Comparative anatomy of the dorsal hump in mature Pacific salmon

Mature male Pacific salmon (Genus Oncorhynchus ) demonstrate prominent morphological changes, such as the development of a dorsal hump. The degree of dorsal hump formation depends on the species in Pacific salmon. It is generally accepted that mature males of sockeye (O. nerka ) and pink (O. gorbuscha ) salmon develop most pronounced dorsal humps. The internal structure of the dorsal hump in pink salmon has been confirmed in detail. In this study, the dorsal hump morphologies were analyzed in four Pacific salmon species inhabiting Japan, masu (O. masou ), sockeye, chum (O. keta ), and pink salmon. The internal structure of the dorsal humps also depended on the species; sockeye and pink salmon showed conspicuous development of connective tissue and growth of bone tissues in the dorsal tissues. Masu and chum salmon exhibited less‐pronounced increases in connective tissues and bone growth. Hyaluronic acid was clearly detected in dorsal hump connective tissue by histochemistry, except for in masu salmon. The lipid content in dorsal hump connective tissue was richer in masu and chum salmon than in sockeye and pink salmon. These results revealed that the patterns of dorsal hump formation differed among species, and especially sockeye and pink salmon develop pronounced dorsal humps through both increases in the amount of connective tissue and the growth of bone tissues. In contrast, masu and chum salmon develop their dorsal humps by the growth of bone tissues, rather than the development of connective tissue.     

Ecological model of interactions between escaped and wild Atlantic salmon Salmo salar

Atlantic salmon in Maine were once abundant but have become depleted, and are listed as endangered under the federal Endangered Species Act. Historically, salmon numbers in Maine may have been as high as 100 000 adults, but habitat loss, pollution and overfishing have contributed to the decline of the species. In 2000, only 110 adults returned to spawn in Maine rivers. Maine produces c. 15 000 metric tons/year of aquacultured Atlantic salmon from a total of nearly 600 coastal net pens. Escapees from these pens may interact with the wild salmon. The dynamics of salmon populations under such conditions are poorly understood. In order to illuminate the role aquaculture may play in such a system, we have developed a model for simulating population trajectories for both wild salmon and competing populations derived from aquaculture escapes. The model simulates a small population of wild salmon based in a stream/estuary system, into which an aquaculture facility is losing fish to escapes. Biological parameters in the model were estimated as much as possible from data in the USFWS report on Maine salmon. We used the model to investigate the consequences of a variety of ecological interactions between the wild and cultured fish including competitive, genetic and disease effects. Initial results indicate that many of these effects allow the aquaculture‐derived population to supplant the wild fish, but that wild populations may still persist under some conditions.     

Chromosome size in salmon and trout

The chromosomes of the Atlantic salmon, Salmo salar (2n=58) are, on average, larger than those of the trout, S. trutta (2n=80). If the difference in chromosome size represents a permanent change in chromosome structure as between the two species the expectation is that the size difference between salmon and trout chromosomes will be maintained in the hybrid. If, alternatively, the size difference between salmon and trout chromosomes is genotypically determined the difference will not be maintained in nuclei of hybrid genotype. Measurements of a specific chromosome, S, of the salmon complement and of another, S ¹, of the trout complement in nuclei of parent species and of the hybrid show that the difference in size is maintained in hybrid nuclei. It is concluded therefore that the size difference between salmon and trout chromosomes is due to structural change rather than to genotypic control.     

Comparative aspects of infectious salmon anemia virus,an orthomyxovirus of fish,to influenza viruses

Infectious salmon anaemia (ISA) is a viral disease that was first recorded in 1984 in farmed Atlantic salmon. The infectious salmon anaemia virus (ISAV) is classified as the type species of the genus Isavirus in the Orthomyxoviridae family and is evolutionary remote to the influenza viruses. The genome consists of eight negative single-stranded RNA segments, and it utilises the same mechanisms as influenza viruses to enter and exit cells. Although a common ancestor of ISAV and other genera of Orthomyxoviruses could be dated back several millions of years, there are still many similarities between ISAV and the influenza viruses regarding morphology, replication cycles and interactions with their respective hosts.     

