Clonal Atlantic salmon × brown trout hybrids produced by gynogenesis

Clonal full-sib progeny groups of Atlantic salmon Salmo salar × brown trout Salmo trutta hybrids were produced by gynogenesis. Eggs obtained from two 3-year-old Atlantic salmon (female) × brown trout (male) F₁ hybrids were activated with UV-irradiated rainbow trout Oncorhynchus mykiss sperm. Fecundity, percentage egg activation and percentage survival to completion of yolk-sac absorption were similar for the two females, and averaged 800 eggs kg⁻¹, 90 and 65%, respectively. Flow cytometric and protein electrophoretic analyses confirmed the progeny to be diploid hybrids. Isogenicity within progeny groups and to the maternal parent was indicated by identical DNA fingerprint patterns detected with multilocus oligonucleotide probes–GATA₍₅₎ and ACTG_(n). Isogenicity was also observed in the gynogenetic progeny of a third female spawned the following year. It appeared that a large portion of the oocytes in females of this hybrid underwent a premeiotic chromosome doubling, or possibly a complete suppression of meiosis. The result was ovulation of diploid eggs, each possessing a full set of both Atlantic salmon and brown trout chromosomes identical to those in the maternal somatic cells. Lines of clonal hybrids could therefore be perpetuated by gynogenesis and would have potential both as experimental animals and in commercial aquaculture.     

Seasonal and spatial microhabitat selection and segregation in young Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L., in a Norwegian river

Seasonal microhabitat selection by sympatric young Atlantic salmon and brown trout was studied by diving. Both species, especially Atlantic salmon, showed seasonal variation with respect to surface and mean water velocities and depth. This variation is partly attributed to varying water flows and water temperatures. In winter the fish sought shelter in the substratum. A spatial variation in habitat use along the river due to different habitat availabilities was observed. Both species occupied habitats within the ranges of the microhabitat variables, rather than selecting narrow optima. It is hypothesized that the genetic basis allows a certain range to the behavioural response. Microhabitat segregation between the two species was pronounced, with brown trout inhabiting the more slow-flowing and partly more shallow stream areas. Atlantic salmon tolerated a wider range of water velocities and depths. Habitat suitability curves were produced from both species. It is suggested that habitat suitability curves that are based on observations of fish occupancy of habitat at median or base flow may not be suitable in habitat simulation models, where available habitat is projected at substantially greater water flows.     

Sperm motility characteristics of wild Atlantic salmon (Salmo salar L.) and sea trout (Salmo trutta m. trutta L.) as a basis for milt selection

The aim of the study was to determine the sperm motility parameters in wild Atlantic salmon and sea trout to define criteria important for selection of milt for controlled fertilisation. Parameters for these species were determined in the fish migrating into north‐western rivers of Poland at spawning time. Eight motility parameters percentage of motile sperm (MOT), curvilinear velocity (VCL), average path velocity (VAP), straight line velocity (VSL), linearity (LIN), straightness (STR), amplitude of lateral head displacement (ALH), beat cross frequency (BCF) and motility duration were subjected to computer‐assisted sperm analysis (CASA). Milt of most individuals studied representing both salmon and trout showed spermatozoa density of 12–22 × 10⁹ ml^?1 and a high percentage of motile sperm (>70%). In general, spermatozoa swim progressively with slightly curved trajectories (mean STR = 70%, LIN = 65%) and velocity VCL of 180 μm s^?1 (salmon) and 190 μm s^?1 (trout), at 10 s post‐activation. Such sperm is easily accessible in the wild populations of salmon and sea trout and is recommended for use in reproduction trials. The spermatozoa of sea trout seem to show a greater tendency to follow curvilinear trajectories than those of salmon, both in the beginning and the final phase of motion. In the first phase of motility, the values and time dependencies of the motility parameters were similar in both species. In the end phase of movement differences in LIN and BCF time dependencies were found in the samples representing the two species. In salmon the linearity and beat cross frequency remained stable in this phase, contrary to the patterns in sea trout for which LIN decreased while BCF increased in the end period of movement. Durations of movement were similar in both species (ranges of 20–40 s).     

