Silver-banded karyotypes of the rainbow trout and the brook trout and their hybrids: disappearance in allotriploids of Ag-NORs originated from the brook trout

Silver-banded karyotypes of the rainbow trout Salmo gairdneri, the brook trout Salvelinus fontinalis, and their hybrids were described. Diploid-type and triploid-type hybrids were obtained. Triploid-type hybrids had two maternal genomes of the rainbow trout and one paternal genome of the brook trout. The rainbow trout had one pair of M or SM with Ag-NORs near the satellite. In the brook trout, Ag-NORs were observed in four pairs of chromosomes, but each cell had different number of chromosomes with Ag-NORs. In all observed cells of triploid-type hybrids, Ag-NORs originated from the brook trout were not recognized, whereas those from the rainbow trout were found.     

Sex linkage in salmonids: Evidence from a hybridized genome of brook trout and arctic charr

In second-generation sparctics (Salvelinus fontinalis × Salvelinus alpinus) backcrossed toS. fontinalis, we have identified tight classical linkage of phenotypic sex withLdh-1, Aat-5, andGpi-3. We designate this locusSex-1 and suggest that it may be the primary sex-determining locus in salmonids. Cumulative salmonid gene-to-centromere map distances for the three biochemical loci put the order as centromere—Ldh-1—(Aat-5 andGpi-3), with the latter two loci being tightly linked. An absence of association of phenotypic sex (presumably Sex-1) with these same three loci and other loci known to be linked to these loci is shown in splakes (S. fontinalis × Salvelinus namaycush) and cutbows (Salmo gairdneri × Salmo clarki). These data imply that the linkage ofSex-1 with these loci is found only inS. alpinus and support the view thatSex-1 lies across the centromere from these three loci inS. alpinus, representing a Robertsonian fusion not found in any of the other four species. A similar specific Robertsonian fusion is argued forS. gairdneri, whereSex-1 may be linked across a centromere to another biochemical locus (Ha). These linkage results and chromosomal observations of other investigators suggest thatSex-1 lies on an information-depauperate arm.     

A linkage map for brown trout (Salmo trutta): chromosome homeologies and comparative genome organization with other salmonid fish

We report on the construction of a linkage map for brown trout (Salmo trutta) and its comparison with those of other tetraploid-derivative fish in the family Salmonidae, including Atlantic salmon (Salmo salar), rainbow trout (Oncorhynchus mykiss), and Arctic char (Salvelinus alpinus). Overall, we identified 37 linkage groups (2n = 80) from the analysis of 288 microsatellite polymorphisms, 13 allozyme markers, and phenotypic sex in four backcross families. Additionally, we used gene-centromere analysis to approximate the position of the centromere for 20 linkage groups and thus relate linkage arrangements to the physical morphology of chromosomes. Sex-specific maps derived from multiple parents were estimated to cover 346.4 and 912.5 cM of the male and female genomes, respectively. As previously observed in other salmonids, recombination rates showed large sex differences (average female-to-male ratio was 6.4), with male crossovers generally localized toward the distal end of linkage groups. Putative homeologous regions inherited from the salmonid tetraploid ancestor were identified for 10 pairs of linkage groups, including five chromosomes showing evidence of residual tetrasomy (pseudolinkage). Map alignments with orthologous regions in Atlantic salmon, rainbow trout, and Arctic char also revealed extensive conservation of syntenic blocks across species, which was generally consistent with chromosome divergence through Robertsonian translocations.     

DEVELOPMENTAL INSTABILITY AND HIGH MERISTIC COUNTS IN INTERSPECIFIC HYBRIDS OF SALMONID FISHES

Interspecific hybrids have been proposed to have reduced developmental stability in comparison to their parental species because the parental genomes have not undergone selection for the maintenance of developmental stability when they occur together. We present data from four interspecific hybrids of salmonid fishes that support this view. Natural hybrids of bull trout (Salvelinus confluentus) with brook trout (Salvelinus fontinalis) and laboratory hybrids of rainbow trout (Salmo gairdneri) with Yellowstone (Salmo clarki bouvieri), westslope (S. c. lewisi), and coastal (S. c. clarki) cutthroat trout all have higher levels of fluctuating asymmetry than either of their parental species raised in the same environment. Thus, the hybrids have reduced developmental stability. The hybrids do not have meristic counts intermediate to the counts of the parental species. The hybrids usually have counts as high as the species with the higher count for those characters that differ between the parental species and often have higher counts for those characters that do not differ between the parental species. We suggest that the tendency for interspecific hybrids to have high meristic counts may be related to differences between the species in the length and timing of the developmental periods during which the counts of the characters are determined.     

