Early predictors of life-history events: the link between first feeding date, dominance and seaward migration in Atlantic salmon, Salmo salar L.

Previous studies have suggested that the earliest fry to emerge from a salmonid redd may have an advantage in the subsequent competition for feeding sites, partly through a 'prior residence' effect. Here we examined whether there was any relationship between the relative date of first feeding and subsequent dominance status and growth in a sibling group of Atlantic salmon, Salmo salar L., fry. Earlier-feeding fry were dominant over their later-feeding siblings (controlling for prior residence), despite not being any larger. However, these early fish soon established and then maintained a size advantage. This led to an increased probability of early-feeding fish migrating to sea at age 1 year (rather than 2 or more). Thus a difference of less than 1 week in the relative timing of first feeding can translate into a year's difference in the timing of migration.     

Prior residence, territory quality and life-history strategies in juvenile Atlantic salmon (Salmo salar L.)

Three groups of juvenile salmon were introduced sequentially into an artificial stream to investigate the effects of prior residence on behaviour and territory choice. Almost half of the first group obtained and defended distinct territories, the other half being constrained to an area approximately the size of one large territory. All of the fish in the subsequent groups, bar one, were also constrained to the same site. Since the fish were of similar size, prior residence alone seemed to influence which individuals obtained territories. However, within the first group, the fish that obtained territories were larger and more aggressive. The territorial fish did not appear to choose the most profitable territories, although they had the greatest opportunity to do so. Since juvenile salmon emerge from their gravel nests fairly synchronously, a time constraint on site sampling is hypothesised: there may be a risk in taking time to sample sites, since these same sites may become occupied with conspecifics. However, fish with territories fed at faster rates than non-territorial fish, possibly because of reduced competition for prey items. Consequently, fish from the first group (containing most of the territorial fish) grew faster than the other two groups. Moreover, most of the territory holders, but only one of the non-territorial fish, reached the threshold size that increases their probability of smolting the following year. This suggests that ability to obtain a defensible territory, primarily through prior residence, influences the age at which juvenile salmon can migrate to sea.     

Analysis of early growth of guppy strains,Poecilia reticulata,with different color patterns

Summary The guppy, Poecilia reticulata, is economically the most important species of freshwater ornamental fish cultured in Singapore. About 30 strains with different color patterns and fin shapes are reared in guppy farms practising monoculture in Singapore. To compare the growth rates of domesticated strains with different color patterns, newborn fry of 11 strains were obtained on the same day from a single farm in Singapore and were reared experimentally in the laboratory for about 100 days. Each strain was distributed randomly into 4 tanks with 30 fish/tank. Weekly weighings of 10 fish/tank were made from 17 to 100 days of age. Three strains were homozygous for the autosomal recessive blond gene which gives rise to a pale yellow background pigmentation (bb). These blond strains had significantly smaller body weights than corresponding ones with the same color pattern but with the wild-type grey-brown background coloration due to the dominant allele (BB). The strains with the red tail pattern due to a dominant X-linked gene (Rdt) had more rapid growth than those with other tail color patterns including the blue, black, green snakeskin and variegated. However, no significant differences were detected among the other color pattern strains. Thus among the strains studied, the blond strains were associated with slower growth while those with the red tail color were associated with faster growth.     

Effects of body size and predators on intracohort competition in wild and domesticated juvenile salmon in a stream

Competition between masu salmon (Oncorhynchus masou) of wild and aquaculture origin was investigated. Fry were individually marked and released in stream enclosures with and without a piscivorous predator. The aim was to assess the effects of predators and salmon body size on survival and growth of the two types of fish under natural conditions. The presence of predaceous Japanese huchen (Hucho perryi) resulted in lower mean growth rates of surviving fry. Relatively large fish survived and grew better than relatively small fish in the absence of predators, but not in their presence. This probably indicates an indirect effect of predation risk on within-cohort competitive behavior among salmon juveniles, with larger fish forced to give up their position as superior competitors. Domesticated fish survived in larger numbers and grew much faster than wild fish, irrespective of predator presence. Comparison with similar field studies revealed a pattern that the pre-experimental environment influenced the outcome of competition between wild and domesticated juvenile salmon. Domesticated fish were superior competitors even in the absence of an initial size advantage, which commonly gives a further advantage to hatchery-raised fish in natural streams. Therefore, caution dictates to avoid the release or escape of such fish into the wild.     