Competition between brown trout and Atlantic salmon parr over pool refuges during rapid dewatering

Variations in distributions and behaviours of Atlantic salmon Salmo salar in allopatry (homogeneous) and in sympatry with brown trout Salmo trutta (mixed) were observed before, during and after 2 day periods of dewatering in a large glass-sided indoor stream at densities typical of Scottish upland streams. Brown trout utilized pools more than Atlantic salmon at normal flows and in both species the majority of fishes moved into pools during dewatering. There was no significant effect of brown trout, which was the more dominant species, on the overall ability of Atlantic salmon to use pool habitat as a refuge during dewatering. Within mixed and homogeneous groups, average feeding levels decreased during dewatering. The highest ranking fish, which was always a brown trout in mixed groups, predominantly monopolized the pool and other individuals in pools adopted a more cryptic, stationary behaviour. Dewatering effectively increased local population density with the result that dominance status became much more important in maintaining food intake, and polarization between the top ranking fish and others increased. During the first day of dewatering, there was extreme behavioural polarization such that the dominant fish exhibited most aggression and least feeding within the group. Among dominant fish on the second day of dewatering, aggression had largely abated and feeding had returned to pretreatment levels despite the reduced average feeding within the group. The main difference between mixed and homogeneous groups was in the behaviour of the most dominant Atlantic salmon, which was near-despotic in allopatry and subordinate to brown trout in sympatry.     

The effect of inorganic and organic pollutants on sperm motility of some freshwater teleosts

Using four different sperm types from brown trout Salmo trutta fario (Salmonidae), chub Leuciscus cephalus (Cyprinidae), burbot Lota lota (Gadidae) and African catfish Clarias gariepinus (Clariidae) the effect of inorganic (cadmium, copper, mercury, lead, zinc and nitrite) and organic (cyclohexane and 2,4‐dichlorophenol) environmental pollutants on sperm motility was investigated. Spermatozoa were activated in double distilled water containing the different test substances and the motility was compared to controls of similar pH. From the investigated motility variables the sperm motility rate and swimming velocity reacted most to the environmental pollutants whereby the changes depended on the species and on the test substance. African catfish spermatozoa were the most resistant, chub and burbot spermatozoa showed medium resistance and brown trout spermatozoa were the most sensitive to the pollutants. With exception of 2,4‐dichlorophenol and zinc the effective concentrations of the tested pollutants exceeded the recommendation for surface waters 100–10·000‐fold and were in a range lethal for the fish themselves. Therefore, it was concluded that fish sperm motility is not a suitable marker for risk assessment of environmental pollutants.     

The effect of temperature on sperm motility and enzymatic activity in brown trout Salmo trutta, burbot Lota lota and grayling Thymallus thymallus

The effect of temperature on sperm motility was investigated in brown trout Salmo trutta, burbot Lota lota and grayling Thymallus thymallus using water and sperm motility prolonging saline solution (SMPS) for motility activation. The effect of temperature (4-20° C) on spermatozoal enzymes for energy supply [malate dehydrogenase (MDH), pyruvate kinase (PK), adenylate kinase (AK)], flagellar movement [Mg(2+) adenosine triposphatase (ATPase)] and oxidative defence [peroxidase (POX)] were measured in S. trutta and L. lota. Temperatures yielding the highest initial sperm motility rates and swimming velocities were 4-6° C for S. trutta [investigated range (IR) = 4-12° C] and L. lota (IR = 2-8° C) and 8-16° C (IR = 4-16° C) for T. thymallus. Motility variables were re-measured after 30 s in S. trutta, after 45 s in T. thymallus and after 60 s in L. lota in water and after 2 min in all investigated species in SMPS. Motility variables were increased by low temperatures and the results differed between water and SMPS. In S. trutta and L. lota, the temperature resulting in highest activities of MDH, PK, AK and ATPase was 4° C. POX had a very narrow temperature optimum at 20° C in both species. This may indicate that the temperature optimum of enzymes of energy supply and flagellar movement are closely related to motility. The present data show that the variables are affected by temperatures in an ecologically relevant range. Too low, as well as too high temperatures affected sperm motility, and the winter spawners (S. trutta and L. lota) have a narrower temperature optimum than the spring spawner T. thymallus.     