The ribosomal RNA genes of three salmonid species

The ribosomal RNA cistrons of three species of trout: Salvelinus namaycush (lake trout), Salvelinus fontinalis (brook trout), and Salmo gairdneri (rainbow trout) were examined by Southern blot analysis of genomic DNA. The repeat length of the cistron of S. namaycush is 26 kb. A repeat-length polymorphism was observed in some of the individual fish examined. These individuals showed 24-kb repeats. In some individuals both forms were present. The restriction maps of the transcribed regions of all three species were similar and showed a site homology with other vertebrate ribosomal RNA genes. Interspecific comparison showed restriction-site differences within the spacer regions examined. A restriction-site polymorphism within the 28 S gene was observed in S. fontinalis. The rDNA of S. namaycush liver showed a high degree of methylation as determined by digestion with the restriction endonucleases MspI and HpaII.     

Cholesterol levels in the blood of Atlantic salmonids

Total cholesterol and high density lipids (HDL) were measured in the blood of Atlantic salmon (Salmo salar), brook trout (Salvelinus fontinalis) and rainbow trout (Salmo gairdneri) from a variety of dietary and environmental regimes. At spawning the total cholesterol of cage reared S. salar was high compared to sea ranched S. salar. Starvation of S. salar in freshwater for 6-11 months depressed total blood cholesterol levels, but not to unusually low levels. A cholesterol supplement of 0.02 g/100 g body weight to a chow diet of 0.38 g/100 g body weight had no significant effect on the total blood cholesterol level in Salvelinus fontinalis. The high proportion of HDL in the blood of S. salar is discussed in terms of its possible importance during the atherosclerotic lesion formation that normally accompanies spawning in these salmon.     

Cytogenetic analysis of pseudolinkage of ldh Loci in the teleost genus salvelinus

Cytological and genetic analyses provide evidence that spontaneous centric fusion and fission can account for curious patterns of pseudolinkage of two LDH loci in males of brook trout (Salvelinus fontinalis) and in the F₁, F₂ and backcross generations of lake trout (S. namaycush) x brook trout hybrids. Intraindividual polymorphisms for acrocentric and metacentric chromosomes in somatic and gonadal tissue of these fish have been related to the proposed polyploid evolution in Salmonidae.     

A genetic linkage map for coho salmon (Oncorhynchus kisutch)

Construction of genetic linkage maps is an important first step for a variety of genomic applications, such as selective breeding in aquaculture, comparative studies of chromosomal evolution and identification of loci that have played key roles in the evolution of a species. Here we present a sex-specific linkage map for coho salmon. The map was constructed using 148 AFLP markers, 133 microsatellite loci and the phenotypic locus SEX . Twenty-four linkage groups spanning 287.4 cM were mapped in males, and 33 linkage groups spanning 429.7 cM were mapped in females. Several male linkage groups corresponded to two female linkage groups. The combination of linkage groups across both sexes appeared to characterize regions of 26 chromosomes. Two homeologous chromosomes were identified based on information from duplicated loci. Homologies between the coho and rainbow trout maps were examined. Eighty-six loci were found to form common linkage relationships between the two maps; these relationships provided evidence for whole-arm fissions, fusions and conservation of chromosomal regions in the evolution of these two species.     

A comparative analysis of the rainbow trout genome with 2 other species of fish (Arctic charr and Atlantic salmon) within the tetraploid derivative Salmonidae family (subfamily: Salmoninae).

