Hatching Time and Alevin Growth Prior to the Onset of Exogenous Feeding in Farmed,Wild and Hybrid Norwegian Atlantic Salmon

The onset of exogenous feeding, when juveniles emerge from the gravel, is a critical event for salmonids where early emergence and large size provide a competitive advantage in the wild. Studying 131 farmed, hybrid and wild Norwegian Atlantic salmon families, originating from four wild populations and two commercial strains, we investigated whether approximately 10 generations of selection for faster growth has also resulted in increased somatic growth prior to the onset of exogenous feeding. In addition, we tested whether relaxed selection in farms has allowed for alterations in hatching time between farmed and wild salmon. Across three cohorts, wild salmon families hatched earlier than farmed salmon families, while hybrid families displayed intermediate hatching times. While the observed differences were small, i.e., 1–15 degree-days (0–3 days, as water temperatures were c. 5–6°C), these data suggest additive genetic variation for hatching time. Alevin length prior to exogenous feeding was positively related to egg size. After removal of egg size effects, no systematic differences in alevin length were observed between the wild and farmed salmon families. While these results indicate additive genetic variation for egg development timing, and wild salmon families consistently hatched earlier than farmed salmon families, these differences were so small they are unlikely to significantly influence early life history competition of farmed and wild salmon in the natural environment. This is especially the case given that the timing of spawning among females can vary by several weeks in some rivers. The general lack of difference in size between farmed and wild alevins, strongly suggest that the documented differences in somatic growth rate between wild and farmed Norwegian Atlantic salmon under hatchery conditions are first detectable after the onset of exogenous feeding.     

Factors affecting river entry of adult Atlantic salmon in a small river

In this study, effects of stock origin, fish size, water flow and temperature on time of river ascent of adult Atlantic salmon Salmo salar were tested. Brood stocks were collected in eight Norwegian rivers situated between 59 and 69° N. The fish were reared to smolts, individually tagged and released in the River Imsa, south-west Norway (59° N). Adults from all stocks approached the Norwegian coast concurrently, but Atlantic salmon ≥70 cm in natural tip length entered coastal water slightly earlier during summer than smaller fish. Atlantic salmon <70 cm, however, ascended the river significantly earlier and at lower water flow and higher water temperature than larger fish. Although largest in size, the fish from the northern populations (62–69° N) ascended the River Imsa almost 1 month earlier than those from the south (59–60° N). They seemed less restricted by the environmental factors than the fish originating from the more southern rivers. There was no apparent trend among years in time of river ascent. Maximum ascent per day occurred at water discharges between 12·5 and 15 m³ s⁻¹ and at water temperatures between 10 and 12·5° C. There was a significant positive correlation between water flow and river ascent during the first part of the upstream run from July to September with best correlation for September, when multiple regression analysis indicated that water temperature had an additional positive effect. Stock origin, fish size and water discharge were important variables influencing the upstream migration of Atlantic salmon in small rivers.     

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.     

Influence of egg vitamin A status and egg incubation temperature on subsequent development of the early vertebral column in Atlantic salmon fry

The effect of egg vitamin A (VA) status and egg incubation temperature on the development of spinal disorders was investigated in Atlantic salmon Salmo salar fry. Atlantic salmon eggs were sorted into two groups with high VA (3·3 ± 0·1 μg retinol g⁻¹ dry mass) and low VA (2·2 ± 0·3 μg retinol g⁻¹ dry mass) status before fertilization and incubated at high (14° C) or low (8° C) temperature from 133 day degrees until the onset of feeding. High egg incubation temperatures increased the concentration of retinol in the eggs: the high VA and high temperature group displayed a significantly higher retinol concentration than the high VA and low temperature group ( P  = 0·001). After hatching, all experimental groups increased their retinol concentration. The source of the increased retinol levels was probably retinal, although astaxanthin may also be a VA precursor after hatching. Atlantic salmon fry incubated at high temperatures had increased amounts of notochord tissue. When measuring morphogenic activity in the notochord using the expression of sonic hedgehog ( shh , mRNA), however, no significant difference was found between the experimental groups. No clear effect of VA status or incubation temperature could be found on the formation of the early vertebral column although Atlantic salmon fry incubated at low temperatures had less regular constrictions of the prospective vertebral column than fry incubated at high temperatures.     