Fertilization dynamics in sockeye salmon and a comparison of sperm from alternative male phenotypes

Sockeye salmon Oncorhynchus nerka sperm maintained high fertility for 10 s after being exposed to water but fertility dropped sharply after that. Eggs also showed limited longevity after being immersed in water, with a large drop in fertility occurring after 20 s of exposure to water. The association between eggs and sperm occurs very rapidly. 80% fertilization success was achieved with 5 s of sperm-egg mixing and c. 25% was achieved with <0·5 s of sperm-egg mixing. Sperm from different male age classes differed little in motility and did not differ at all in their propensity to fertilize eggs. The results have important implications for mate choice and sperm competition.     

Timing matters: species‐specific interactions between spawning time,substrate quality,and recruitment success in three salmonid species

Substratum quality and oxygen supply to the interstitial zone are crucial for the reproductive success of salmonid fishes. At present, degradation of spawning grounds due to fine sediment deposition and colmation are recognized as main factors for reproductive failure. In addition, changes in water temperatures due to climate change, damming, and cooling water inlets are predicted to reduce hatching success. We tested the hypothesis that the biological effects of habitat degradation depend strongly on the species‐specific spawning seasons and life‐history strategies (e.g., fall‐ vs. spring‐spawners, migratory vs. resident species) and assessed temperature as an important species‐specific factor for hatching success within river substratum. We studied the species‐specific differences in their responses to such disturbances using egg‐to‐fry survival of Danube Salmon (Hucho hucho), resident brown trout (Salmo trutta fario), and migratory brown trout (Salmo trutta lacustris) as biological endpoint. The egg incubation and hatching success of the salmonids and their dependence on temperature and stream substratum quality were compared. Hatching rates of Danube salmon were lower than of brown trout, probably due to higher oxygen demands and increased interstitial respiration in spring. Increases in maximum water temperature reduced hatching rates of resident and migratory brown trout (both fall‐spawners) but were positively correlated with hatching rates of Danube salmon (a spring‐spawner). Significantly longer incubation periods of resident and migratory brown trout coincided with relatively low stream substratum quality at the end of the egg incubation. Danube salmon seem to avoid low oxygen concentrations in the hyporheic zone by faster egg development favored by higher water temperatures. Consequently, the prediction of effects of temperature changes and altered stream substratum properties on gravel‐spawning fishes and biological communities should consider the observed species‐specific variances in life‐history strategies to increase conservation success.     

Seasonal and Diel Changes in Behaviour,Microhabitat use and Preferences by Young Pool-dwelling Atlantic Salmon,Salmo salar,and Brown Trout,Salmo trutta

There was a pronounced decline in activity of young pool-dwelling Atlantic salmon, Salmo salar, and brown trout, Salmo trutta, as the water temperatures dropped in the autumn and early winter, and the fish switched from a predominantly diurnal towards a nocturnal activity pattern. Such a switch in activity pattern has previously been observed in young brown trout, but the present study is the first documentation for juvenile Atlantic salmon under natural conditions. Juvenile fish fed actively even when water temperatures were below 0°C, although foraging behaviour at near-freezing temperatures was recorded exclusively during night surveys. This indicates that other proximate factors, in addition to water temperature, affect the activity of young salmon and trout in rivers. Trout kept feeding positions significantly higher above bottom than salmon in August and September, but both species reduced the height above bottom at the onset of winter, possibly due to reduced swimming performance and lowered food availability in the upper part of the water column.     

Comparison of the physical properties of real and artificial salmonid eggs and of their performance when drifting in an experimental stream channel

The paper compares three batches of brown trout (Salmo trutta Linnaeus) eggs, two batches of Atlantic salmon (Salmo salar Linnaeus) eggs and the artificial trout eggs described by Ottaway (1981), in terms of their measurable physical characteristics. Attention was given to the effects of temperature and of the stage of development of the real eggs. Comparisons were also made between brown trout eggs and artificial eggs of the rate of fall in a water column, of the value and speed of attainment of terminal velocity when falling through water and of the pattern of settlement within an experimental stream channel.The main measurable characteristics of brown trout eggs (after water hardening) and of artificial eggs (values in parentheses) were: diameter 0.52–0.54 mm (0.54), fresh weight 0.078–0.086 g (0.09), density 1.071–1.075 g ml^–1 (1.066), and volume 0.073–0.081 ml (0.085). There was no evidence of any major change in these values as egg development proceeded. Salmon eggs had a similar density to trout and artificial eggs but had 30 to 70% greater weight and volume. The water-hardening of eggs caused their dry matter content to fall from 37–39% to 31–34% and there were corresponding increases in volume and fresh weight.The rate of fall of trout eggs and artificial eggs through water when timed from rest at the water surface to a depth of 125 cm was similar for both types of egg at 10 °C. This rate rose with increasing temperature at the same low rate of c. 0.02 cm s^–1 °C^–1 for both types of egg. Both types of egg had a terminal velocity of c. 8.8 cm s^–1 and both achieved terminal velocity in less than 1.5 s after release.The patterns of settlement of the two types of eggs in an experimental channel were similar.     

