Individual variation in early life-history traits in brown trout

Abstract – We studied the variability of early life history traits in individually raised progeny of brown trout Salmo trutta L. Eggs from two dams were each fertilized with sperm from 10 sires, producing 20 full-sib families. Each individual progeny was followed from fertilization until death without exogenous food. Egg size was positively correlated with a number of life-history traits, especially length and yolk sac volume at hatching and length at death, and these traits were strongly influenced by maternal effects. We found significant additive genetic variance for growth rate, length at death and life span without external food. Thus these early life-history traits may be modulated by natural selection. We found positive phenotypic and genetic correlations between most early life-history traits. However, life span without external food was negatively correlated with growth rate, indicating a genetic trade-off. ^Note     

The importance of egg size and egg energy density for early size patterns and performance of Arctic charr Salvelinus alpinus

The primary target of selection in most breeding programmes in aquaculture is to promote rapid growth while delaying maturation. In most cases, little is known about how such selection programmes may affect maternal provisioning of offspring and early development. Under favourable and predictable condition, which is typically the case in fish farming, females generally produce more but smaller eggs than wild females. By using a continuous range of egg size originating from females of a fourth generation of Icelandic domesticated Arctic charr Salvelinus alpinus (Linnaeus, 1758), we assessed the relative importance of maternal effects and individual egg size for energy density of individual egg and several offspring traits from fertilization until 138 days post fertilization (dpf). We found that egg size and related energy density varied more among than within females. Although egg size was not correlated with energy density (at 1 dpf) or survival (at hatching and 138 dpf), it was positively correlated with most offspring traits throughout ontogeny. We further did not find any temporal variation in the correlation with the offspring traits during ontogeny. Within clutch variation in growth, yolk sac volume and hatching time were under the influence of direct female effects (maternal or genetic effects). These results show how egg size relates to individual performance in Arctic charr and constitute a baseline for understanding how domestication may alter offspring phenotypes early in life. These findings are discussed with regard to their consequences for aquaculture of Arctic charr and salmonids in general.     

Maternal variation in juvenile survival and growth of triploid hybrids between female rainbow trout and male brown trout and brook charr

Related sib‐groups of rainbow trout × brown trout and rainbow trout × brook charr triploid hybrids and monospecific diploid and triploid rainbow trout controls were obtained from a common set of rainbow trout dams. On the basis of hybrid juvenile performances, 10 sib‐groups were selected and the corresponding diploid rainbow trouts were raised up to adult stage. Females from each group of rainbow trout were used to produce a second generation of hybrid progeny, the performances of which were analysed for grandmaternal variation and relation with first‐generation relatives. Results showed that hybrid traits (alevin yield and weight, survival and growth of fingerlings) were strongly influenced by maternal origin, and could be correlated to those of rainbow trout controls, but that maternal abilities had a low rate of inheritance. It was concluded that little improvement can be expected through selective breeding within parental populations.     

Parentally acquired differences in resource acquisition ability between brown trout from alternative life history parentage

Dominance hierarchies, where they exist, affect individual food acquisition ability and fitness, both of which have the potential to influence life history pathways. Juvenile salmonids exhibit clear dominance hierarchies in early life. As one of the drivers for the adoption of alternative life histories in salmonids is the relative rate of resource acquisition, there is potential for juvenile behaviour to influence the subsequent life history strategy of the individual. Lacustrine brown trout, Salmo trutta, exhibit a multitude of life histories which includes among others the piscivorous (ferox) life history where individuals grow to large size and have delayed maturity and benthivorous and pelagic life histories where individuals grow to much smaller sizes, however mature earlier. Using a number of observable characteristics of dominance, this study compared differences in behaviour between size‐matched pairs of progeny, reared under common garden conditions which are derived from alternative, co‐existing life history strategy parents. We found that first‐generation progeny of ferox trout were more aggressive, acquired more food, had lighter skin pigmentation and held more desirable positions than the progeny of benthivorous brown trout in an experimental stream system. Ferox trout progeny were dominant over benthivorous brown trout progeny in 90% of trials in dyadic contests. Given such clear differences in dominance, this study indicates that parentally acquired dominance‐related differences, passed through either, or both, of genetic and nongenetic (e.g. maternal effects) means, are likely a contributing factor to the continued maintenance of distinct life history strategies of brown trout.     

Are environmental conditions in Finnish streams limiting to early life-history survival in the nonnative rainbow trout?