Adult spawners: A critical period for subarctic Chinook salmon in a changing climate

Concurrent, distribution-wide abundance declines of some Pacific salmon species, including Chinook salmon (Oncorhynchus tshawytscha), highlights the need to understand how vulnerability at different life stages to climate stressors affects population dynamics and fisheries sustainability. Yukon River Chinook salmon stocks are among the largest subarctic populations, near the northernmost extent of the species range. Existing research suggests that Yukon River Chinook salmon population dynamics are largely driven by factors occurring between the adult spawner life stage and their offspring's first summer at sea (second year post-hatching). However, specific mechanisms sustaining chronic poor productivity are unknown, and there is a tremendous sense of urgency to understand causes, as declines of these stocks have taken a serious toll on commercial, recreational, and indigenous subsistence fisheries. Therefore, we leveraged multiple existing datasets spanning parent and juvenile stages of life history in freshwater and marine habitats. We analyzed environmental data in association with the production of offspring that survive to the marine juvenile stage (juveniles per spawner). These analyses suggest more than 45% of the variability in the production of juvenile Chinook salmon is associated with river temperatures or water discharge levels during the parent spawning migration. Over the past two decades, parents that experienced warmer water temperatures and lower discharge in the mainstem Yukon River produced fewer juveniles per spawning adult. We propose the adult spawner life stage as a critical period regulating population dynamics. We also propose a conceptual model that can explain associations between population dynamics and climate stressors using independent data focused on marine nutrition and freshwater heat stress. It is sobering to consider that some of the northernmost Pacific salmon habitats may already be unfavorable to these cold-water species. Our findings have immediate implications, given the common assumption that northern ranges of Pacific salmon offer refugia from climate stressors.     

Amino acid cues emanating from Pacific salmon eggs and ovarian fluid

The eggs of salmonid fishes are an important food source for many aquatic predators that detect eggs using olfaction. Moreover, chemicals from eggs and ovarian fluid aid sperm cells in detecting and locating eggs for fertilization, and ovarian fluid is attractive to conspecific males. Thus chemicals from eggs and ovarian fluid may facilitate reproduction but may also attract egg predators. The authors sampled mature females of three Pacific salmon species – Chinook (Oncorhynchus tshawytscha), coho (Oncorhynchus kisutch) and sockeye (Oncorhynchus nerka) – and determined the proportional representation of amino acids, potent fish odorants, from their eggs and ovarian fluid (Chinook and coho salmon only). They then tested juvenile coho salmon, an egg predator, for responses to ovarian fluid and egg odours using the electro-olfactogram (EOG) recording technique. The amino acid compositions of the salmon species were significantly and positively correlated with each other, and the interspecific differences were comparable to those between individuals of the same species. The egg water samples were, on average, dominated by lysine, alanine and glutamine (12.6%, 12.4% and 10.9%, respectively). The ovarian fluid samples were dominated by lysine (20.5%), followed by threonine (9.7%), glycine (9.2%) and arginine (8.8%). EOG recordings demonstrated the ability of juvenile coho salmon to detect the chemical traces of eggs and ovarian fluid. It is concluded that salmon eggs are a potent source of odours for potential predators but likely not highly differentiated among salmon species.     

Reproductive isolation following reintroduction of Chinook salmon with alternative life histories

We evaluated reproductive isolation of Chinook salmon (Oncorhynchus tshawytscha) life history types that have been reintroduced to northern Idaho, USA. Analysis of 1003 samples at six microsatellite loci revealed strong reproductive isolation between ocean- and stream-type Chinook salmon (fall and summer spawn timing, respectively) within the Clearwater River sub-basin (F _ST = 0.148, P < 0.00001). Very little evidence for gene flow among the two life history types was observed as assignment tests correctly assigned 99.6% of individuals in reference collections to either ocean- and steam-type Chinook salmon. Assignment of naturally reared juveniles indicated that both life history types were present with 24.1% stream-type and 75.9% ocean-type. Previous studies suggest high levels of divergence among the two life history types in natural populations, and our study verifies the persistence of reproductive isolation among types following colonization of habitat. Successful colonization of new habitat by (re)introduced species is likely influenced by diversity in life history types and this strategy has lead to naturally spawning populations in a variety of available habitats in the Clearwater River. As many populations of O. tshawytscha are listed as threatened or endangered under the U.S. Endangered Species Act, hope for recovery lies not only in effective management and habitat improvement, but adaptability of this species.     

