Changes in the rate of growth in a stunted roach Rutilu rutilus population

Data is presented concerning the growth of roach in a South Lancashire lake. Results from a sample of roach in May 1969 indicate the slowest growth rate for which published records are available. A comparison of the length/age and weight/age relationships between data for May 1969 and May/June 1971 yields significant between-year differences indicating an increase in the growth rate during that period. A study of the growth history of individual year classes using back-calculated values from operculum measurements suggests that for both sexes the observed increase in the rate of growth commenced after the 1965–67 period during which very weak year classes were produced. The mean length of females appears to have been larger than that of males from the age of four onwards but the 95% confidence intervals are almost totally overlapping for each pair of values and a statistical comparison of growth rates between the sexes indicates no significant differences. Data for the sexes combined is compared with published results from other British and Continental waters.     

Comparative observations on the growth rate and year class strength of roach Rutilus rutilus L. in two Cheshire lakes, England

In this study some comparative data on the growth rate and year class composition of roach in two Cheshire meres (Rostherne and Tatton) are given. The populations are characterised by year class instability and high growth rates, both of which are most extreme in Tatton Mere. The growth of roach in Tatton Mere is the highest yet recorded from Britain.
The concurrence of two (1969, 1973) strong year classes in both lakes, suggests that climate is an important factor which conditions year class strength. However, although there is some correlation between year class strength and the water temperature index (>14° C) the ultimate cause of year class variability remains obscure. There is no evidence to suggest that year class failure is due to parasitic infestation by Ligula intestinalis or to predation by perch upon roach fry. Year class failure may be due to the mortality of eggs related to the time of spawning. The observed difference in growth rate between the two populations may be attributed to a difference in density caused by the disparity of year class representation in Rostherne Mere and Tatton Mere. However, despite these differences, the net biomass achieved (36–39 kg ha⁻¹) may be similar in both lakes. Superimposed on the effects of density an influence of temperature is noted.     

The occurrence and natural alleviation of stunting in a population of roach, Rutilus rutilus (L.)

Two sudden and pronounced changes in the growth rate of roach, Rutilus rutilus (L.), in Slapton Ley, Devon, have been observed between 1972 and 1976. At first the growth rate declined to the point at which the population became stunted, then subsequently it improved again. These changes were correlated with population density; the numbers of roach were increasing during the late 1960's/early 1970's, and this increase was substantially reinforced by very strong year classes in 1972, and, to a lesser extent, in 1973. The increase in population density resulting from the strong 1972 year class was apparently adequate to accelerate the onset of the stunting observed. The improvement in growth rate in 1976 followed immediately after extensive mortalities amongst the roach during 1975, caused by an epidemic of the parasite Ligula intestinal is (L.). Some improvement in the growth of O group roach had been evident during 1975, but this was not observed in the population. Ford-Walford plots showed a decline in the value of L after the onset of stunting in 1972.
Differences in the reproductive biology of the fish were found following the alleviation of stunting; the gonads of both sexes achieved a greater relative weight, and correspondingly the fecundity of the females was higher, although this latter effect became progressively less marked with increasing size. It was not apparent amongst the largest individuals. There was also some evidence of a decline in the proportion of fish maturing at minimum size. The changes in the reproductive biology were associated with the improvement in conditions for individual fish following the decline in population density.     

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.     

A contribution to the knowledge of the stunted growth of perch (Perca fluviatilis L.) in Holland

Summary The stunted growth of the Dutch perch was studied. To calculate the age and former growth the winterrings of the scales were used. The excellent growth of perch in some parts of Holland in contrast to the bad growth reported by foreign authors is discussed. It is pointed out that the latter growth may be ascribed to the existence of false winterrings, from which one can get rid by dipping the scales in a HCl-solution.It appeared that the initial growth of the stunted perch up to about 13 cm is good. The stunted growth afterwards must be ascribed to a shortage of prey-fish. Especially the smelt seems to be very important. As, however, in Holland young roach are abundant it seems incomprehensible why these cannot serve for suitable perch-food. To investigate this problem the conduct of hunting perch was studied in an aquarium.The results of the aquarium-experiments were in agreement with the change in food-diet at a certain length discussed by other authors; also with the assumption that middle-sized perch do not prey upon smaller congeners.Concerning the preying upon roach it became clear that a single perch will scarcely or not succeed in seizing a roach. The hunting is accomplished by cooperation in open water stretches. Amidst plants perch does not hunt. Hence the stunted growth of perch in Holland must be ascribed to the many plants in Dutch waters, between which the roach may easily escape hunting by perch.A short comparison of prey-sizing by wels, pike pike-perch and perch is made.     

