Differences in susceptibility of anadromous and resident stocks of Arctic charr to infections of Gyrodactylus salaris, under experimental conditions

The ability of Gyrodactylus salaris , an important pathogen of the Atlantic salmon Salmo salar , in Norway, to infect anadromous and resident stocks of the Arctic charr, Salvelinus alpinus , has been examined in the laboratory. Resident charr (Korssjoen stock) exposed to heavily infected salmon, were considered innately resistant as they lost their infections within 21 days when individually isolated. Isolated anadromous harr (Hammerfest stock) remained infected for up to 150 days, although most infections disappeared within 30–50 days. In many cases the parasite population grew initially, but growth was limited after 20–30 days and infections subsequently disappeared. At the same time, shoals of 50 anadromous charr, swimming in the tanks containing the individually isolated fish in floating cages, remained infected for up to 280 days. Charr isolated from these shoals after 115 days and subsequently monitored individually lost their infections within 30 days, although the parasite persisted within the shoals for a further 75–135 days. This suggests that G. salaris , persisted on shoaling charr despite an immune response which led to the elimination of parasites from isolated hosts. The Hammerfest stock of anadromous charr supports G. salaris , in the laboratory, and the extended period of survival on this host suggests that charr may be important in the epidemiology of G. salaris , in northern Norway.     

Differences in the host resistance of Atlantic salmon, Salmo salar L., stocks to the monogenean Gyrodactylussalaris Malmberg, 1957

The susceptibility and resistance of hatchery-reared salmon parr, native to the rivers Neva (U.S.S.R. Baltic Sea), Alta (northern Norway) and Lone (western Norway) (both eastern Atlantic Ocean), to Gyrodactylus salaris from Norway, was examined. The level of resistance to the parasite was assessed from counts, made on anaesthetized salmon, ofthe numbers of G. salaris after an initial experimental exposure (2 weeks) to G. salaris-infected salmon. Three experiments, all in water at c. 12° C, were carried out: (1) 50 Alta and 50 Neva salmon, initial mean parasite intensity c. 12; (2) 50 Lone and 50 Neva salmon, initial mean parasite intensity c. 60; (3) 10 Lone and 10 Neva salmon individually isolated, initial intensity one gravid G. salaris . In both the Norwegian salmon stocks, the G. salaris infrapopulations steadily increased during the experimental period of 5 weeks, in contrast to a prominent decline in the Neva salmon stock, after, respectively: (Exp. 1) week 3, average peak intensity 32.6; (Exp. 2) week 2, average peak intensity 58.7; and (Exp. 3) week 3, average peak intensity 6.3. The hatchery-reared Baltic Neva stock demonstrated both an innate and an acquired resistance towards G. salaris , in contrast to the highly susceptible, Norwegian Alta and Lone salmon stocks.     

The susceptibility of grayling (Thymallus thymallus) to experimental infections with the monogenean Gyrodactylus salaris

The pathogenic monogenean Gyrodactylus salaris infecting Atlantic salmon (Salmo salar) is found to attach and reproduce under laboratory conditions on several species in the subfamily Salmoninae other than the Atlantic salmon. The gyrodactylid species Gyrodactylus thymalli infecting grayling (Thymallus thymallus) in another subfamily, Thymallinae, is previously said to be very similar to G. salaris based on morphometry and genetical analysis which prompted the present laboratory experiments to test the susceptibility and resistance of grayling to G. salaris. All 0+ and 1+ grayling became infected with G. salaris during the experimental infection procedure. However, both innate resistant and susceptible grayling were found. In susceptible individually isolated fish, parasite reproduction lasted for more than 35 days. Parasite reproduction also occurred among grouped grayling as judged from the duration of infection of more than 50 days. However, grayling susceptibility as judged from G. salaris reproduction, was very limited. Hence, the results indicate significant biological differences between the function of Atlantic salmon and grayling as host for G. salaris. The grayling is interpreted as unable to sustain G. salaris in nature which implies that G. thymalli is not conspecific with G. salaris. However, G. salaris dispersal by grayling cannot be excluded.     