The effect of acid water on the hatching of salmonid eggs

The survival of salmon Salmo salar , sea trout and brown trout Salmo trutta eggs in acid water in the range pH 3·5±7·0 was investigated over a period of three winters. The effects of fertilising the eggs in acid water at pH 4·0, 4·5 and 5·0, and dilution of the test media were also studied. There was considerable yearly variation in the numbers of eggs which hatched, but there were no marked differences between the three species in the tolerances of the eggs to acid water; pH 3·5 was lethal within 10 days to all the eggs, but at pH 4·5 and higher pH levels there was no obvious difference in hatching attributable to acidity.     

Interspecific and intraspecific variations in egg hatching for British populations of Taeniopteryx nebulosa and Brachyptera risi (Plecoptera: Taeniopterygidae)

Adults were obtained from three populations of Taeniopteryx nebulosa and four populations of Brachyptera risi ; their eggs were incubated at seven constant temperatures (range 3.8–22.1°C). There were interspecific, but not intraspecific, differences in adult life-span, mean number of eggs laid per female, hatching success and egg incubation periods. The optimum temperature for hatching success and the range over which at least 50% of the eggs hatched were lower for T. nebulosa (6.5°C, 2.7–15.0°C) than for B. risi (9.0°C, 5.1–15.8°C). No eggs hatched at 22.1°C. The relationship between incubation period (d days) and water temperature (T°C) was given by; d = 326.4 T^−1.015 for T. nebulosa , d = 824.0 T^−0.739 for B. risi . Both equations successfully predicted incubation periods for eggs placed in a stream.
Hatching success and incubation periods were similar to those already published for a Norwegian population of T. nebulosa . The lack of significant intraspecific variation suggests that the genotypes associated with the variables examined in this study have remained remarkably stable in these two species in spite of the geographical isolation of their different populations.     

Inter- and intra-population morphological differences between wild and farmed Atlantic salmon juveniles

Whether population-specific morphological differences were detectable in small (26–52 mm) Atlantic salmon Salmo salar parr reared under similar conditions was tested. Discrimination based on morphological characters was total (100%) between the fish of farmed origin (from AquaGen) and four wild fish populations. Between the four wild populations the corresponding discrimination was 59·8–86·3%. The inter-population variation in morphological characters was larger than the intra-population variation. The fish originating from the local populations at Driva and Innfjord were narrower in body form, whereas fish from the AquaGen and Innfjord populations had smaller and less pointed heads with smaller eyes. The Driva population fish had the smallest mouth while the longest pectoral fin was found in the Bjoreio population, the river that also has the largest fall gradient. Population-specific morphological characters were thus detectable among Atlantic salmon parr relatively rapidly after yolk absorption.     

Temporal and spatial variation in growth of juvenile Atlantic salmon

Spatial and temporal variation in length‐at‐age and environmental factors affecting variation in growth rate of juvenile Atlantic salmon Salmo salar were studied using data from a long‐term study in the River Stjørdalselva, central Norway. Mean annual instantaneous growth rate among 1+ and 2+year juvenile Atlantic salmon varied between 0·59 and 1·50 g g⁻¹ year⁻¹ and mean instantaneous daily growth rate of young‐of‐the‐year (YOY) varied between 0·013 and 0·033 g g⁻¹ day⁻¹. Between year variation in growth was larger than the within year intra‐watercourse spatial variation. For YOY and 1+year Atlantic salmon, a major part of the observed between year variation in growth rates was explained by variation in mean daily water discharge and spring temperature. For 2+year juvenile Atlantic salmon, mean daily water discharge and cohort density were the only variables to significantly explain variation in growth rates. A large part of the within water‐course spatial variation could not be explained by temperature variations and juvenile Atlantic salmon in the uppermost areas of the river, experiencing the lowest ambient temperatures during the growth period, displayed the highest growth rates. Within the baselines set by temperature, biotic and abiotic factors connected to water flow regime and variation in food availability are suggested to be a major determinants of the temporal and spatial variation in juvenile Atlantic salmon growth rates.     

The role of food particle size in the growth of juvenile Atlantic salmon (Salmo salar L.)