The effect of domestication on some life history traits of sea trout and Atlantic salmon

During the period 1968–1991, certain morphological traits of Atlantic salmon Salmo salar and sea trout S. trutta have been recorded regularly at the hatchery at Älvkarleby, central Sweden. Total body length, weight (for females both before and after stripping), number of eggs, egg size and date of ovulation. A smaller data set for fish marked and released as smolts, providing information about total body length of released smolts, time spent at sea and body size of the recovered adults, was also available for analysis. According to theory and empirical data, the process of artificial breeding results in an evolutionary divergence of the cultured strain from the wild phenotypic norm. The reason for such a divergence is that both natural and sexual selection pressures are altered or relaxed during the process of artificial breeding, as well as random genetic processes, such as founder effects and in- and outbreeding. A path analysis of both species and sexes revealed that the size of the released smolts had increased during the study period. Time spent in sea has decreased for both female and male sea trout, but not for Atlantic salmon. Adult body size for female and male trout have increased as well as female trout condition factor. The increase found in egg size of both species was greater for the salmon than for the trout, indicating that female salmon invest more in egg size with increasing body size. These results support the view that domestication probably has a significant and selective impact on the life history traits of the two salmonid species studied.     

Comparing upriver spawning migration of Atlantic salmon Salmo salar and sea trout Salmo trutta

Radio tagged wild Atlantic salmon Salmo salar(n = 30) and sea trout Salmo trutta(n = 19) were simultaneously released from a sea pen outside the mouth of the River Lærdalselva and their migration to spawning areas was recorded. The distance from the river mouth to a position held at spawning ranged from 2 to 24 km and did not differ between the species (mean ± s .d . 15·9 ± 4·3 and 14·9 ± 5·2 km for Atlantic salmon and sea trout, respectively). The duration of the migration phase, however, was significantly shorter for Atlantic salmon than for sea trout (8–12 days, respectively). All Atlantic salmon migrated straight to an area near the spawning ground, whereas 50% of the sea trout had a stepwise progression with one or more periods with erratic movements before reaching the spawning area. After the migration phase, a distinct search phase with repeated movements up‐ and downstream at or close to the position held at spawning was identified for the majority of the fishes (75%, both species). This search phase was significantly shorter for Atlantic salmon than for sea trout (mean 13–31 days, respectively). Mean ± s .d . length of the river stretch used during the search phase was larger for sea trout (3·3 ± 2·5 km) than for Atlantic salmon (1·2 ± 0·9 km). A distinct holding phase, with no movements until spawning, was also observed in the majority of the Atlantic salmon (80%, mean duration 22 days) and sea trout (65%, mean duration 12 days). For both species, a weak, non‐significant trend was observed in the relationship between time spent on the migration phase, and time spent on the search (r² = 0·43) and holding phase (r² = 0·24). There was a highly significant decrease, however, in the duration of the holding phase with an increase in the time spent on the search phase (r² = 0·67).     

Temporal and spatial segregation of spawning in sympatric populations of Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L.

Based on data from Norwegian streams with sympatric populations of Atlantic salmon and brown trout, it is suggested that temporal segregation is the main mechanism segregating Atlantic salmon and brown trout during spawning. Peak spawning of trout was about 15 days earlier than that of salmon. Physical factors, such as water depth, water velocity and distance from the river banks segregate spawning sites of salmon and trout poorly. Gravel sizes of the redds of salmon and trout were significantly different, though with a considerable overlap, and mean egg depth of salmon and trout were 0.18 and 0.12 m, respectively, probably attributable to the different size of spawners of salmon and trout. None of the temporal or spatial parameters analysed segregate spawners of salmon and trout completely. Species determination of eggs and alevins from the redds showed no interspecific superimposition of redds. It is, therefore, concluded that low survival of hybrids after hatching does not explain the low frequency of hybrids observed in sympatric populations of salmon and trout.     