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.     

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     

Ecological changes in a lake fishery and their effects on a stunted roach Rutilus rutilus population

Data for two sampling years (1969/70 and 1970/71) is presented to illustrate the changing composition, density and biomass of the macroinvertebrate fauna in a South Lancashire lake. Six faunal groupings are recognised for a non-parametric analysis of the data and significance levels are given for the between-station and between-year differences recorded. The mean density of macroinvertebrates (1.2 mm mesh sieve) estimated for the lake as a whole increased by 28.5% in the second year (2519 m⁻²-3237 m⁻²) whereas the estimated mean biomass in g m⁻² increased by 86% wet weight (8.61−16.01) and 196% dry weight (1.74−5.15). Molluscs contributed substantially to the increased standing crop which was accompanied by increased development of macrophytes.
A comparative study of the diets of roach in two size groups in winter and summer is described. The variety of food items consumed was found to be greater than in a preliminary study in May 1969 and, for the roach sampled, the average quantity of food consumed per fish increased after the 1969/70 winter. The possible relationships between invertebrate standing crop, roach diet and previously reported changes in the growth of the roach are discussed and an explanation is offered concerning the sequence of biological changes observed in the lake.     

Relationship between scale growth checks, circuli formation rate and somatic growth in Rutilus rutilus (L.) a fish farm-reared cyprinid

Check formation on scales of roach Rutilus rutilus was examined during their production cycle at a fish farm in England through analysis of circuli patterns. Regular check formation was associated with the movement of fish from one type of grow-out facility to another; this resulted in a sudden shift in growth rate and the formation of a new check. As these had the characteristics of annual marks, their formation potentially invalidates their use as structures to determine the age of individuals during recapture events that may follow their introduction to the wild. At low growth rates, the number of circuli was constant throughout the year; this situation changed when fast growth rates were achieved. Five methods of backcalculation were also validated. When the proportionality between the body length and scale radii was weak, backcalculation methods were poor in determining length at check formation.     

Aspects of the ecology of bream, Abramis brama (L.), in a gravel pit lake and the effects of reducing the population density

The growth of bream, Abrumis brumu (L.), in a gravel pit lake was considered to be good in 1982/1983 (L_x=663cm, K=0.105). Theadult breampopulationwasdominated by the 1973and 1976 year-classes. These year classes still formed most of the spawning population in 1987, but had grown very little (<2 cm) in the previous 5 years. In November 1987, 158 kg ha⁻¹ of bream were removed from this lake and in I988 the remaining bream population showed an increase in growth (mean length by 2.7 cm and mean weight by 0.5 kg) and condition (1.87 in 1987 to 2.09 in 1988) in one growing season. This additional growth of an ageing stunted bream population is considered to be the result of an increased food supply after reducing the fish density.     

Reproductive and growth strategies of the red roach, Rutilus arcasii (Steindachner, 1866), in two contrasting tributaries of the River Duero, Spain

The life-histories of red roach were assessed in the R. Moros and the R. Ucero, two contrasting tributaries of the R. Duero. Growth occurred from April to October and Ucero red roach were larger than those in the Moros from their second year onwards. Condition increased with gonad development and abruptly decreased after spawning in both populations. The condition of Ucero fish recovered during the summer, but that of the Moros fish remained low. The same year classes (1980 and 1982) dominated both populations. River Moros red roach spawned in a 4-week period between the end of April and early June, while the spawning season of Ucero red roach lasted until August. The length to fecundity relationship was similar for both populations, and fecundity and ovum size increased with fish length. The life-history traits of these red roach populations are discussed in relation to current life-history theories.     

Observations on the age, growth, reproduction and food of the roach Rutilus rutilus (L.) in two rivers in southern England

Scales and opercular bones from 632 roach from the River Stour were used for age and back-calculated growth determinations. The scales had clearer inner annuli but operculars clearer outer annuli in fish more than nine years old. The annuli were laid in late May or early June at the beginning of the growth period. Growth was minimal between November and April. Roach from both rivers grow faster than those in most other European waters. Female roach grow faster than males; River Frome roach faster than those from the Stour. Spawning occurred in May and elaboration of gonads between September and May. Immature roach have an annual cycle in condition with a maximum in June and a minimum in early Spring. The condition of mature females is affected by the gonad cycle. The fecundity of Stour roach is represented by the formula: log egg number=4.43 log length (mm)—1.69. Approximately half of the Stour males attained sexual maturity at age III and most of the rest by age IV. Half of the females were mature at age IV and the remainder by age V. Both brood success and growth rate varied from year to year but independently of one another. Most Stour roach ate aquatic insect larvae and molluscs but algae were more frequent in the diet of larger fish.     