Social status and growth rates as determinants of site attachment in juvenile Atlantic salmon

This paper describes an experimental study of the effects of food supply, growth rates and social interactions on homing by juvenile Atlantic salmon Salmo salar in response to displacement. Groups of five fish were housed in a section of an artificial stream and given either rations allowing maximum growth (the rich condition) or 0·1 of this amount (the poor condition); daily specific growth rates were significantly higher in the rich condition. After a 6-day settlement period, the fish were captured, displaced downstream and their movements recorded over the next 3 h. Prior to displacement, the fish showed a high degree of site fidelity and high levels of aggression. Dominant fish and those with stronger site attachment grew faster prior to displacement, these effects being independent. Following displacement, 24% of all fish returned to their previously favoured site and stayed there, 23% returned home initially, but subsequently moved on, 5% settled in a new site and 49% failed to move. The distribution of responses was identical for the rich and poor conditions, but fish that homed were dominant and had grown faster during the pre-displacement period.     

Factors affecting the outcome of territorial contests between hatchery and naturally reared coho salmon parr in the laboratory

In aquarium experiments using coho salmon as a model species, prior residents dominated intruders of the same size but intruders with a 6% length advantage were equally matched against prior residents. Prior winning experience (distinct from individual recognition) also strongly influenced competitive success and overcame a prior residence effect. Coho salmon reared in a hatchery dominated size-matched fish from the same parental population reared in a stream. Hatchery-reared salmon also dominated naturally spawned salmon, even when the wild salmon were prior residents. Thus the combined effects of greater size and rearing experience of hatchery-produced salmon were sufficient to overcome a wild salmon's advantage of prior residence. Efforts to rehabilitate salmonid populations must consider such behavioural interactions if displacement of wild fish is to be prevented.     

Hierarchical status and colour preference in Nile tilapia (<Emphasis Type="Italic">Oreochromis niloticus</Emphasis>)

We studied the colour preference of isolated Nile tilapia (Oreochromis niloticus) and whether previous residence or body size can affect environmental colour choice. In the first phase, a cylindrical tank was divided into five differently coloured compartments (yellow, blue, green, white and red), a single fish was introduced into the tank and the frequency at which this fish visited each compartment was recorded over a 2-day study period. An increasingly larger fish (approx +2 cm in length each time) was then added into the tank on each of days 3, 5 and 7 (=four fish in the tank by day 7), and the frequency at which each fish visited the different compartments of the tank was observed twice a day to obtain visit frequency data on the differently sized fishes. This experiment was replicated six times. In the first phase, the solitary fish established residence inside the yellow compartment on the first and second days. Following the introduction of a larger fish, the smaller fish was displaced from the occupied compartment. Nile tilapia possibly shows this preference for yellow as a function of its visual spectral sensitivity and/or the spectral characteristics of its natural environment. Moreover, body size is an important factor in determining hierarchical dominance and territorial defence, and dominant fish chose the preferred environmental colour compartment as their territory.     

Effects of crowding and availability of food on growth rate of fry in the European and Chinese races of the common carp

Fry of the European and Chinese races of the common carp and interracial crossbred fry were grown together in a series of ponds for 28 days after hatching. The degree of crowding and the amount of available natural food were intentionally varied in these experimental ponds. This resulted in a large variation in mean growth between ponds, ranging from 0·1 to 5·2 g per fish during the 28 days growth period. Under conditions of very slow growth, resulting from severe crowding and lack of natural food, fry of the Chinese carp grew several times faster than European carp. At lower densities and when natural food was more abundant, resulting in a faster rate of growth, this difference between the two races disappeared. Crossbred fry between the Chinese and European carp grew faster than either of their parents under all conditions tested.     