The effect of particle size of a commercial pelleted feed on the growth of Atlantic salmon from first feeding alevins to first and second year smolts was investigated using 20-day feeding experiments. Ten experiments were performed over a two year period, each comprising six groups of fish separately selected from stock populations. Each group was presented with one of six sizes of food particle ranging from larger than the respective mean mouth breadth (100% feed size), through 50%, 25%, 12·5%, 6·25% to 3·125%. Experiments were performed in six radial flow/circumferential drain tanks under ambient photoperiod and water temperature. Growth rate was found to be closely related to feed size. Maximum growth in each case was shown only on one size of particle; larger and smaller sizes resulted in reduced growth. The particle size for maximum growth increased in direct proportion to fish length. Fish from 4·2 to 20·3 cm in length showed maximum growth on particle diameters 0·022 to 0·026 × fish fork length (PFR). First feeding alevins were found to show comparable growth rate on particle diameters 0·0115 to 0·090 PFR. Some seasonal variation in growth response was indicated. The results are discussed in relation to developmental and seasonal effects. Possible factors affecting the energetics of prey size related differential growth are discussed.     

Reproductive biology of the endangered Oxleyan pygmy perch Nannoperca oxleyana Whitley

The reproductive biology of the Oxleyan pygmy perch Nannoperca oxleyana is described from simultaneous studies of wild populations in north-eastern New South Wales and mature fish held in aquaria. In the wild, 50% of males and females matured at total lengths of 24·0–25·9 and 28·0–29·9 mm, respectively. The species displays sexual dichromatism during the spawning season, with males developing more intense red and brown fin and body colouration, and black pelvic fins. Captive male broodfish displayed territoriality during the breeding season, closely guarding sites within artificial, plant-like substrata in which pairs of fish spawned adhesive eggs. Protracted serial spawning of wild and captive fish occurred from September to April and May at mean water temperatures ≥16·6° C and day length ≥10·7 h. Captive broodfish spawned on an average of 57% of days during the 256 day spawning period. Gonado-somatic indices averaged 0·7% for all ripe males and 4·1–4·2% for all ripe females collected. Mean total and batch fecundities of captive females were 1323 eggs per fish and 7·8 eggs per fish per day, respectively, and relative fecundity was 587 eggs g⁻¹ of body mass. Batch fecundity of wild females was estimated at 7·8 eggs per fish. The adaptive significance of this reproductive strategy in a harsh, variable environment is discussed.     

Do Norwegian Atlantic salmon feed in the northern Barents Sea? Tag recoveries from 70 to 78° N

Three tagged Atlantic salmon Salmo salar were recaptured as subadults or adults (1·4–3 kg) between 70·5 and 78° N in the western Barents Sea, two of which originated from the Alta Fjord region in northern Norway and one from the Drammen River, south-eastern Norway. An additional tag was recovered from the stomach of a Greenland halibut Reinhardtius hippoglossoides captured south-west of Bear Island at >600 m depth; this tag was from a smolt released in the River Alta 1 month earlier. These are the northernmost tag recoveries reported for Atlantic salmon, and indicate that Norwegian Atlantic salmon, especially the fish from northern populations, may use the northern Barents Sea as a feeding area during part of their life cycle.     

Emergence of Baltic salmon, Salmo salar L., in relation to temperature: a laboratory study

Emergence pattern and developmental status at emergence of Baltic salmon fry from the Umeälven hatchery stock (63°50'N, 20°25'E) were studied at 6, 10 and 12°C in the laboratory. The number of days and degree days from hatching to 50% emergence decreased exponentially with increasing temperature. Synchronization of emergence increased with temperature. Optimal temperature for incubation of yolk-sac alevins was 10°C, which resulted in the largest fry at emergence and the lowest death rate. Fry kept at 6°C had the lowest mean weight and at 12°C the highest death rate. The fry emerged at an earlier developmental state with more yolk at 12°C than at 6°C. The Baltic salmon had a faster developmental rate during the gravel-phase, as compared to more southern Atlantic salmon populations.     