Juvenile salmonid populations in a temperate river system track synoptic trends in climate

Widespread declines among Atlantic salmon (Salmo salar) and brown trout (Salmo trutta) over recent decades have been linked to pollution, exploitation and catchment modification, but climate change is increasingly implicated. We used long‐term, geographically extensive data from the Welsh River Wye, formerly a major salmon river, to examine whether climatically mediated effects on juveniles (>0+) might contribute to population change. Populations of Atlantic salmon and brown trout fell across the Wye catchment, respectively, by 50% and 67% between 1985 and 2004, but could not be explained by pollution because water quality improved during this time. Stream temperatures, estimated from calibrations against weekly air temperature at eight sites, increased by 0.5–0.7 °C in summer and 0.7–1.0 °C in winter, with larger tributaries warming more than shaded headwaters. Rates of winter warming were slightly greater after accounting for the effect of the North Atlantic Oscillation (1.1–1.4 °C). However, warming through time was smaller than measured variations among tributaries, and alone was insufficient to explain variations in salmonid density. Instead, population variations were best explained in multilevel mixed models by a synoptic variate representing a trend towards hotter, drier summers, implying interactions between climate warming, varying discharge and fluctuations in both brown trout and salmon. Taken alongside recent data showing effects of warming on survival at sea, these data suggest that Atlantic salmon might be jeopardized by future climatic effects in both their marine and freshwater stages. Effects on nondiadromous brown trout also imply climatically mediated processes in freshwaters or their catchments. Climate projections for the United Kingdom suggest that altered summer flow and increasing summer temperatures could exacerbate losses further in these species, and we advocate management actions that combine reduced abstraction with enhanced riparian shading.     

Day and night drift-feeding by juvenile salmonids at low water temperatures

Drift-feeding salmonids in boreal streams face temperatures below physical optima for extensive periods of the year. Because juvenile salmonids react to low water temperatures by becoming nocturnal, knowledge about their foraging ability at low light intensities in cold water is needed to accurately estimate energy intake during non-summer conditions. In a laboratory stream channel, we studied temperature effects on the drift-feeding behaviour of juvenile Atlantic salmon, brown trout, and European grayling in simulated daylight and moonlight at temperatures ranging from 2 °C to 11 °C. Prey capture probability was positively related to temperature, but the temperature dependence did not agree with predictions of the Metabolic Theory of Ecology. Furthermore, reaction distance was positively related to temperature for the three species, which may be one of the underlying mechanisms responsible for the temperature effects on prey capture probability. Overall, the three species had similar capture rates at the different temperature and light levels, although there were species differences. European grayling had a slightly higher prey capture probability than brown trout, and brown trout had a shorter reaction distance than Atlantic salmon and European grayling. These results have implications for both energetics-based drift-foraging theory and for studies of winter ecology.     

Inter- and intrapopulation variation in temperature sum requirements at hatching in Norwegian Atlantic salmon

The duration of the ontogenetic development of the eggs up to hatching time was studied under stable water temperatures for a total of 47 families (the offspring of a single pair of spawners) from eight wild populations and one domesticated stock of Norwegian Atlantic salmon. Eggs from each family were incubated at stable water temperatures of 7·3 and 9·4° C. The number of days required to 50% hatching differed significantly between populations for both experimental temperatures together and for each temperature separately. The interpopulation and intrapopulation variation each accounted for about 50% of the overall variation. Significant differences were found for the days required to 50% hatching between eggs of the domestic stock from The Norwegian Salmon Breeding Company and two natural populations and between some of the wild populations. It is concluded that the time required from fertilisation to hatching in Atlantic salmon is a population-specific adaptive trait.     