The effects of intra- and inter-specific competition on the survival and growth of stocked juvenile Atlantic salmon, Salmo solar L., and resident trout, Salmo trutta L., in an upland stream

The relative effects of inter- and intra-specific competition on the survival and growth of stocked salmon were investigated in an upland trout stream during summer and winter sampling periods. The stream was divided into two areas by an impassable fish barrier, and trout were removed from the upstream section prior to 2 years of salmon stocking. Salmon fry stocked into the cleared area survived more than twice as well and grew significantly larger than those stocked into the area containing trout and older salmon. Intra-specific competition from older salmon in the second year of stocking in the cleared area significantly reduced the survival and growth of the O+ salmon. However, these were still significantly larger and survived better than those in the control area where inter-specific competition from trout was maintained. Some immigration of trout to the cleared area occurred; these showed greatly enhanced growth rates compared to those in the control area, reflecting low intra-specific trout competition in the former. Inter-specific competition effects of older salmon on both trout fry growth and survival were also detected, although the latter did not become apparent until the winter. This is discussed in terms of the relative importance of biotic and abiotic regulating mechanisms. Evidence of allopatric niche segregation is also discussed, since salmon in the cleared area did not have a biomass equivalent to that in the area which also contained trout.     

The effects of intra- and inter-specific competition on the distribution of stocked juvenile Atlantic salmon, Salmo salar L., in relation to depth and gradient in an upland trout, Salmo trutta L., stream

The relative effects of inter- and intra-specific competition on the distribution of stocked salmon in relation to depth and gradient were investigated in an upland stream during two summer sampling periods. The stream was divided into two areas by an impassable fish barrier, and trout were removed from the upstream section prior to 2 years of salmon stocking. A small amount of trout re-immigration to the cleared area occurred. Under sympatric conditions in the downstream section the fry of both species were significantly more abundant in shallow water, whereas the yearling and older fish tended to inhabit the deeper, slow-flowing areas. Under near allopatric conditions in the cleared section salmon fry changed their distribution to include deeper areas, with shallow, fast-flowing water becoming the least preferred habitat. This trend for a wide distribution of salmon fry over all the available habitat in the upstream section was maintained in the second year when parr were present but trout remained at very low densities. It was therefore concluded that high inter-specific competition from trout was responsible for restricting the distribution of salmon fry to shallow habitat in the control area. Intra-specific competition from older salmon apparently only affected the growth and survival of salmon fry. The regulating mechanisms involved in these inter- and intra-specific effects are discussed in terms of competition for stream resources and predation. Recommendations based on the findings are made for stocking and habitat management of salmonid waters.     

Polyunsaturated fatty acid profiles of whole body phospholipids and triacylglycerols in anadromous and landlocked Atlantic salmon (Salmo salar L.) fry

We compared the fatty acid compositions and gains of whole body triacylglycerols (TAG) and phospholipids (PL) in anadromous and landlocked Atlantic salmon (Salmo salar) fry, of the same age, fed the same commercial marine oil-rich diet over a 42-day feeding trial. The landlocked strain exhibited significantly (P<0.05) higher growth rate and feed efficiency, due principally to a higher fat retention, particularly of monounsaturated and saturated fatty acids (SFA). n-3 and n-6 long-chain polyunsaturated fatty acid (PUFA) gains and retentions were significantly higher (P<0.05) in the landlocked fry. Great similarities were found in the fatty acid profiles of whole body TAG of both strains. However, marked genotypic differences were observed in the PUFA profiles of whole body PL fractions. The total PUFA, n-3 PUFA and docosahexaenoic acid (DHA) level in PL was significantly higher (P<0.05) while the SFA level, and the PUFA C18/C20 and eicosapentaenoic acid/arachidonic acid ratios were significantly lower (P<0.05) in the anadromous fry than in landlocked fry. Our results indicate that the level of DHA in salmon PL is under strong genetic control and that the capacity for incorporation, and possibly for the conversion of dietary n-3 and n-6 PUFA, is higher in the landlocked strain.     