The survival of semi‐wild, wild and hatchery‐reared Atlantic salmon smolts of the Simojoki River in the Baltic Sea

The recapture rate and survival of hatchery‐reared Atlantic salmon Salmo salar stocked as 1 year‐old parr (semi‐wild) with that of hatchery‐reared Atlantic salmon stocked as 2 year‐old smolts and wild smolts of Atlantic salmon in the northern Baltic Sea were compared. This was done through tagging experiments carried out in 1986–1988 and 1992. The recapture rate of the semi‐wild groups varied from 1·0 to 13·1%, being similar in 3 tagging years and lower in 1 year than that of the wild groups (1·7–17·0%). The recapture rate of the semi‐wild groups was similar (in 2 years) or higher (in 2 years) than that of the hatchery‐reared groups stocked as smolts (1·3–6·3%). The survival of semi‐wild smolts during the sea migration was as high as that of wild Atlantic salmon of an equal size and two to three times higher than hatchery‐reared Atlantic salmon stocked as smolts. The survival rate was positively associated with smolt size. The suitability of hatchery‐reared parr and smolts in the management of reduced Atlantic salmon stocks is compared.     

The carotenoids of eggs of wild and farmed Atlantic salmon, and their changes during development to the start of feeding

The only carotenoid detected in newly fertilized eggs of wild Atlantic salmon, Salmo salar, from western Scotland was astaxanthin at a concentration [μg carotenoid g^?1 wet wt of eggs, mean ±S.D. (number of parental females)] of 6.2±1.2(7) in 1982, 6.4±1.8(20) in 1983, and 7.6 ± 13(6) in 1984. In eggs of farmed Atlantic salmon the only carotenoid detected was canthaxanthin at concentrations which varied significantly between farms depending on the level of synthetic canthaxanthin in the broodstock diet. Thus on two farms using feed with 50 μgg^?1, the levels were 11.8 ± 3.4(7) and 12.3 ± 2.9(6), while on two farms using 75μgg^?1 the levels were 18.7 ± 5.0(9) and 21.2 ± 2.7(21). The levels in eggs of one-seawinter fish (grilse) did not differ from those of two-seawinter fish reared on the same farm and diet. During development from newly fertilized egg to fry at the end of yolk-sac absorption, the quantity of carotenoid present per individual decreased, presumably as a result of metabolism. Despite large differences in quantity present, the quantity so metabolized was fairly constant at 2–4 μg carotenoid g^?1 original egg weight for eggs from two-seawinter farmed and wild salmon, except that in eggs from farmed grilse it was 7 μg g^?1. In fry from wild eggs, 99.14% of the remaining carotenoid was present in the integument (skin and fins) as astaxanthin, astaxanthin monoester and astaxanthin diester. In fry from farmed salmon eggs, 47 ± 8% of the carotenoid present was found in the unused yolk oil droplets and in the liver, and 37 ± 6% was found in the integument as canthaxanthin and an unidentified metabolite of canthaxanthin. These findings explain visible colour differences between fry from wild parents and fry from canthaxanthin-fed farmed parents, particularly in the fins, liver and residual oil droplets. The canthaxanthin metabolite was also found, together with canthaxanthin, in the skin of farmed adults fed canthaxanthin. Preliminary tests showed it to be unchanged by saponification but reduced by sodium borohydride. For eggs from the three farms incubated under the same conditions in the same season, percentage mortality both to the eyed stage and between hatching and first feeding varied significantly between farms, but percentage mortality between the eyed stage and hatching did not do so. Results combined from two seasons for eggs from three farms and one wild source showed that egg mortality between fertilization and the eyed stage was not significantly different between wild and farmed salmon, but mortality between the eyed stage and hatching, and between hatching and first feeding, were both significantly higher in farmed salmon than in wild salmon. Such differences could not be explained simply by the large differences in egg carotenoid content, but were almost certainly due to factors such as broodstock nutrition, broodstock management, and stripping and fertilization procedures.     

Thermal variation near the thermal optimum does not affect the growth,metabolism or swimming performance in wild Atlantic salmon Salmo salar

Typically, laboratory studies on the physiological effects of temperature are conducted using stable acclimation temperatures. Nonetheless, information extrapolated from these studies may not accurately represent wild populations living in thermally variable environments. The aim of this study was to compare the growth rate, metabolism and swimming performance of wild Atlantic salmon exposed to cycling temperatures, 16–21°C, and stable acclimation temperatures, 16, 18.5, 21°C. Growth rate, metabolic rate, swimming performance and anaerobic metabolites did not change among acclimation groups, suggesting that within Atlantic salmon's thermal optimum range, temperature variation has no effect on these physiological properties.     