The host specificity of Gyrodactylus salaris Malmberg (Platyhelminthes, Monogenea): susceptibility of Oncovhynchus mykiss (Walbaum) under experimental conditions

An experimental epidemiological approach was chosen to study the survival and infection dynamics of Gyrodactylus salaris on juvenile rainbow trout, Oncorhynchus mykiss , in the laboratory. A marked heterogeneity in the host stock was apparent. The rainbow trout could be divided into three groups on the basis of parasite survival and infection pattern on individually isolated fish: (1) hosts receptive to initial parasite attachment, but unreceptive to parasite establishment and reproduction; (2) hosts moderately susceptible to parasite establishment and reproduction, but which, after a period of restricted parasite population growth, responded, recovered and eliminated the parasites; and (3) hosts very susceptible to parasite infection and reproduction, but which, after a period of significant parasite population growth, responded, recovered and eliminated the parasites. These different patterns are considered to reflect genetic differences between host individuals. Parasite aggregation was also shown to be an important factor in the outcome of the host-parasite association. The parasites were finally eliminated on the individually isolated hosts, but not on hosts maintained in batches and so host population size and immigration of fresh. previously unexposed, hosts appeared to be important for growth and maintenance of the parasite population. The parasite was not found to cause host mortality. Rainbow trout was a suitable host for G. salaris , capable of transmitting the parasite to new localities as a consequence of stocking programmes or migratory behaviour.     

Susceptibility of Baltic and East Atlantic salmon Salmo salar stocks to Gyrodactylus salaris (Monogenea)

The susceptibility of a Baltic salmon stock Salmo salar (Indals?lv, central Sweden) to Norwegian Gyrodactylus salaris (Figga strain, central Norway) was experimentally tested and compared with previously obtained results on East Atlantic salmon (Lierelva, SE Norway). Contrary to expectation, the Baltic salmon, which had no prior exposure to this parasite strain, appeared almost as susceptible as the Norwegian salmon parr that naturally experience G. salaris-induced mortality. Individually isolated salmon of both stocks sustained G. salaris infections with little evidence of innate resistance. A few individuals of the Indals?lv stock controlled their infection from the beginning, but overall there was considerable heterogeneity in the course of infection in both stocks. On individual hosts, G. salaris growth rates declined steadily throughout the infection, a trend which was particularly marked amongst the Lierelva stock. On shoaling Lierelva fish, there was some evidence of reduced parasite population growth towards the end of the infection; this was not apparent in Indals?lv fishes. These results reflect a growing awareness that not all Baltic salmon may be resistant to Norwegian G. salaris, and that Norwegian and Baltic G. salaris strains may differ in virulence. Consequently, management decisions concerning this parasite-host system should be based upon the actual, and tested, susceptibility of stocks under consideration and not upon identification of stocks as either Atlantic or Baltic.     

The susceptibility of Salvelinus fontinalis (Mitchill) to Gyrodactylus salaris Malmberg (Platyhelminthes; Monogenea) under experimental conditions

The host specificity of Gyrodaclylus Solaris is examined experimentally with respect to its ability to infect the brook trout, Salvelinus fontinalis . The parasite readily attached to and reproduced on parr of this host and infections grew for c. 20 days from first monitoring (c. 30 days from first infection) before declining. Parasites could persist on this host for up to 70 days before finally disappearing. The pattern of infection resembled that seen in many other gyrodactylid species on their normal hosts, and suggested the action of a host response, In this respect infections of G. salaris on parr of S. fontinalis , anadromous Salvelinus alpinus, Oncorhynchus mykiss, Thymallus thymallus and Baltic Salmo salar follow a normal pattern, while infections of Norwegian S. salar are unusual in a continued unchecked growth, until the host dies, under pooled laboratory conditions.     