Ovarian fluid enhances sperm movement in Arctic charr

Like the spermatozoa of most other fish species spawning in fresh water, Arctic charr Sahelinus alpinus sperm were short-lived (mean 42 s) after activation and their swimming speed declined rapidly during this period, e.g. from a mean speed of 106 um s⁻¹ at 10 s after activation in fresh water to 21 μm s⁻¹ only 20 s later. Ovarian fluid significantly influenced sperm longevity (duration of forward mobility), per cent motility, swimming speed and the linearity of sperm movement. All of these variables generally increased as the concentration of ovarian fluid increased from 0 to 50%, even though ovarian fluid is more than three times as viscous as fresh water. It is concluded that ovarian fluid enhances sperm movement in this species and thus has the potential to influence both fertilization success and the outcome of sperm competition.     

Cryopreservation of sperm from the endangered formosan landlocked salmon (Oncorhynchus masou formosanus)

The Formosan landlocked salmon (Oncorhynchus masou formosanus) are at a high risk of extinction, and the sustained maintenance of the population will soon depend on aquaculture systems, which use cryopreservation of spermatozoa to increase genetic diversity. We investigated the effectiveness of dimethyl sulfoxide (DMSO), dimethyl-acetamide (DMA), and methanol as cryoprotectants in combination with 300 mM glucose as extender on the freezing of Formosan landlocked salmon spermatozoa. We also evaluated the morphological changes of Formosan landlocked salmon spermatozoa after their immediate dilution in the 300 mM glucose-DMSO extender and after freeze-thawing. The spermatozoa frozen with DMSO as a cryoprotectant showed significantly higher post-thaw motility and fertility than spermatozoa frozen with DMA or methanol. The fertilization capacity of frozen-thawed Formosan landlocked salmon was comparable to that of fresh spermatozoa. Intersubspecies fertilization trials between cryopreserved Formosan landlocked salmon spermatozoa and Amago salmon eggs showed high fertilization rates. Based on the findings, the potential value of using sperm bank to safeguard this endangered species is discussed.     

Effects of body size on sympatric shelter use in over‐wintering juvenile salmonids

The study tests two hypotheses: (1) the degree of shelter dominance in Atlantic salmon Salmo salar and brown trout Salmo trutta increases progressively with increasing size differential between heterospecific fish in a pair and (2) shelter dominance, standardized to size differential, correlates with aggression. The results support the first but not the second hypothesis, suggesting that the fitness consequences of high growth performance during the summer are likely to become evident during winter. At this time of year, when mortality is high among both Atlantic salmon and brown trout, shelter dominance may increase the chances of survival.     

Static habitat partitioning and dynamic selection by sympatric young Atlantic salmon and brown trout in south-west England streams

Direct underwater observation of micro‐habitat use by 1838 young Atlantic salmon Salmo salar [mean L_T 7·9 ± 3.1(s.d.) cm, range 3·19] and 1227 brown trout Salmo trutta (L_T 10·9 ± 5·0 cm, range 3·56) showed both species were selective in habitat use, with differences between species and fish size. Atlantic salmon and brown trout selected relatively narrow ranges for the two micro‐habitat variables snout water velocity and height above bottom, but with differences between size‐classes. The smaller fishes <7 cm held positions in slower water closer to the bottom. On a larger scale, the Atlantic salmon more often used shallower stream areas, compared with brown trout. The larger parr preferred the deeper stream areas. Atlantic salmon used higher and slightly more variable mean water velocities than brown trout. Substrata used by the two species were similar. Finer substrata, although variable, were selected at the snout position, and differences were pronounced between size‐classes. On a meso‐habitat scale, brown trout were more frequently observed in slow pool‐glide habitats, while young Atlantic salmon favoured the faster high‐gradient meso‐habitats. Small juveniles <7 cm of both species were observed most frequently in riffle‐chute habitats. Atlantic salmon and brown trout segregated with respect to use of habitat, but considerable niche overlap between species indicated competitive interactions. In particular, for small fishes <7 cm of the two species, there was almost complete niche overlap for use of water depth, while they segregated with respect to water velocity. Habitat suitability indices developed for both species for mean water velocity and water depth, tended to have their optimum at lower values compared with previous studies in larger streams, with Atlantic salmon parr in the small streams occupying the same habitat as favoured by brown trout in larger streams. The data indicate both species may be flexible in their habitat selection depending on habitat availability. Species‐specific habitat overlap between streams may be complete. However, between‐species habitat partitioning remains similar.