Size-dependent attack rate and handling capacity: inter-cohort competition in a zooplanktivorous fish

Two size‐dependent processes, metabolic requirements and foraging capacity, heavily influence the competitive ability of organisms. We studied the size‐dependent competitive ability of roach (Rutilus rutilus) in a laboratory experiment to determine the attack rate of roach as a function of roach and zooplankton sizes. The estimated size‐dependent attack rates, size‐dependent metabolic demands and handling capacities were subsequently used to interpret the outcome of a competition experiment between two size classes of roach. Furthermore, size‐dependent attack rates were implemented in an optimal foraging model to predict consumption rates and zooplankton selection to reveal the mechanisms behind competition.
The attack rate first increased with roach size to a maximum around 160 mm to thereafter decrease. Based on this result, we predicted that, small (150 mm) roach had a double advantage in competition for zooplankton in the pond experiment due to their higher attack rate and their lower metabolism compared to large (230 mm) roach. As expected, small roach depressed total zooplankton biomass to a higher extent than large roach, included more zooplankton in their diet and consumed smaller zooplankton. Predicted smallest size class of zooplankton in the diet was close to the observed. Also as expected, large roach fed more on the benthic resource and depressed the benthic resource to a larger extent than small roach. Large roach affected large zooplankton to a higher extent during the first part of the experiment, which could be related to their overall higher handling capacity. The higher impact of large roach on large zooplankton during the first part of the experiment, in turn, resulted in a lower estimated mass intake of zooplankton by small roach in the mixed treatment compared to small roach only treatments. Both small and large roach had a lower growth rate in the mixed treatment compared to single size class treatments. We relate the lower growth rate of small roach in the mixed treatment to large roach's higher efficiency on benthic resources and on large zooplankton during the first part of the experiment. In correspondence with diet data, large roach preferred the shore area of the pond with more benthic invertebrates and were also found more often close to the bottom. Although our results are explainable by exploitative interactions, the fact that the presence of large roach affected the feeding position of small roach points to that social interactions were also involved. Overall, our study implies that a mechanistic understanding is crucial for the interpretation of competition experiments, especially in systems with size‐structured interactions.     

Interspecific dominance amongst three species of coarse fish in Slapton Ley, Devon

Marked changes in the relative numbers of roach Rutilus rulilus (L.), rudd Scardinius erylhroplhalmus (L.) and perch Perca fIuviatilis L. , in Slapton Ley, Devon, have been observed over the period 1967–1978. Historically, the lake had been dominated by rudd and perch; significant numbers of roach were not thought to be present until 1967. From then onwards the roach population exhibited a considerable expansion, apparently replacing the rudd, which had virtually disappeared by 1974. In 1975 extensive mortalities of roach were caused by the pseudophyllidean cestode Ligula intestinalis L. During 1976 and 1977 larger number of rudd, belonging to the 1975 and 1976 year classes, were present, suggesting that the roach mortalities might be giving the rudd a chance to recover. However, very few rudd were present during 1978, possibly because lower incidences of ligulosis during 1976–1978 had allowed the roach to recover from the main outbreak of the disease in 1975, and to replace the rudd for the second time. Evidence from trap catches indicated that the perch population had shown a consistent decline between 1970 and 1977. The major phase of this decrease occurred between 1971 and 1973, when the roach population was increasing the most rapidly. Comparisons were made with changes in fish populations in other localities. The major reason for the changes observed was thought to be a competitive relationship between the plankton feeding younger stages, with roach being the dominant species, although other factors may also have been involved.     

The dynamics of a population of roach (Rutilus rutilus (L.)) in a shallow lake: is there a 2-year cycle in recruitment?

Recruitment success of roach varied dramatically between 1978 and 1985 in Alderfen Broad, a small lake in eastern England. All size classes of roach feed to a significant extent upon zooplankton, but the underyearling fish have the greatest effects upon the abundance, species composition and mean size of zooplankton. During years of good recruitment (1979, 1981, 1983 and 1985) when the 0 + age group was abundant, they showed poor growth as a result of the depression of their prey populations. Older fish also tended to grow poorly in these years and may have been less fecund the following year. In years of poor recruitment (1980, 1982 and 1984), with the release of the depressive effect upon the zooplankton exerted by underyearling fish, the older size classes tended to grow well with higher fecundity the following season, giving rise to good recruitment of underyearling fish, even when the number of spawners was low. The evidence indicates that there is a 2-year cycle of roach recruitment in Alderfen and this will be described.     