Sequence analysis of a polymorphic Mhc class II gene in Pacific salmon

Polymorphism of the nucleotide sequences encoding 149 amino acids of linked major histocompatibility complex (Mhc) class II 131 and 132 peptides, and of the intervening intron (548–773 base pairs), was examined within and among seven Pacific salmon (Oncorhynchus) species. Levels of nucleotide diversity were higher for theB1 sequence than forB2 or the intron in comparisons both within and between species. For the codons of the peptide binding region of the BI sequence, the level of nonsynonymous nucleotide substitution (d_N) exceeded the level of synonymous substitution (d_S) by a factor of ten for within-species comparisons, and by a factor of four for between-species comparisons. The excess of d_N indicates that balancing selection maintains diversity at this salmonidMhc class II locus, as is common forMhc loci in other vertebrates. Levels of nucleotide diversity for both the exon and intron sequences were greater among than within species, and there were numerous species-specific nucleotides present in both the coding and noncoding regions. Thus, neighbor-joining analysis of both the intron and exon regions provided phylogenies in which the sequences clustered strongly by species. There was little evidence of shared ancestral (trans-species) polymorphism in the exon phylogeny, and the intron phylogeny depicted standard relationships among the Pacific salmon species. The lack of shared allelicB1 lineages in these closely related species may result from severe bottlenecks that occurred during speciation or during the ice ages that glaciated the rim of the north Pacific Ocean approximately every 100 000 years in the Pleistocene.The nucleotide sequence data reported in this paper have been submitted to the GenBank nucleotide sequence database and have been assigned the accession numbers U34692-U34720     

Annual changes in the population size of the salmon louse Lepeophtheirus salmonis (Copepoda: Caligidae) on high-seas Pacific Salmon (Oncorhynchus spp.), and relationship to host abundance

The population size of the salmon louse, Lepeophtheirus salmonis, was monitored annually in the summers of 1991–1997 by examining six species of Pacific salmon (Oncorhynchus spp.) caught by surface long-lines in oceanic offshore waters of the North Pacific Ocean and Bering Sea. The annual copepod population size on all salmonids caught was estimated by combining the calculated number of copepods carrying on each salmonid species. The copepod population fluctuated markedly from year to year, which resulted largely from marked annual changes in abundance of pink salmon (O. gorbuscha). Since pink salmon were most frequently and heavily infected and since their abundance changed every year, the copepod population was high in the years when this salmonid species was abundant, but low when it was rare. On the contrary, chum salmon (O. keta) did not show high prevalence and intensity of infection, but the annual abundance of this host species was consistently high, i.e. chum salmon carried many copepods every year. Copepods on other salmonid species (sockeye salmon O. nerka, coho salmon O. kisutch, chinook salmon O. tshawytscha, and steelhead trout O. mykiss) constantly formed a small percentage of the total copepod population. Both chum and pink salmon are the most important hosts in terms of their substantial contribution to support the copepod population, but the importance as hosts of each species is definitely different between the species. Chum salmon is a stable important host supporting the copepod population at a relatively high level every year, while the number of copepods on pink salmon annually exhibits marked fluctuations, and this salmonid species is regarded as an unstable important host.     