Selection on increased intrinsic growth rates in coho salmon, Oncorhynchus kisutch

Substantial evidence from the animal kingdom shows that there is a trade-off between benefits and costs associated with rapid somatic growth. One would therefore expect growth rates under natural conditions to be close to an evolutionary optimum. Nevertheless, natural selection in many salmonid species appears to be toward larger size and earlier emergence from spawning redds, indicating a potential for increased growth rate to evolve. We tested how selection for genetic variants (growth hormone transgenic coho salmon, Oncorhynchus kisutch, with more than doubled daily growth rate potential relative to wild genotypes) depended on predator timing and food abundance during the early period of life (fry stage). In artificial redds, fry of the fast-growing genotypes showed a highly significant developmental shift, emerging from gravel nests approximately two weeks sooner, but with an 18.6% reduced survival, relative to wild-genotype fry. In seminatural streams, fry of the fast-growing genotypes suffered higher predation than those of wild genotypes when predators were present at the time of fry emergence, but this difference was less pronounced when food was scarce. In streams where predators were introduced after emergence, fry survived equally well regardless of food availability. Surviving fry grew faster in habitats provided with more food, and fast-growing genotypes also grew faster than wild genotypes when predators arrived late and food was abundant. Fewer fish migrated downstream past a waterfall when food availability was high and in the presence of predators, and wild-genotype fry were more likely to migrate than fry of the fast-growing genotypes. After being returned to the experimental streams after migration, fast-growing genotypes survived equally well as those of the same genotypes that did not migrate, whereas migrating wild genotypes experienced higher mortality relative to those of the same genotypes that did not migrate. Comparisons of growth rates between siblings retained under hatchery conditions and those from habitats with the fastest growth in the experimental stream revealed that growth rates were similar for wild genotypes in both environments, whereas the fast-growing genotypes in the streams only realized 90% of their growth potential. The present study has shown that a major shift in developmental timing can alter critical early stages affecting survival and can have a significant effect on fitness. Furthermore, ecological conditions such as food abundance and predation pressure can strongly influence the potential for fast-growing variants to survive under natural conditions. The large-scale removal of many predatory species around the world may augment the evolution of increased intrinsic growth rates in some taxa.     

Evidence of differences in growth and food intake regulation in different genetic strains of channel catfish

Norris and USDA-103 strains of channel catfish Ictalurus punctatus were compared for growth rate and food conversion ratio under satiation feeding and restricted feeding (1% body weight day⁻¹) regimes. At the start of the experiment Norris fish weighed 2·8 g, USDA-103 fish weighed 14·0 g. Therefore, a regression of the loge of specific growth rate against the log_e of mean body size with an empirically derived fixed slope of -0·37 was used to compare growth rates. Under both feeding regimes the USDA-103 strain had faster specific growth rates and more efficient food conversion. In subsequent studies, voluntary food intake of size matched fish (60 g average) from these two strains was compared using a radiographic method. Fish were acclimatized to tank conditions for 3 weeks prior to voluntary food intake measurement. Half of the groups were deprived of food for 2 days prior to food intake measurement, while the remaining groups were fed 1% body weight day⁻¹. The USDA-103 strain fish ate significantly more food and grew faster than the Norris strain fish. Previously fasted Norris fish subsequently ate more than their fed counterparts, whereas the fed USDA-103 fish consumed more food than the fasted USDA-103 group. When the USDA-103 strain fish were deprived of food for 4 , 2 or 0 days, all groups subsequently consumed between 4·5 and 5·0% of body weight in one meal. The USDA-103 fish, unlike the Norris fish were not stimulated to consume more after short-duration fasting. Taken together, these results suggest that there are genetic differences in growth, food conversion ratio and regulation of food intake between Norris and USDA-103 strains.     

An experiment on faster growth of salmon Salmo salar (L.) in a Scottish stream

The experiment was made in an attempt to modify the usual relationship in which young trout grow faster than young salmon in streams in which they occur together. A stretch of a trout stream was stocked with advanced salmon eggs, which produced fry earlier than the trout eggs laid naturally. The salmon grew faster than the trout and were longer than the trout at the end of the growing season. The mean length of 77.7 mm attained by the salmon is the largest known size reached by salmon in their first year when feeding on natural food supplies in streams in Scotland. Survival rate from egg planting to production of salmon of this mean length was high.     

