, as measured in terms of both their pathogenicity and maturity, has shifted from the 1 + to the 0+ roach.     

Resource limitation during early ontogeny: constraints induced by growth capacity in larval and juvenile fish

The presence of and mechanisms behind density-dependent growth and resource limitation in larval and juvenile stages of organisms with high mortality such as fish are much debated. We compare observed consumption and growth rates with maximum consumption and growth rates to study the extent of resource limitation in young-of-the-year (YOY) roach (Rutilus rutilus) and perch (Perca fluviatilis). Diet, habitat use, consumption rate and growth rate were measured under varying YOY fish densities over 2 years in four lakes. In the first year, YOY roach and perch were studied under allopatric conditions. Experimental addition of perch roe in the second year also allowed study of YOY of the two species under sympatric conditions in two of the lakes. The diet of YOY roach was dominated by cladoceran zooplankton and YOY roach habitat use was restricted to the shore region in both years. This restricted habitat use did not involve any cost in foraging gain in the first year as consumption and growth rates were very close to maximum rates. During the second year, when the two species coexisted, resources were limited in late season, more so in the littoral than in the pelagic habitat in one lake while the reverse was the case in the other lake. The diet of YOY perch was also dominated by zooplankton, and with increasing perch size the proportion of macroinvertebrate prey in the diet increased. After hatching, YOY perch first utilized the pelagic habitat restricting their habitat use to the shore after 1 to several weeks in the pelagic zone. During the larval period, perch were not resource limited whereas juvenile perch were resource limited in both years. The fact that YOY perch were more resource limited than YOY roach was related to the higher handling capacity and lower attack rate of perch relative to roach, rendering perch more prone to resource limitation. Estimates of resource limitation based on consumption rates and growth rates yielded similar results. This supports the adequacy of our approach to measure resource limitation and suggests that this method is useful for studying resource limitation in organisms with indeterminate growth. Our results support the view that density-dependent growth is rare in larval stages. We suggest that density-dependent growth was absent because larval perch and roach were feeding at maximum levels over a wide range of larvae densities. Received: 14 June 1999 / Accepted: 29 October 1999     

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.     

Allopatric and sympatric diversification within roach (Rutilus rutilus) of large pre‐alpine lakes

Intraspecific differentiation in response to divergent natural selection between environments is a common phenomenon in some northern freshwater fishes, especially salmonids and stickleback. Understanding why these taxa diversify and undergo adaptive radiations while most other fish species in the same environments do not, remains an open question. The possibility for intraspecific diversification has rarely been evaluated for most northern freshwater fish species. Here, we assess the potential for intraspecific differentiation between and within lake populations of roach (Rutilus rutilus)—a widespread and abundant cyprinid species—in lakes in which salmonids have evolved endemic adaptive radiations. Based on more than 3,000 polymorphic RADseq markers, we detected low but significant genetic differentiation between roach populations of two ultraoligotrophic lakes and between these and populations from other lakes. This, together with differentiation in head morphology and stable isotope signatures, suggests evolutionary and ecological differentiation among some of our studied populations. Next, we tested for intralacustrine diversification of roach within Lake Brienz, the most pristine lake surveyed in this study. We found significant phenotypic evidence for ecological intralacustrine differentiation between roach caught over a muddy substrate and those caught over a rocky substrate. However, evidence for intralacustrine genetic differentiation is at best subtle and phenotypic changes may therefore be mostly plastic. Overall, our findings suggest roach can differ between ecologically distinct lakes, but the extent of intralacustrine ecological differentiation is weak, which contrasts with the strong differentiation among endemic species of whitefish in the same lakes.     