Spatial synchrony in the response of a long range migratory species (Salmo salar) to climate change in the North Atlantic Ocean

A major challenge in understanding the response of populations to climate change is to separate the effects of local drivers acting independently on specific populations, from the effects of global drivers that impact multiple populations simultaneously and thereby synchronize their dynamics. We investigated the environmental drivers and the demographic mechanisms of the widespread decline in marine survival rates of Atlantic salmon (Salmo salar) over the last four decades. We developed a hierarchical Bayesian life cycle model to quantify the spatial synchrony in the marine survival of 13 large groups of populations (called stock units, SU) from two continental stock groups (CSG) in North America (NA) and Southern Europe (SE) over the period 1971–2014. We found strong coherence in the temporal variation in postsmolt marine survival among the 13 SU of NA and SE. A common North Atlantic trend explains 37% of the temporal variability of the survivals for the 13 SU and declines by a factor of 1.8 over the 1971–2014 time series. Synchrony in survival trends is stronger between SU within each CSG. The common trends at the scale of NA and SE capture 60% and 42% of the total variance of temporal variations, respectively. Temporal variations of the postsmolt survival are best explained by the temporal variations of sea surface temperature (SST, negative correlation) and net primary production indices (PP, positive correlation) encountered by salmon in common domains during their marine migration. Specifically, in the Labrador Sea/Grand Banks for populations from NA, 26% and 24% of variance is captured by SST and PP, respectively and in the Norwegian Sea for populations from SE, 21% and 12% of variance is captured by SST and PP, respectively. The findings support the hypothesis of a response of salmon populations to large climate‐induced changes in the North Atlantic simultaneously impacting populations from distant continental habitats.     

Changes in polyamines during embryonic development of Atlantic salmon, Salmo salar.

1. Polyamine composition of eggs and alevins from cultured and wild Atlantic salmon, Salmo salar, was compared during early development. 2. Eggs collected from wild stock had higher amounts of bound and free putrescine, spermidine and spermine than those collected from cultured stock. 3. Cadaverine was present in bound form in cultured and wild stocks before and after fertilization, respectively. However, free cadaverine was detected in both stocks throughout development. 4. Atlantic salmon embryos and alevins can synthesize polyamines which could be associated with their growth and development.     

Temperature-dependent development rates of cod Gadus morhua eggs

Temperature development relationships were determined for batches of Irish Sea cod Gadus morhua eggs incubated in flow-through incubators. Hatching began 16·4 days after fertilization (DAF) at 6° C, 10·3 DAF at 8° C, 9·4 DAF at 10° C and 7·4 DAF at 12° C. Egg mortality increased at the higher temperatures, but survival was >80%. Results were compared with published data at four comparable stage end points: the end of blastula, the end of gastrula, the point of growth of the embryo completely surrounding the yolk and the point when 50% of the eggs were hatched. All the studies showed a curvilinear relationship between age at stage and temperature. There was a 12 day inter-study difference in time to 50% hatch at 2° C and 4 day difference at 10° C. There were no consistent trends that differentiated eastern v. western, or northern v. southern populations. A single model for cod egg incubation time from fertilization to 50% hatch was derived based on data from six cod populations, but it is recommended that individual stock relationships should be used where possible.     

Initial feeding time of Atlantic salmon,Salmo salar,alevins compared to river flow and water temperature in Norwegian streams

Synopsis The time of initial feeding of Atlantic salmon,Salmo salar, alevins in ten geographically widespread Norwegian streams was estimated theoretically by combining data on spawning time and models describing the time from fertilization to hatching and from hatching to initial feeding at different temperatures. The point of initial feeding was correlated with water flow and water temperature regimes. Initial feeding was avoided in all rivers during spring peak flow, and before water temperature reached 8°C. Two different strategies were indicated: (1) initial feeding may take place before the culmination of the spring flow or (2) after the peak spring flow. The choice of strategy depends on temperature and flow regimes in each river.     

The use of genetic marking to assess the reproductive success of mature male Atlantic salmon parr (Salmo salar, L.) under natural spawning conditions

The reproductive success of mature male Atlantic salmon parr ( Salmo salar ) under natural spawning conditions was estimated using the polymorphism at the MEP - 2 * locus as a genetic marker. The percentage of eggs per redd fertilized by parr varied considerably over the five redds examined (0·9–27.7%, mean 10·8%), but a gametic contribution from mature parr was detected in each case. Parr reproductive success has important implications for the population structure and evolution of the Atlantic salmon through its effect on gene flow.