Experimental study on population dynamics in the guppy, Poeclia reticulata (Peters). Effect of shelters on the increase of population density

To study mass culture of the guppy, Poecilia reticulata (Peters) for use in mosquito control, the population dynamics of the fish were studied in 10001 polycarbonate tanks with and without shelters for fry, under conditions of constant temperature and filtration of water, and sufficient food supply. In the tank with shelters, the maximum density attained was 5.84 fish per 1 at the 77th week, when the initial population had been 25 males and 25 females. After this period the density began to decrease and dropped to 1.04 fish per 1 at the 105th week. Conversely, in the tank without shelters, almost no increase in number took place when the population started with either 50 adult fish or 25 fry. The sex ratio of the guppies in the 10001 tank with shelters varied. In particular, females increased in. relation to male when the fry which had been produced in large quantities became sexually mature. The intervals between spawning averaged 38 days with a range of 26 to 90 days. An equation, log Y =0.0443X-0.2968 expresses the relationship between the number of young in the brood and the body length of the females.     

Treatment with thiamine hydrochloride and astaxanthine for the prevention of yolk-sac mortality in Baltic salmon fry (M74 syndrome).

Two practical methods are reported for treating feral Baltic salmon with thiamine hydrochloride against M74 syndrome (abnormally high yolk-sac fry mortality of the Baltic salmon). Both bathing of the yolk-sac fry in thiamine hydrochloride (1000 mg l-1, 1 h) and a single intraperitoneal injection given to the female brood fish (100 mg kg-1 fish) during the summer 3 mo before stripping were shown to elevate the whole body total thiamine concentration in the fry. Both treatments were also shown to be effective in preventing mortality due to M74 syndrome. The effect of bathing the yolk-sac fry was shown to be dose-dependent. The results support the view that there is a causal relationship between the thiamine status of the yolk-sac fry and M74 mortality. An intraperitoneal injection of astaxanthine suspension administered to the female brood fish (11 mg kg-1 fish) in the summer 3 mo before stripping elevated the astaxanthine concentration in the eggs but did not affect mortality due to M74 syndrome. An interaction between astaxanthine and thiamine may occur in the developing embryo or yolk-sac fry, however. No association could be demonstrated between the various thiamine hydrochloride treatment practices and hepatic cytochrome P450 dependent 7-ethoxyresorufin-O-deethylase (EROD) activity in the yolk-sac fry. An injection of thiamine hydrochloride into the peritoneal cavity of wild Baltic salmon females could be used to raise thiamine concentrations in their offspring in the rivers. The effect on smolt production in Finnish Baltic salmon rivers needs to be investigated further, however.     

Water balance trumps ion balance for early marine survival of juvenile pink salmon (Oncorhynchus gorbuscha)

Smolting salmonids typically require weeks to months of physiological preparation in freshwater (FW) before entering seawater (SW). Remarkably, pink salmon (Oncorhynchus gorbuscha) enter SW directly following yolk absorption and gravel emergence at a size of 0.2 g. To survive this exceptional SW migration, pink salmon were hypothesized to develop hypo-osmoregulatory abilities prior to yolk absorption and emergence. To test this, alevins (pre-yolk absorption) and fry (post-yolk absorption) were transferred from FW in darkness to SW under simulated natural photoperiod (SNP). Ionoregulatory status was assessed at 0, 1 and 5 days post-transfer. SW alevins showed no evidence of hypo-osmoregulation, marked by significant water loss and no increase in gill Na?/K?-ATPase (NKA) activity or Na?:K?:2Cl? cotransporter (NKCC) immunoreactive (IR) cell frequency. Conversely, fry maintained water balance, upregulated gill NKA activity by 50 %, increased the NKA α1b/α1a mRNA expression ratio by sixfold and increased NKCC IR cell frequency. We also provide the first evidence of photoperiod-triggered smoltification in pink salmon, as fry exposed to SNP in FW exhibited preparatory changes in gill NKA activity and α1 subunit expression similar to fry exposed to SNP in SW. Interestingly, fry incurred larger increases in whole body Na? than alevins following both SW and FW + SNP exposure (40 and 20 % in fry vs. 0 % in alevins). The ability to incur and tolerate large ion loads may underlie a novel mechanism for maintaining water balance in SW prior to completing hypo-osmoregulatory development. We propose that pink salmon represent a new form of anadromy termed "precocious anadromy".     