Characterisation of a low pathogenic form of Gyrodactylus salaris from rainbow trout

Gyrodactylus salaris was isolated from rainbow trout in a Danish freshwater trout farm, and a laboratory population of this particular parasite form was established on rainbow trout. Challenge infections were performed using different salmonid strains and species, including East Atlantic salmon Salmo salar (from the Danish River Skjern?), Baltic salmon S. salar (from the Swedish River Ume Alv) and rainbow trout Oncorhynchus mykiss (from the Danish rainbow trout farm Fousing). These were compared to infection studies on the Norwegian Laerdalselva parasite form kept under exactly the same conditions in the laboratory. The Danish G. salaris form had low virulence towards both Atlantic and Baltic salmon, whereas rainbow trout proved susceptible to the parasite. The Danish G. salaris form was able to maintain a very low infection on East Atlantic salmon, but not on the Baltic salmon, which eliminated the infection within 2 wk. Rainbow trout developed infection intensities ranging up to several hundred parasites per host. The host colonization patterns of the parasite differed clearly from those of previous studies on microhabitats of the Norwegian form of G. salaris. A comparative study on morphological characters (opisthaptoral hard parts) from the Danish parasite form and Norwegian G. salaris showed no significant differences. Selected genes comprising internal transcribed spacers 1 and 2 (ITS), ribosomal RNA intergenic spacer (IGS) and cytochrome c oxidase subunit I (COI) regions were cloned and sequenced. Five sequenced ITS clones from 5 individuals of the Danish strain consistently revealed a single base substitution compared to ITS sequences from all other known species and strains of Gyrodactylus. Mitochondrial COI gene sequences demonstrated that the Danish G. salaris form is closely similar to the Laerdalselva parasite form found in Norway. The IGS sequences were highly variable, but very similar to those obtained from German isolates of G. salaris.     

Identification of genetic markers associated with Gyrodactylus salaris resistance in Atlantic salmon Salmo salar

Gyrodactylus salaris Malmberg, 1957 is a freshwater monogenean ectoparasite of salmonids, first recorded in Norway in 1975 and responsible for extensive epizootics in wild Atlantic salmon Salmo salar L. The susceptibility of different populations of Atlantic salmon to G. salaris infection differs markedly, with fish from the Baltic being characterised as relatively resistant whereas those from Norway or Scotland are known to be (extremely) susceptible. Resistance to Gyrodactylus infection in salmonids has been found to be heritable and a polygenic mechanism of control has been hypothesised. The current study utilises a 'Quantitative trait loci' (QTL) screening approach in order to identify molecular markers linked to QTL influencing G. salaris resistance in B1 backcrosses of Baltic and Scottish salmon. Infection patterns in these fish exhibited 3 distinct types; susceptible (exponential parasite growth), responding (parasite load builds before dropping) and resistant (parasite load never increases). B1 backcross fish were screened at 39 microsatellite markers and single marker-trait associations were examined using general linear modelling. We identified 10 genomic regions associated with heterogeneity in both innate and acquired resistance, explaining up to 27.3% of the total variation in parasite loads. We found that both innate and acquired parasite resistance in Atlantic salmon are under polygenic control, and that salmon would be well suited to a selection programme designed to quickly increase resistance to G. salaris in wild or farmed stocks.     

Brown trout Salmo trutta released to support recreational fishing in a Norwegian fjord

One- and two-year-old hatchery reared juveniles of seven freshwater resident and anadromous populations of Scandinavian brown trout were released in the outer and inner part of the Oslofjord and in the River Akerselva, flowing through the city of Oslo. Recapture rates were highest (mean 20.3%) for river released fish and lowest for those released in the outer (16.8%) and inner (12.1%) fjord. In general, recapture rate increased with fish size at release (r=0.76). When released in fresh water, most of the recaptures were from fresh water and when released in the fjord, most recaptures were from the fjord. In general, freshwater resident stocks showed a higher degree of freshwater residency than anadromous stocks. However, mean migratory distance was longer for freshwater resident than anadromous fish. Trout moved longer distances at sea when released in the outer than in the inner fjord. Specific growth rate and size at recapture varied among release sites and stocks; they were highest for fish released in the outer fjord and lowest for river-released trout. There was no consistent difference in sea growth between freshwater resident and anadromous stocks. Estimated total yield was highest for fish released in the outer fjord, whereas there was no significant difference in yields between trout released in the river and the inner Oslofjord.     