The nonnative rainbow trout Oncorhynchus mykiss has been an unsuccessful invader in North European streams, although it has been widely introduced. Here we studied whether early life history stages (egg incubation and hatching, first overwintering) act as filters for the establishment of hatchery rainbow trout. Survival of hatched alevins was approximately 80%, whereas only 47% of the embryos survived. However, the latter value was impacted by the high number of unfertilized eggs. Correlation coefficients with embryo survival rate and environmental variables (pH and temperature) were statistically insignificant. In the overwintering experiments, the survival of rainbow trout was 93%. The growth was generally slowed during the winter, but in the spring the growth of rainbow trout exceeded that of the native brown trout. Our data demonstrated that the survival and growth of rainbow trout during early life-history stages were relatively high and comparable to those of the native brown trout. Based on the variables considered in our study, our results suggest that environmental conditions during early life-history stages are not detrimental for rainbow trout in the study streams.     

Total population density during the first year of life as a major determinant of lifetime body‐length trajectory in marble trout

Abstract – The conditions experienced early in life can strongly influence life‐history trajectories in a variety of animal species. Here, we use data from four isolated populations of the endangered stream‐dwelling salmonid marble trout (Salmo marmoratus Cuvier 1817) living in the Soca and Idrijca river basins (Slovenia) to explore the influence of the total density experienced during and after the first year of life by marble trout year‐classes on body length of marble trout through the lifetime. Analyses were performed by pooling together the stream‐specific datasets to cover a wider range of densities. Mean body length of marble trout year‐classes through the lifetime (from age 1+ to 5+) was negatively related to total density of marble trout during the first year of life. The relationship between density during the first growth period and body length through the lifetime was well described by negative power curves. Total population density after the first year of life was not correlated with body length, thus suggesting that body growth trajectories are heavily determined early in life. Given size‐dependent sexual maturity and egg production in marble trout, the relationship between density early in life and lifetime individual growth may have strong implications in terms of population dynamics and regulation of population size.     

Effects of Egg Size Differences on Juvenile Weight between and within Lots in Rainbow Trout Oncorhynchus mykiss

Effects of egg size on weight and growth of juvenile rainbow trout Oncorhynchus mykiss were investigated within and between lots, using a genetic variant with "golden" phenotype as a marker. Analysis of between-lots variation corroborated earlier findings that egg weight influence on the juvenile body weight disappears after a few months of life. In contrast, within-lot effects of egg size were greater and persisted over a longer time. Egg size did not affect subsequent survival or specific growth of the juvenile trout. The genetic marker caused some detrimental pleiotropic effects on eyed egg yield and juvenile body weight, but not on juvenile survival or specific growth.     

Fecundity, egg size and total egg volume differences in 12 stocks of rainbow trout, Oncorhynchus mykiss Richardson

Abstract. Measurements of egg number, egg size and total egg volume in relation to post-stripped weight, made on the eggs stripped from 12 separately maintained farm stocks of 4-year-old rainbow trout. Oncorhynchus mykiss Richardson, revealed highly significant differences between populations when analysed by regression and covariance statistical techniques. All of the stocks showed increases in fecundity with increasing fish size, with 61% (coefficient of determination or r²) of the variation in fecundity of the 12 stocks explained by the common regression on parental fish size. Comparison of the regressions of the different stocks revealed that the rate of increase of fecundity with increasing fish weight was constant throughout the weight range (i.e. the regressions had similar slopes) although there were highly significant differences in elevation, with some stocks producing almost twice as many eggs as some of the others. By contrast, the relationship of egg size with fish weight was much more variable with three of the stocks showing no significant regression of egg size on fish weight and a further two stocks with data poorly fitted to the regressions. Of the remaining seven stocks only one showed a significant difference in slope of the regression with modest increases in fish weight being associated with large increases in egg size. Only four of the remaining stocks of fish showed significant differences in the elevation of their regressions, i.e. the egg sizes were significantly different after the potentially conflicting influence of fish size was partitioned using covariance techniques. Assessments of total egg volume produced far improved regressions on post-strip weight with 75% (r²) of the variation in the pooled data of the 12 stocks being described by the line of the common regression. In general, there appeared to be a “trade-off” between egg number and size with the most and least fecund fish producing somewhat smaller and larger eggs respectively. All 12 regressions of totai egg volume on fish weight showed a common slope. However, there were significant differences in elevation of the regressions, with some of the stocks of trout producing up to a 55% greater volume of eggs. There appeared to be no relationship between the differences in egg number, size or total egg volume and the type or geograpfiical location of the farms on which the stocks were held. Although it remains to be determined to what extent these differences in reproductive performance are Inheritable, it is recommended that these characteristics should be taken into account by hatcheries in tbe design of long-term broodstock selection programmes for rainbow trout.     