Feeding and Growth Response of Roach, Rutilus rutilus, to Alarm Substance

Feeding and growth responses of roach from three size classes to alarm substance (Schreckstoff) were quantified in laboratory experiments. Larger fish (60.0–80.0 mm in length) reacted stronger to treatment than two smaller sized groups (35.0–45.0 and 46.0–55.0 mm) lowering feeding rate by 80 and 40 and 50%, respectively. The reduction in feeding rate of larger fish caused decrease in growth rate in length and weight, while the lowered consumption of smaller fish caused only reduction in growth rate in weight. Condition factor of exposed to alarm substance small sized roach was lower than that of the control individuals and roach from other two size classes, both, treated and untreated. The difference in growth response to a danger of predation has its roots probably in different metabolism and growth rates of small and large fish. Small fish have higher metabolic rate and less lipid reserves than larger ones, therefore they are probably forced to feed to be able to grow. Also, small sized roach is more vulnerable to predation than large sized fish, thus growing fast seem to be crucial for survival in a risky environment. Study shows that small roach trade off their safety against food, feeding in risky environment to sustain fast growth. This ability of fast outgrowing of a dangerous, vulnerable to predators, size increases survival of juveniles in dangerous environment.     

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.     

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.     

Response of young<Emphasis Type="Italic"> Tsuga heterophylla</Emphasis> to deer browsing: developing tools to assess deer impact on forest dynamics

We used dendroecology to describe and understand the consequences of deer browsing on regenerating western hemlock (Tsuga heterophylla). We compared tree shape, growth rate, height and age at four different sites in Haida Gwaii (British Columbia, Canada) that had trees representative of the range of deer impact on trees: (1) trees showing no sign of browsing, (2) escaped trees which were still browsed below the browse line and (3) stunted and heavily browsed trees. Repeated and intense browsing resulted in the small size, compact heavily ramified shape of stunted trees and in the short compact and ramified lower branches of escaped trees. These contrasted with the shape of non-browsed trees, a shape that was also found in escaped trees above the browse line. Before release, all browsed trees experienced stagnation in growth characterised by narrow rings (0.3 mm/year) and a small annual height increment (2.5 cm/year). At release, growth rate increased and stabilised: rings were wider (1.3 mm/year) and annual height increments were greater (10.5 cm/year). Non-browsed trees had a mean ring-width of 1.3 mm/year and an annual height increment of 22 cm/year. Delay in tree recruitment caused by deer varied from site to site. It had been about 15 years for the escaped trees and is estimated at 30–40 years for the stunted trees. Spatial variation in deer impact may reflect spatial variation of browsing pressure resulting from local differences in the availability of preferred forage or to differences in tree chemical defences/nutritional values.     

Ontogeny of sexual development in the roach (Rutilus rutilus) and its interrelationships with growth and age

The roach (Rutilus rutilus) has become a sentinel species for the study of sexual disruption in wild fish populations as a consequence of exposure to endocrine disrupting chemicals (EDCs). Little is known, however, about the normal ontogeny of sexual development in this species. Here, we analyzed the ontogeny of sexual development in captive‐bred roach and assessed how growth rate and fish size affected the timing of both sexual differentiation and sexual development over a 2‐year period. Ovarian differentiation was first recorded at 68 days post‐fertilization (dpf) and this preceded testicular differentiation (first recorded at 98 dpf). In contrast, sexual maturation occurred at an earlier age in males (300 dpf) compared with females (728 dpf). No differences in body size (length or weight) were recorded between male and female roach until the fish were 415 dpf. Studies on three populations of roach which grew at different rates showed that the timing of sexual differentiation was highly variable and more related to fish size than to fish age. Time to sexual maturation was also variable among populations but, subsequent to their first year of life, gonadal status was less well associated with fish size. Interestingly, the sex ratio of the population was biased towards females in populations that grew more rapidly during early life. The findings presented here provide a valuable foundation of work to support both field‐ and laboratory‐based assessments on the effects of EDCs, and other stressors, on sexual differentiation and development in the roach. J. Morphol., 2008. © 2008 Wiley‐Liss, Inc.