Sperm ultrastructure and a new type of spermiogenesis in two species of Pimelodidae, with a comparative review of sperm ultrastructure in Siluriformes (Teleostei: Ostariophysi)

During spermiogenesis, the spermatids of the pimelodid species Pimelodus maculatus and Pseudoplatystoma fasciatum show a central flagellum development, no rotation of the nucleus, and no nuclear fossa formation, in contrast to all previously described spermatids of Teleostei. These characteristics are interpreted as belonging to a new type of spermiogenesis, named here type III, which is peculiar to the family Pimelodidae. In P. maculatus and P. fasciatum, spermatozoa possess a spherical head and no acrosome; their nucleus contains highly condensed, homogeneous chromatin with small electron-lucent areas; and a nuclear fossa is not present. The centriolar complex lies close to the nucleus. The midpiece is small, has no true cytoplasmic channel, and contains many elongate and interconnected vesicles. Several spherical to oblong mitochondria are located around the centriolar complex. The flagellum displays the classical axoneme (9+2) and no lateral fins. Only minor differences were observed among the pimelodid species and genera. Otherwise, spermiogenesis and spermatozoa in the two species of Pimelodidae studied exhibit many characteristics that are not found in other siluriform families, mainly the type III spermiogenesis.     

Experimental microevolution: transplantation of pink salmon into the European North

Human-mediated translocations of species beyond their native ranges can enhance evolutionary processes in populations introduced to novel environments. We studied such processes in several generations of pink salmon Oncorhynchus gorbuscha introduced to the European North of Russia using a set of morphological and life-history traits as well as molecular genetic markers with different selective values: protein-coding loci, mtDNA, microsatellites, and MHC. The introduction of reproductively isolated pink salmon broodlines of odd and even years yielded different results. The odd-year broodline established self-reproducing local populations in many rivers of new range, but sustainable changes in external morphology, reproduction, and life-history, as well as the impoverishment of the gene pool occurred. Their successful colonisation of the new range resulted in specialisation manifested in the rapid directional shifts in some highly heritable phenotypic traits accompanied by increased homozygosity at molecular markers as a consequence of genetic drift and selective processes. The returns of transplanted pink salmon of even-year broodline decreased sharply already in the second generation, but there was no marked reduction of genetic diversity. Our data, as well as the analysis of the history of all pink salmon transplantations beyond the species range, demonstrate comparatively greater success of introduced odd-year broodline and permit to assume different adaptive plasticity of the even- and odd-year broodlines in pink salmon, what is most likely determined by differences in their evolutionary histories. Population genetic data suggest that the even-year broodline probably diverged from the odd-year broodline relatively recently and, due to the founder effect, may have lost a part of its genetic variation with which adaptive plasticity potential is associated.     

The genetic population structure of lacustrine sockeye salmon, <Emphasis Type="Italic">Oncorhynchus nerka,</Emphasis> in Japan as the endangered species

Lacustrine sockeye salmon (Oncorhynchus nerka) are listed as an endangered species in Japan despite little genetic information on their population structure. In order to clarify the genetic diversity and structure of Japanese populations for evaluating on the bottleneck effect and an endangered species, we analyzed the ND5 region of mitochondrial DNA (mtDNA) and 45 single nucleotide polymorphisms (SNPs) in 640 lacustrine sockeye salmon in Japan and 80 anadromous sockeye salmon in Iliamna Lake of Alaska. The genetic diversity of the Japanese population in both mtDNA and SNPs was significantly less than that of the Iliamna Lake population. Moreover, all Japanese populations had SNP loci deviating from the HWE. In spite of low genetic diversity, the SNP analyses resulted that the Japanese population was significantly divided into three groups. These suggest that Japanese sockeye salmon populations should be protected as an endangered species and genetically disturbed by the hatchery program and transplantations.     

Hybrids between chum Oncorhynchus keta and pink Oncorhynchus gorbuscha salmon: age,growth and morphology and effects on salmon production

Mature hybrids between chum salmon Oncorhynchus keta and pink salmon Oncorhynchus gorbuscha, which were identified by an intermediate colour pattern, were caught at the Kurilsky Hatchery, Iturup Island, Russia. Most of them were female and 3 years old (a partial freshwater year and 2 marine years), which is intermediate between the ages of maturity of the parental species. The hybrids exceed both parental species in the rate of growth, are large in size and robust and might successfully compete for mating in the wild or be chosen for artificial reproduction. The ratio of the scale length over width, R, is oblate (R < 1), whereas scales of the parental species are prolate (R > 1). From scale analyses, the c.v. in body size of hybrid females at the second marine year is twice that of O. keta, which suggests developmental instability in the hybrid. A dynamic model predicted that continuing hybridization at a low rate does not produce a substantial hybrid load due to selection against advanced‐generation hybrids and backcrosses. A high hybridization rate, however, may be an additional risk for genetic management and should be taken into account in programmes of artificial reproduction of Pacific salmon Oncorhynchus spp., although such hybrids might have commercial use in confined production systems.     