Male reproductive suppression in the cooperatively breeding fish Neolamprologus pulcher

In most cooperative breeders, dominants suppress the reproductionof subordinates. However, two previous studies of Neolamprologuspulcher, a cooperatively breeding cichlid fish, have suggestedthat socially subordinate helper males sneak fertilizationsfrom dominant breeding males. If such sneaking does occur, boththeoretical work and empirical studies of other fish speciessuggest that sperm competition will select for increased reproductiveinvestment by sneaker males, relative to more dominant males.To address these issues, we quantified gonadal investment andsperm characteristics of 41 N. pulcher male breeders and 62male helpers from 55 groups in Lake Tanganyika. Gonadal investmentfollowed patterns consistent with reproductive suppression,with breeders having considerably larger testes masses thanhelpers. Breeders also had faster and longer swimming spermand a higher percentage of motile sperm compared to helpers.However, sperm characteristics of large helpers were similarto those of breeders, but these same helpers had lower testesmasses. Thus, large helpers had sperm that were physiologicallyequivalent to that of breeders, but their relatively small gonadsimply that they were reproductively suppressed.     

The relative influence of prior residency and dominance on the early feeding behaviour of juvenile Atlantic salmon

In many species where social hierarchy mediates conflict over resources, dominant individuals monopolize food, shelter and reproductive opportunities. The benefits of social dominance, however, can often be offset by a prior residence advantage, whereby individuals arriving first in a new habitat obtain, and subsequently defend, the most profitable sites. We investigated the relative influence of these two factors on the acquisition of feeding territories by juvenile Atlantic salmon,Salmo salar L., by placing groups of six individuals of known dominance rank sequentially into an experimental arena with feeding sites of varying quality. Dominants had an advantage over subordinates in the percentage of time spent in a good-quality feeding site and individual feeding rate. There was also a significant time of arrival effect: those individuals that arrived first in a habitat monopolized the resource. The two effects were of approximately equal strength, so that late-arriving dominant fish had similar success as prior resident subordinates. These results indicate that both dominance and prior residence are important in the feeding behaviour of juvenile Atlantic salmon, and that the two factors can have independent and additive effects.Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

Social dominance, prior residence and the acquisition of profitable feeding sites in juvenile Atlantic salmon

Twelve groups of 10 Atlantic salmon Salmo salar parr were screened for dominance in small tanks with a single localized food source using a serial removal method and the top, middle and bottom two fish in each group were classified as high, medium and low social status, respectively. These 72 ranked fish were weighed and formed into four groups or waves, each consisting of six fish of each status category. The fish were given dye marks according to status and wave and were then introduced into an artificial stream in four waves over a 12-day period. Patterns of settlement/emigration were recorded over a 2-month period, at the end of which the fish were removed from the stream, identified and reweighed, and daily specific growth rates calculated. Fifty-one fish established feeding stations in the flume, the remainder emigrating via a downstream trap. The average growth rate over the 2-month period was 0·87% day⁻¹. Probability of settlement and growth rates following settlement were significantly higher in fish from the earlier waves. In contrast, salmon that were dominant in the initial trial were less likely than subordinate fish to settle in the experimental stream and showed lower postsettlement growth rates. The possibility that fish have individually consistent behavioural profiles that promote success in different competitive environments is discussed with reference to published literature for other groups. A small-scale trial at the end of the main experiment confirmed the prior residence effect and demonstrated that 1 day was sufficient for such an effect to be evident.