Environmental correlates of population variables of perch (<Emphasis Type="Italic">Perca fluviatilis</Emphasis>) in boreal lakes

We examined relationships among perch population variable parameters in two types of lakes, lakes with perch (P-lakes, n?=?15) and lakes with perch and roach (PR-lakes, n?=?10) using redundancy analysis (RDA) to relate population variables to environmental factors. Effects from environmental factors were tested for significance by means of permutation tests (Monte Carlo). Three factors, pH, altitude and fraction of roach (by number) in the gill net catches, explaining 47.9% of the variation, had significant effects on perch population variables. The significance of pH was improved by partialing out the effect of conductivity and roach. Similarly, the significance of altitude was improved by partialing out the effect of pH and roach, and the significance of roach was improved by partialing out the effect of pH and altitude. When the fraction of pike in the catch was included in the analysis, the effect of roach was not significant and vice-versa, as roach and pike fractions were correlated with each other. The effect of pike was significant when roach was not included, but the effect was not as strong as the effect of roach. A biplot was constructed by plotting population variables on the first and second RDA axis, with arrows showing five selected environmental factors. Growth of 3+ to 5+ perch was positively related to pH and altitude, perch catch per unit effort was negatively related to pH and altitude, and age specific perch weight was negatively related to fraction of roach. The relationship between growth of 2+ perch and pH was not as strong as the relationship between pH and the growth of older perch. Moreover, the growth of 2+ perch was negatively related to the fraction of roach, probably due to competition between young zooplankton feeding perch and roach.     

Seasonal variations in the vertical distribution and buoyancy of Microcystis aeruginosa Kütz. emend. Elenkin in Rostherne Mere,England

Distribution and buoyancy studies in Rostherne Mere (depth: 30m) show that the life-cycle of Microcystis is intimately related to the cycle of thermal stratification: colonies overwinter on the bottom muds, but migrate to the epilimnion in summer, returning again in late autumn or winter.     

How many fish populations in Finland are affected by acid precipitation?

Synopsis The number of fish populations affected or lost from small lakes in southern and central Finland due to acid precipitation is estimated. Tolerance limits (pH and labile aluminum) of common fish species were obtained from a fish status survey of 80 lakes. These tolerance values were used to estimate the proportion of affected lakes from the water chemistry data of 783 statistically selected lakes. The proportion of anthropogenically acidified lakes was estimated by calculating pre-acidification pH and aluminum concentrations of the lakes, using a steady-state model based on water chemistry. The number of fish populations for which acidification has affected growth or population structure was estimated at between 2200 and 4400. Out of these, the number of fish populations that have disappeared due to acid precipitation would be about 1000–2000. Almost 60% of the affected or lost populations are roach, Rutilus rutilus, the most sensitive of the common fish species in small lakes in southern and central Finland. Less than 15% of the damaged population is European perch, Perca fluviatilis, the most common species. This is due to the substantially higher tolerance of perch to acidified water in comparison with roach.     

Can migrants escape from density dependence?

Migration is thought to maximize growth by enabling individuals to escape from density dependence, but this has rarely been tested at the individual level in natural populations. We employed linear mixed modeling of the spacing between consecutive scale growth rings to reconstruct individual growth profiles of a paradigmatic fish migrant, the sea trout (Salmo trutta) and related these to estimates of year class strength over a 13‐year period. Variation in scale growth was 1.3 times greater among individuals than within individuals in freshwater and 10 times greater at sea. Scale growth was inversely related to year class strength, both in freshwater (before migration) and at sea (after migration). Competition for patchily distributed resources is the most plausible explanation of the negative density‐dependent growth observed in freshwater and, to a lesser extent, in the marine environment. Our study provides some of the strongest evidence for a role of density dependence in determining partial migrations because although migrants can maximize growth by moving into the sea, they do not appear to become free from density dependence constraints completely. This has implications for conservation and suggests that sea trout and other anadromous fish displaying partial migrations may not be best managed on a river by river basis, but rather from a broader, coastal perspective.     

Distribution and variation in the growth of roach, Rutilus rutilus (L.), and dace, Leuciscus leuciscus (L.), in a river catchment in south-west England

The distribution and growth of roach and dace in the R. Exe catchment was studied. Distribution conformed to the classic theories of zonation, although human interference extended or reduced the range of species. Distinct differences were found in the growth rates of roach and dace between different regions of the catchment. Both species achieved their fastest growth in zones where the river topography portrayed their 'preferred' habitat characteristics, i.e. the fast-flowing middle and slow-flowing lower reaches for dace and roach, respectively. Female roach were found to grow considerably faster than male fish, whilst no sexual differentiation in growth rates was found in dace. Considerable fluctuations in year class strength were observed in both species.