Molecular faunistics of accidental infections of <Emphasis Type="Italic">Gyrodactylus</Emphasis> Nordmann, 1832 (Monogenea) parasitic on salmon <Emphasis Type="Italic">Salmo salar</Emphasis> L. and brown trout <Emphasis Type="Italic">Salmo trutta</Emphasis> L. in NW Russia

Salmon Salmo salar L. and brown trout S. trutta L. juveniles were examined for the presence of accidental monogenean ectoparasitic species of Gyrodactylus Nordmann, 1832 in the Baltic and White Sea basins of Russian Karelia in order to estimate the frequency of host-switching attempts on an ecological timescale. To collect phylogeographical information and for exact species identification, the parasites were characterised by nuclear internal transcribed spacer sequences of rDNA (ITS) and, for some species, also by their mitochondrial DNA (CO1 gene) sequences. Four accidental Gyrodactylus species were observed on salmon and brown trout. A few specimens of G. aphyae Malmberg, 1957, the normal host of which is the Eurasian minnow Phoxinus phoxinus (L.), were observed on lake salmon from the Rivers Kurzhma (Lake Kuito, White Sea basin) and Vidlitsa (Lake Ladoga, Baltic basin). G. lucii Kulakovskaya, 1952, a parasite of the northern pike Esox lucius L., was observed on salmon in the Kurzhma. In the River Vidlitsa, two specimens of G. papernai Ergens & Bychowsky, 1967, normally on stone loach Barbatula barbatula (L.), were found on salmon. On anadromous White Sea salmon in the River Pulonga in Chupa Bay, a few salmon parr carried small colonies of G. arcuatus Bychowsky, 1933, which were shown to have originated from the local three-spined stickleback Gasterosteus aculeatus L. consumed as prey. No specimens of Gyrodactylus salaris Malmberg, 1957 were observed, although the Pulonga is the nearest salmon spawning river to the River Keret', which is heavily infected with introduced G. salaris. In the River Satulinoja, Lake Ladoga, three specimens of G. lotae Gusev, 1953, from burbot Lota lota (L.), were collected from a single brown trout S. trutta. All nonspecific gyrodactylid infections on salmonids were judged to be temporary, because only a few specimens were observed on each of the small number of infected fishes. The prevalence of endemic G. salaris was also low, only 1% (Nfish = 296) in Lake Onega and 0.7% (Nfish = 255) in Lake Ladoga, while brown trout specific Gyrodactylus species were not observed on any of the 429 trout examined from the Ladoga basin. The host-specific and unspecific burden of Gyrodactylus spp. on these 'glacial relict' populations of salmon and brown trout was very low, suggesting a generalised resistance against the co-evolved freshwater parasite community, or some kind of 'vaccination' effect. These hypotheses deserve further testing.     

Assessing the reproductive contributions of sympatric anadromous and freshwater‐resident brook trout

The δ¹³C, δ¹⁵N and δ³⁴S isotope values of newly emerged, brook trout Salvelinus fontinalis alevins (free‐swimming individuals beginning exogenous feeding) were examined to determine if progeny of anadromous female spawners could be detected and their contributions to reproduction assessed in river systems with mixed migration strategies. Separation of anadromous and freshwater resident and immigrant sources of progeny could be detected primarily using δ¹³C values before alevins reached a size of 28 mm fork length and c . 20 days after exogenous feeding began. Among 10 populations, the contributions of anadromous females ranged from 0 to 42% of newly emerged alevins. Correlations between anadromous contributions and estimates of total brook trout density and alevin production were unexpectedly negative.     