Life‐history variation in a species complex of nonmigratory galaxiids

Life‐history theory predicts that the optimal strategy in the trade‐off between egg size and number varies in relation to resource availability and environmental disturbance. We assessed interspecific differences in egg size, fecundity and other life‐history traits in a species complex of stream‐resident galaxiid fish, which are distributed across a range of contrasting habitat types on the South Island, New Zealand. Oocyte size, fecundity and reproductive effort were measured from gravid females collected immediately before spawning. Proxy measures of stream productivity, flow variability and predation pressure were extracted from modelled data sets. A suite of different egg sizes were identified across species within the Galaxias vulgaris complex, with mean oocyte volume differing by up to 133% between species. The species with the smallest eggs showed mean size‐relative fecundities 246% higher than the species with the largest eggs. A significant negative relationship was found between species’ mean egg size and size‐relative fecundity, suggesting a trade‐off between these traits. Species with larger eggs had larger maternal body size, lower reproductive effort and delayed maturity compared to ‘small‐egg’ species. Consistent with the predictions of life‐history theory, species with larger eggs, lower size‐relative fecundity, lower reproductive effort and delayed maturity were associated with low productivity, stable streams, whereas species exhibiting the opposite set of traits occurred in relatively productive but disturbed systems.     

A genetics analysis of the performance of three rainbow trout broodstocks

The reproductive performance of three domesticated rainbow trout broodstocks spawning at 2 years of age was analysed using data collected over a period of 5 years. Data were collected for post-spawning body weight, volume of eggs spawned, egg size, egg number and fertility to the eyed egg stage. There were significant differences among the stocks attributable to both genetic and environmental causes. In addition, there were marked differences between year-classes, isolated from each other due to 2-year spawning. The differences appeared to be genetic in origin and demonstrated the effect of sampling the original parents.Heritabilities estimated from full-sib comparisons indicated male weight was more influenced by environmental stress at spawning than was female weight. Heritability estimates for egg volume, egg size and egg number ranged from 0.05 to 0.76 for individual stocks, but when averaged for the three stocks the range was from 0.32 to 0.52. Egg size was found to have the lowest heritability of the three traits. The heritability of fertility was found to be low and was the lowest for the stock with the poorest fertility.Genetic and phenotype correlations among the traits were also estimated. The relationship between body weight and egg volume and size were both positive as was that between egg volume and egg size. It was determined that egg volume was the principal determinant of egg number and that the relationship of number to size could be negative. The latter evidence was obtained by artificially adjusting the egg production traits to a constant female body size. The result suggested that females that produce larger numbers of eggs are likely to produce smaller eggs unless the change is associated with a marked increase in body weight. It is recommended that selection for egg size be included in all selection programs and that egg number be ignored in any selection program designed to increase body size.     

Phenotypic plasticity for life history traits in a stream population of the threespine stickleback, Gasterosteus aculeatus L.

Abstract – The life history traits of breeding size, clutch size, egg size and relative clutch mass were examined for evidence of plasticity within a set of six annual samples of threespine stickleback, Gasterosteus aculeatus L., females from south-central Alaska, USA. Three samples (from 1992, 1996 and 1999) were of stickleback native to, and living within, a stream environment, while three (from 1995, 1996 and 1999) represented fish living within a pond environment recently colonized by the stream stickleback. Significant differences between stream and pond fish were found for all four traits. For most traits, pond-living females showed greater variation both across and within years than did stream-living females. Although extremely rapid evolution within the pond, or genetic drift caused by low founding population size, could not be completely ruled out, trait changes across years in both environments were interpreted as representing adaptive plasticity.     

Preliminary report on hybridization experiments in trout — Growth and survival of F1 hybrids

Hybridization experiments were carried out by crossing females of common rainbow trout with the males of albino rainbow trout. Survival rates in egg, hatchling, fry and yearling stages, and the annual growth rate were higher than those of either parent.Reciprocal crosses with females of albino rainbow trout and males of common rainbow trout showed poor survival rate. Gonadial development of these hybrids was normal.     