Genetic mapping of Y-chromosomal DNA markers in Pacific salmon

Sex chromosomes in fish provide an intriguing view of how sex-determination mechanisms evolve in vertebrates. Many fish species with single-factor sex-determination systems do not have cytogenetically-distinguishable sex chromosomes, suggesting that few sex-specific sequences or chromosomal rearrangements are present and that sex-chromosome evolution is thus at an early stage. We describe experiments examining the linkage arrangement of a Y-chromosomal GH pseudogene (GH-Y) sequence in four species of salmon (chum, Oncorhynchus keta; pink, O. gorbuscha; coho, O. kisutch; chinook, O. tshawytscha). Phylogenetic analysis indicates that GH-Y arose early in Oncorhynchus evolution, after this genus had diverged from Salmo and Salvelinus. However, GH-Y has not been detected in some Oncorhynchus species (O. nerka, O. mykiss and O. clarki), consistent with this locus being deleted in some lineages. GH-Y is tightly linked genetically to the sex-determination locus on the Y chromosome and, in chinook salmon, to another Y-linked DNA marker OtY1. GH-Y is derived from an ancestral GH2 gene, but this latter functional GH locus is autosomal or pseudoautosomal. YY chinook salmon are viable and fertile, indicating the Y chromosome is not deficient of vital genetic functions present on the X chromosome, consistent with sex chromosomes that are in an early stage of divergence.     

Risk management of non-target fish taxa in the Yakima River Watershed associated with hatchery salmon supplementation

Hatchery cultured salmon have the potential to interact strongly with other valued fish taxa (non-target taxa; NTT) in the natural environment. Monitoring and managing adverse interactions between hatchery supplemented salmon and NTT is one unique characteristic of a hatchery salmon supplementation program in the Yakima River, Washington. In this study, we evaluate impacts of spring Chinook salmon Oncorhynchus tshawytscha and coho salmon O. kisutch reintroduction to 15 NTT after 11 years of stocking approximately one million yearling smolts annually in the upper Yakima Basin between 1999 and 2009. Our risk management monitoring indicated changes in important response variables for NTT were within acceptable limits. Rigorous pre-implementation planning likely prevented many undesirable ecological impacts from the hatchery supplementation program. We illustrate a number of important features associated with risk management of hatchery and wild fish interactions. First, pre-project planning can eliminate many risks of concern and substantially reduce the need for risk containment during project implementation. Second, the sieve approach for monitoring impacts provided an acceptable balance between monitoring effort and risk containment ability, although in some cases, we would not detect impacts of interest. Third, rare and disbursed species that cannot be monitored effectively benefit from risk averse hatchery release strategies. Fourth, risk containment monitoring programs can be used to refute unsubstantiated claims of undesirable impacts. In short, our experience suggests that risk management of ecological interactions can occur by using a combination of pre-project adjustments through risk assessment and risk reduction, and by cost-effective risk containment monitoring and management.     

Parental relatedness and major histocompatibility effects on early embryo survivorship in Atlantic salmon

Salmon have provided key insights into the relative influence of natural and sexual selection on major histocompatibility complex (MHC) variation. Natural selection on salmon MHC genes has been demonstrated in pathogen studies, and there is evidence of MHC-based mate choice (sexual selection). We tested whether parental MHC genes affect survivorship of juvenile Atlantic salmon (Salmo salar) by quantifying the influence of parental genome-wide relatedness and MHC genotype on survivorship to the swim-up stage. Thirteen microsatellite loci were used to estimate the influence of genome-wide relatedness between parents on offspring survivorship and MHC genotypes were determined by sequencing part of the class IIβ gene. Our results revealed no significant relationship between early offspring survivorship and genome-wide relatedness, predicted MHC heterozygosity, or MHC allelic similarity. Overall, our data are consistent with the contention that excess MHC heterozygosity in Atlantic salmon juveniles is due to sexual selection as well as differential survival of offspring due to MHC genotype.     