Effects of seawater-acclimatization and release sites on survival of hatchery–reared brown trout Salmo trutta

To test whether seawater–acclimatization of hatchery–reared anadromous and freshwater resident brown trout before release increased the survival of adults, smolts were retained 0, 2, 4 and 8 weeks in sea water before release. Total recapture rate increased for smolts retained 4 and 8 weeks in sea water before release relative to the controls. This trend was more pronounced for Freshwater resident than for anadromous stocks, Offspring of anadromous fish stayed longer at sea than offspring of freshwater resident fish. Recapture rates in fresh water were higher for brown trout released in the river than in the fjord in the R, Drammen area, but not in the R. Imsa. In both cases, most fish were recaptured in the sea. Moving into the R. Imsa (relative to other rivers) appeared higher for fish released at the mouth of the river (93%) than in the fjord (47%). Judged from the recapture rates, sea survival appeared to be the same whether released in the fjord or at the mouth of the river.     

Genetic population structure of Norwegian brown trout

Biochemical genetic variation in populations of anadromous and resident brown trout, Salmo trutta L., was studied. Altogether 50 Norwegian populations were screened for 32 enzyme loci. Genetic polymorphism was found at the following 11 loci: AAT-4 * (E.C. 2.6.1.1), CK-1 * (E.C. 2.7.3.2), G3PDH-2 * (E.C. 1.1.1.8), IDHP-2 * (E.C. 1.1.1.42), LDH-5 * (E.C. 1.1.1.27), MDH-2 * (E.C. 1.1.1.37), MDH-3/4 * (E.C. 1.1.1.37), MEP-2 * (E.C. 1.1.1.40), GPI-2 * (E.C. 5.3.1.9). GPI-5 * (E.C. 5.3.1.9) and PGM-1 * (E.C. 5.4.2.2), giving an overall polymorphism of 34%, ranging from 3.7 to 29.6% among individual populations. The average calculated heterozygosity ranged from 1.4 to 10.2% among populations. Genetic heterogeneity was observed among anadromous populations, and significant differences in allelic frequencies were found between anadromous populations in neighbouring watercourses, among resident populations and between anadromous and resident populations inhabiting the same watercourses. Significant heterogeneity was also found among 12 populations from Lake Mjøsa, with a major division between the western and eastern populations of the lake. Differences in allelic frequencies were found between wild stocks and their hatchery derivatives, and between different hatchery derivatives originating from the same wild population. In some cases release of hatchery populations into wild stocks may have influenced the genetic characteristics of wild stocks. The data support the hypothesis of eastern as well as western postglacial colonization lines for Norwegian brown trout.     