Prediction of Genetic Variation in Stocks of Offspring and Its Correlation with Viability and Growth Rate

This study tested the accuracy of predictions of genetic variation with Genassemblage 1.0 software and investigated whether offspring with greater genetic variation have higher survivability in aquaculture conditions than offspring with less genetic variation. To achieve these aims, microsatellite polymorphism was used to determine genetic variation in rainbow trout, Oncorhynchus mykiss, spawners and progeny. Spawning pairs were assembled based on the estimated genetic variation of their offspring. The offspring were divided into two groups: (H) group had higher expected values of heterozygosity (0.93) and allelic richness (41), whereas (L) group had lower expected values (0.63 and 31). The heterozygosity, survivability, and growth rate of these groups were recorded. The observed heterozygosity was 0.93 in (H) group and 0.62 in (L) group. (H) group had slightly higher survivability (89.8%) than (L) group (87.6%), but this difference was not significant (p ≥ 0.05), possibly due to residual tetrasomy in the rainbow trout genome. In the offspring, predicted and observed variations were very similar, indicating that Genassemblage 1.0 accurately predicts genetic variation in progeny.     

Comparative ecologies of domestic and naturalised rainbow trout in northern Lake Huron

Northern Lake Huron (Ontario, Canada) supports the largest concentration of freshwater salmonid cage culture farms in Canada and receives inputs of domestic‐strain rainbow trout (Oncorhynchus mykiss) through escapement. We assessed the potential effects of these domestic fish on the naturalised rainbow trout of this ecosystem by comparing their respective ecologies. Mature adults were sampled from spawning tributaries, primarily in spring, and to a lesser extent in fall. Fish of domestic origin comprised ~80% of rainbow trout sampled from sites near cage culture facilities but <20% of rainbow trout sampled from more distant sites. Domestic adults in spawning condition (gametes free‐flowing) were present in the spawning tributaries in both spring and fall sampling seasons, whereas naturalised fish in spawning condition were only observed in the spring. Domestic adults were younger and smaller (in length), appeared to have shorter lifespan and had a higher male to female ratio compared with their naturalised counterparts. Growth rates (change in length with age) of naturalised and domestic females in the wild were similar, but domestic males grew more slowly than naturalised males. Domestic females also produced smaller eggs than naturalised females. Food web positions (inferred from δ¹³C and δ¹⁵N) of domestic and naturalised fish were very similar but varied more strongly with body size in the former. Domestic‐strain rainbow trout of cage culture origin can survive, grow and attempt to spawn in northern Lake Huron and have the potential to compete for food, mates and spawning habitat with naturalised rainbow trout.     

Paternal variation in juvenile survival and growth of the triploid hybrid between female rainbow trout (Oncorhynchus mykiss Walbaum) and male brown trout (Salmo trutta L.)

Abstract Related sib‐groups of rainbow trout × brown trout triploid hybrid and monospecific brown trout controls were obtained from a common set of brown trout sires. On the basis of hybrid juvenile survival and growth, 10 sib‐groups were selected and the corresponding brown trouts were raised up to adult stage. Males from each group of brown trout were used to produce a second generation of hybrid progeny, the performances of which were analysed for grandpaternal variation and relation with first generation relatives. Results showed that hybrid traits were influenced by paternal and grandpaternal origin, but that there was little relation between hybrids and brown trouts, or between successive generations of hybrids.     

Covariation between diploid and triploid progenies from common breeders in rainbow trout, Oncorhynchus mykiss (Walbaum)

The covariation between diploid and triploid progenies from common breeders was investigated in various progeny-testing experiments where either dams or sires were sampled from rainbow trout, Oncorhynchus mykiss (Walbaum), stocks. Triploidiza tion was found to frequently reduce the performance in the traits studied: body length and weight, growth, coefficient of condition and pyloric caeca number. Triploidization also generated significant interactions with the parental breeding value. These interactions were caused in part by the familial variance not being the same in triploids as in diploids, but also by actual ranking differences between diploid and triploid familial performances. However, the effect of these interactions was minor as compared with the amount of variation common to both ploidy levels (genetic correlations averaged 0.7–0.9). Therefore, selection of diploid breeders appeared efficient enough for improving triploid progeny, unless family selection methods including triploid progeny testing were preferred for other reasons. Lastly, it was observed that variances from maternal origin tended to be larger in triploids, whereas variances from paternal origin tended to be smaller, as compared with diploids. This point was discussed referring to the genetic make-up of triploids and in the scope of dams and sires for selective breeding.     