Ultrastructure of melanocytes from retina and choroid of the Pacific salmon

The ultrastructure of the retina and choroid cells in three species of the Pacific salmon, Oncohrynchus gorbuscha, O. keta and O. masou, was studied. The structure of retina pigment cells is similar in all the three species, only a small difference was found in the percentage of lengthened and rounded forms of melanosomes. Melanocytes of the masu salmon differ also in the structure of their nuclei. The pigment cells of choroid differ from those of retina by a more extended form of melanosomes and by the presence of less rounded melanosomes. In the chum salmon retina we found electron lucent "contact vesicles", whose assignment is open to discussion. In retina pigment cells of the masu salmon smolts ready for migration, the activity of Golgi appararus rises, mitochondria and mature melanosomes increase in number. The choroid pigment cells are slightly swollen, their processes more often and deeper penetrate into the walls of vessels, down to the endothelium. Results of the experiment with the application of an artificial magnetic field (AMF) have shown that the retina and choroid pigment cells in the masu salmon fry react to the field of a certain direction. The phenomenon of magnetosensitivity of pigment cells is discussed in addition to their possible involvement in magnetoreception.     

Gonadotropic cells in the Atlantic salmon,Salmo salar

Summary The gonadotropin-producing cells (GTH-cells) in the Atlantic salmon were studied light and electron microscopically before, during and after spawning, and after injections of luteinizing hormone-releasing hormone (LH-RH). The double immunofluorescent technique was applied using rabbit anti-carp GTH as the first antibody. Numerous immunofluorescent cells were observed throughout the pars distalis, but very few in the pars intermedia. These cells are basophilic and PAS-positive, and ultrastructurally classified as globular gonadotropes. Only one gonadotropic cell type could be identified; its size, morphology and fine structure vary considerably. In the same specimen the GTH-cells can be predominantly globular or vesicular in appearance, depending on the reproductive phase of the fish. At spawning and after LH-RH injection, many GTH-cells reach a vacuolar stage; the content of the vacuoles is not immunofluorescent. Another cell type, which resembles GTH-cells in semithin sections, did not show gonadotropic properties; its nature and functional significance are unknown. In addition, the present study revealed an increase in the synthetic and exocytotic activity of prolactin cells after LH-RH injections. It is suggested that LH-RH mediates this effect via LH and eventually via estradiol.The authors are indebted to Dr. N. Johansson and superintendent N. Steffner for generously providing facilities at their institutes in Älvkarleby. They also wish to thank Mrs. Y. Lilliemarck and Mrs. V. Lundin for their skillful technical assistance. The grants made available by the Swedish Natural Science Research Council (No. 2124-037) are also gratefully acknowledged. Thanks are also due to Mrs. S.M. McNab for linguistic assistance     

Discovery and characterization of single nucleotide polymorphisms in coho salmon,Oncorhynchus kisutch

Molecular population genetic analyses have become an integral part of ecological investigation and population monitoring for conservation and management. Microsatellites have been the molecular marker of choice for such applications over the last several decades, but single nucleotide polymorphism (SNP) markers are rapidly expanding beyond model organisms. Coho salmon (Oncorhynchus kisutch) is native to the north Pacific Ocean and its tributaries, where it is the focus of intensive fishery and conservation activities. As it is an anadromous species, coho salmon typically migrate across multiple jurisdictional boundaries, complicating management and requiring shared data collection methods. Here, we describe the discovery and validation of a suite of novel SNPs and associated genotyping assays which can be used in the genetic analyses of this species. These assays include 91 that are polymorphic in the species and one that discriminates it from a sister species, Chinook salmon. We demonstrate the utility of these SNPs for population assignment and phylogeographic analyses, and map them against the draft trout genome. The markers constitute a large majority of all SNP markers described for coho salmon and will enable both population‐ and pedigree‐based analyses across the southern part of the species native range.