Host specificity dynamics: observations on gyrodactylid monogeneans

The directly transmitted viviparous gyrodactylids have high species richness but low morphological and biological diversity, and many species are recorded from only a single host. They therefore constitute a guild of species ideal for studies of the evolutionary significance of host specificity. The group has the widest host range of any monogenean family, being found on 19 orders of bony fish. However, individual species range from narrowly specific (71% of 402 described species recorded from a single host) to extremely catholic (Gyrodactylus alviga recorded from 16 hosts). Gyrodactylid-host interactions extend from 60 mya (G. lotae, G. lucii) down to 150 years (G. derjavini on Oncorhynchus mykiss). Co-evolution with the host is comparatively rare within the gyrodactylids, but host switching or ecological transfer is common, and has been facilitated by the mixing of fish strains that followed glaciation. In this review, we consider the factors responsible for gyrodactylid specificity patterns, using examples from our work on salmonid gyrodactylids including G. salaris, responsible for major epidemics on wild Atlantic salmon (Salmo salar) in Norway since 1975, and G. thymalli from grayling and G. derjavini from trout.G. salaris has a wide host range with highest population growth rates on Norwegian salmon strains. However, growth rates are variable on both host strains and species, because of the multitude of micro- and macro-environmental factors influencing parasite mortality and fecundity. A better predictor of performance is the proportion of fishes of a strain which are innately resistant to the parasite, a measure which is negatively correlated with the time to peak infection in a host strain. Population growth rate is also negatively correlated with age of infection; the initial rate, therefore, predicts best the suitability of a fish as host for G. salaris. The host response to gyrodactylids appears to be the same mechanism in all salmonids with innate resistance as one end of a spectrum, but influenced by stress and probably under polygenic control. Hybrid experiments show that performance of G. salaris on a host is heritable, and usually intermediate between that of the parents. This host response mechanism, coupled with the initial parasite population growth on a fish, determines the host specificity, i.e. whether the fish will be susceptible, a responder or innately resistant. The use of population growth rate parameters allows comparison of different hosts as a resource for a gyrodactylid. In the case of G. salaris, East Atlantic and Baltic strains of Atlantic salmon are core hosts, but other salmonids can physiologically sustain infections for considerable periods, and may be important in parasite dispersal and transmission. A further group of non-salmonid fishes are unable to sustain G. salaris reproduction, but can act as transport hosts.Population growth parameters are very labile to stressors and environmental factors, particularly temperature and salinity, and also other aspects of host ecology and water quality. These factors may also influence the spectrum of hosts that can be infected under particular conditions, and probably favoured ecological transfer of gyrodactylids between host species in periglacial conditions. G. salaris may still be undergoing post-glacial range expansion (aided by anthropogenic spread) as shown by the increase in the species range over the last 25 years. The origin of G. salaris, G. teuchis and G. thymalli is discussed in relation to glacial refugiums during the last ice age.     

Analysis of ribosomal RNA intergenic spacer (IGS) sequences in species and populations of Gyrodactylus (Platyhelminthes: Monogenea) from salmonid fish in Northern Europe

The intergenic spacer (IGS) region of ribosomal RNA genes was amplified and sequenced from a variety of Gyrodactylus specimens collected from wild and farmed Atlantic salmon Salmo salar, rainbow trout Oncorhynchus mykiss, and grayling Thymallus thymallus, from various locations in Northern Europe. Phylogenetic analysis of the sequences confirmed the distinction between G. salaris Malmberg, 1957 and G. thymalli Zitnan, 1960, supporting their validity as separate species. G. salaris adapted to rainbow trout are also distinct from the parasites found on Atlantic salmon, supporting the existence of a rainbow-trout form that was initially identified on the basis of morphological differences. Analysis of the IGS did not provide good resolution of different populations of G. salaris sensu stricto, but was consistent with epidemiological evidence which indicates that introduction of the parasite to Norway was recent and limited. The IGS may be helpful in distinguishing forms of G. salaris that are pathogenic to Atlantic salmon from those that are not.     

Genetic architecture of growth and early life‐history transitions in anadromous and derived freshwater populations of steelhead

Heritabilities of growth, precocious maturation and smolting were measured in 75 families of juvenile steelhead or rainbow trout Oncorhynchus mykiss , progeny of within and between line matings (crosses) of wild, anadromous steelhead and wild, resident (lake) rainbow trout originally derived from the same anadromous stock 70 years earlier. The tagged yearling progeny were combined by line in common freshwater rearing containers and graded into three categories: mature, smolt or rearing (undifferentiated) at age 2 years. Heritabilities of precocious male maturity, smolting and growth were moderate to high, and the genetic correlation between growth and smolting was low. Smolting and precocious male maturity were highly variable among families within lines and significantly different between lines. Each of the four lines produced significant numbers of smolts at age two. Smolting and maturation were negatively genetically correlated, which may explain the persistence of smolting in the lake population despite strong selection against lake smolts; balancing selection on male maturation age may help to maintain variation for smolting. The high heritability of smolting, coupled with the inability of smolts that leave the lake to return to it indicates that the genetic potential for smolting can lie dormant or be maintained through a dynamic interaction between smolting and early maturation for decades despite complete selection against the phenotype. The results have significant implications for the preservation of threatened anadromous stocks in fresh water and the inclusion of resident fish of formerly anadromous populations, currently trapped behind long‐standing barriers to migration, as one component of the same population.     