Interspecific differences in early life‐history traits in a species complex of stream‐resident galaxiids

Fish differentially provision resources to eggs and larvae to optimise survival during the critical early life‐history period. Resource allocation is limited by a trade‐off between egg size and fecundity, and the optimal strategy varies with habitat type. This study examines the consequences of egg size differences for the early life‐histories of four closely related galaxiid species which occur in contrasting habitat types on the South Island, New Zealand. Headwater species had substantially larger water‐hardened eggs, longer incubation times, and newly hatched larvae were on average up to 41% longer than lower catchment species. Significant interspecific differences in gape diameter, eye diameter and myomere depth were also observed. Swimming ability was positively associated with larval length at hatch. Interspecific differences in length and swimming ability were generally maintained throughout the larval period, despite larvae being reared under relatively benign conditions where many other studies suggest initial differences should disappear. These results demonstrate the consequences of differential maternal provisioning to the egg for larval traits. The larger larvae of headwater species are likely to be more resistant to starvation and have improved foraging ability compared to lower catchment species; traits which are likely to confer them survival advantages in the low productivity, food‐scarce environments they occupy. The smaller larvae of lower catchment species are likely to be a consequence of females investing in fecundity in these relatively resource‐rich streams.     

A novel breeding design to produce genetically protected homogenous fish populations for on‐growing

We present a novel KING breeding design to produce genetically protected homogeneous fish material for commercial producers from a breeding nucleus. KING F2‐production population is established from the nucleus, first through full‐sib mating within two unrelated high‐quality families to produce inbred F1‐progeny and then resolving the inbreeding in F2 through mating of the unrelated F1‐individuals. Owing to a small number of founders and the inbred F1‐parents, the remaining additive genetic variance is 37.5% of the original. This restricts the use of F2‐progeny to establish new breeding programmes, thereby protecting the genetic material of the nucleus. The theoretical calculations show that a concomitant decrease of phenotypic variance is possible. However, the reduction is considerable only for traits with high heritability (h² > 0.50). The method was tested with rainbow trout. The results revealed that the mean body weight of the KING‐progeny was similar, but, surprisingly, phenotypic variation (especially due to residual variance) was higher compared with either their outbred control group or the nucleus breeding population. Although further evaluation of this breeding design is needed, the results suggest that while genetic protection is achieved, the efficiency of the method to reduce phenotypic variation is limited for economically important traits with low‐to‐moderate heritability.     

Competitive effects between rainbow trout and Atlantic salmon in natural and artificial streams

Competition with non‐native species may impede the restoration of native species, but differences in competitive abilities among intraspecific native populations may make some populations more suitable for reintroduction than others. Here, juvenile Atlantic salmon (Salmo salar) from two allopatric populations (LaHave and Sebago) being used for reintroduction into Lake Ontario were placed into two natural stream sites differing in the presence of ecologically similar rainbow trout (Oncorhynchus mykiss). We assessed the effects of competition in the natural streams on fitness‐related traits and habitat use of the Atlantic salmon. We then compared these effects to those observed in artificial streams from a previous study. Atlantic salmon in natural streams had reduced fitness‐related traits that were associated with suboptimal microhabitats in the presence of rainbow trout, but utilised optimal microhabitats in their absence. In the presence of rainbow trout, the two Atlantic salmon populations exhibited comparable recapture proportions to each other, but the individuals from the Sebago population had better performance (body size and condition) than those from the LaHave population. Responses of both Atlantic salmon populations to competition with rainbow trout were generally similar in both direction and magnitude when compared to results from the artificial stream study. The combined results suggest that native species restoration efforts should be focused on candidate populations that are ecologically suitable to reintroduction environments, as well as on suitable habitats that do not contain exotic competitors. Moreover, this study highlights the value of controlled experiments in artificial environments for predicting fitness‐related performance in natural environments.     

Production of Spotless Allotriploids from Female Non-Spotted Rainbow Trout (Houraimasu), Oncorhynchus mykiss,and Male Amago Salmon,O. rhodurus

ABSTRACT

A new phenotype, spotless allotriploids, was introduced into the aquaculture industry. Spotless allotriploids were produced from female non-spotted rainbow trout, Oncorhynchus mykiss, and male ama-go salmon, landlocked type of O. rhodurus. These allotriploids combine the favorable characteristics of the good flavor of amago salmon, faster growth (compared to that of amago salmon), and absence of spots, preferred by the market. The relationship between the non-spot character (gene) and the external appearance of the allotriploids was traced by analyzing the erythrocyte major axis, the allozymes, and the progeny test in each maternal genotype. Results indicate that spotless allotriploids can be produced by using female dominant homozygotic (hh) non-spotted rainbow trout. When females are heterozygotic (hn) non-spotted rainbow trout, the offspring produced are both spotted and spotless allotriploids in approximately equal proportions.