Demonstrating freedom from Gyrodactylus salaris (Monogenea: Gyrodactylidae) in farmed rainbow trout Oncorhynchus mykiss

This paper describes an approach to demonstrate freedom of individual rainbow trout farms from Gyrodactylus salaris Malmberg, 1957. The infection status of individual farms is relevant should G. salaris be introduced into a country or zone previously known to be free of the parasite. Trade from farms where G. salaris may have been introduced would be restricted until freedom had been demonstrated. Cage, fish and parasite sample sizes were calculated based on the minimum detectable prevalence (P*), test characteristics, population size, and Type I and II errors. Between 5 and 23 cages per farm would need to be sampled to demonstrate freedom at a cage level P* of 10%. The number of fish sampled per cage depended mainly on the test sensitivity (probability of correctly identifying an infected fish). Assuming a test sensitivity of 99% at the fish level, 59 fish per cage are needed (P* = 5%). Since G. salaris may exist in mixed infection with G. derjavini, testing a sample of gyrodactylid parasites may not result in the parasite being detected when present. Test sensitivity at the fish level depends on the number of gyrodactylids on the fish, the proportion of which are G. salaris and the number examined. Assuming a P* of 5% (i.e. G. salaris are at least 5% of the gyrodactylid population), between 20 and 73 parasites per fish would need to be sampled (depending on abundance) to maintain the Type I error at 0.01 (thus a fish level test sensitivity of 99%). This work identifies the critical information, and further research, needed to assess freedom from G. salaris with a known level of confidence; this is essential to provide a sound scientific basis for decision-making about disease control measures.     

A small number of anadromous females drive reproduction in a brown trout (Salmo trutta) population in an English chalk stream

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Genetic differentiation between freshwater resident and anadromous brown trout, Salmo trutta, within watercourses

Freshwater resident and anadromous life history forms of brown trout, Salmo trutta L., sampled from each of three watercourses were analysed for biochemical genetic variation. Ten enzyme systems encoded by a putative 30 loci were analysed. Significant genetic differences were found between the two life history forms in all three watercourses. Further, significant genetic differences were also found between the three anadromous populations analysed, although two of them are separated by a distance along the shore line by less than 2 km. The findings focus on the need to reconsider the conservation and management practice of wild and hatchery-reared stocks.     

Life history response and age-specific tolerance to starvation in Brachionus plicatilis O.F. Müller (Rotifera)

To examine the life history response and age-specific tolerance to starvation in the rotifer Brachionus plicatilis O.F. Müller, we carried out two series of individual culture experiments. In the first experiment, rotifers were fed until each of the ages of 1-4 days, and were then starved during the rest of their lifetimes. The control group was fed throughout their lifespans. Rotifers stopped active reproduction just after the onset of food deprivation, and showed shorter subsequent survival times when they were starved at older ages. The finding that the larger the number of offspring produced before food deprivation, the shorter the subsequent lifetime under starvation, appeared to reflect a trade-off with the cost of reproduction. In the second experiment, newborns were starved until each of the ages of 1-5 days, and were fed thereafter. The lifespans of the rotifers starved up to the age of 3 days were not statistically different from those that were not starved. Although the starved rotifers began to reproduce once fed again, their lifetime fecundity decreased significantly from that of the non-starved group. Based on these results, it was suggested that the reproductive suppression caused by starvation would cause rotifers to have a longer lifespan to allow for future reproduction.