We updated the genetic map of rainbow trout (Oncorhynchus mykiss) for 2 outcrossed mapping panels, and used this map to assess the putative chromosome structure and recombination rate differences among linkage groups. We then used the rainbow trout sex-specific maps to make comparisons with 2 other ancestrally polyploid species of salmonid fishes, Arctic charr (Salvelinus alpinus) and Atlantic salmon (Salmo salar) to identify homeologous chromosome affinities within each species and ascertain homologous chromosome relationships among the species. Salmonid fishes exhibit a wide range of sex-specific differences in recombination rate, with some species having the largest differences for any vertebrate species studied to date. Our current estimate of female:male recombination rates in rainbow trout is 4.31:1. Chromosome structure and (or) size is associated with recombination rate differences between the sexes in rainbow trout. Linkage groups derived from presumptive acrocentric type chromosomes were observed to have much lower sex-specific differences in recombination rate than metacentric type linkage groups. Arctic charr is karyotypically the least derived species (i.e., possessing a high number of acrocentric chromosomes) and Atlantic salmon is the most derived (i.e., possessing a number of whole-arm fusions). Atlantic salmon have the largest female:male recombination ratio difference (i.e., 16.81:1) compared with rainbow trout, and Arctic charr (1.69:1). Comparisons of recombination rates between homologous segments of linkage groups among species indicated that when significant experiment-wise differences were detected (7/24 tests), recombination rates were generally higher in the species with a less-derived chromosome structure (6/7 significant comparisons). Greater similarity in linkage group syntenies were observed between Atlantic salmon and rainbow trout, suggesting their closer phylogenetic affinities, and most interspecific linkage group comparisons support a model that suggests whole chromosome arm translocations have occurred in the evolution of this group. However, some possible exceptions were detected and these findings are discussed in relation to their influence on segregation distortion patterns. We also report unusual meiotic segregation patterns in a female parent involving the duplicated (homeologous) linkage group pair 12/16 and discuss several models that may account for these patterns.     

Genetic variation,inheritance, and quaternary structure of malic enzyme in brook trout (Salvelinus fontinalis)

Electrophoretic variation is described for malic enzyme (ME) for the first time in brook trout (Salvelinus fontinalis). Since the quaternary structure of ME was not clear from examination of banding patterns in brook trout alone, ME phenotypes in rainbow trout (Salmo gairdneri) × brook trout hybrids as well as in esocid species demonstrated that ME is tetrameric. A model of two duplicated loci is proposed to account for the observed variation. One locus (ME-2) is fixed and one locus (ME-1) is variable with three electrophoretically distinct alleles; the protein products of ME-1 are reduced in activity relative to the protein products of ME-2. Joint segregation was examined between ME-1 and ten other biochemical loci in brook trout, and between ME-1, ME-2, and nine other biochemical loci in a splake—lake trout (Salvelinus namaycush) × brook trout hybrid—backcross. All pairwise examinations showed random assortment except ME-2 with an isocitrate dehydrogenase locus (IDH-3), which showed complete linkage in the splake backcross. This may be due to a chromosomal aberration.Authorized for publication as Paper No. 5599 in the Journal Series of The Pennsylvania Agricultural Experiment Station, University Park, Pennsylvania, in cooperation with the Benner Spring Fish Research Station, The Pennsylvania Fish Commission, Bellefonte, Pennsylvania. M.S. was supported by an NSF Graduate Fellowship.     

Comparative Recombination Distances among Zea Mays L. Inbreds, Wide Crosses and Interspecific Hybrids

Recombination distances and linkage heterogeneity were compared among a wide range of maize inbreds, wide crosses and maize X teosinte hybrids. Twelve maize and four teosinte races were backcrossed to stocks fixed for rare marker alleles on chromosome arm 1L. Recombination fraction estimates were higher for exotic germplasm than for either U.S. maize or maize X teosinte crosses. Serrano, Tuxpeno and a US-adapted inbred line of tropical origin, NC300, exhibited enhanced recombination. Three of the four maize X teosinte hybrids had little or no recombination between two loci. The observed recombination ``shrinkage' resulted from an apparent inversion in the vicinity of the Amp1 locus. Average recombination distances among common marker loci for composite maps were highly variable, even when map construction was restricted to maize germplasm of similar origins.     

Occasional recombination of a selfish X‐chromosome may permit its persistence at high frequencies in the wild

The sex‐ratio X‐chromosome (SR) is a selfish chromosome that promotes its own transmission to the next generation by destroying Y‐bearing sperm in the testes of carrier males. In some natural populations of the fly Drosophila neotestacea, up to 30% of the X‐chromosomes are SR chromosomes. To investigate the molecular evolutionary history and consequences of SR, we sequenced SR and standard (ST) males at 11 X‐linked loci that span the ST X‐chromosome and at seven arbitrarily chosen autosomal loci from a sample of D. neotestacea males from throughout the species range. We found that the evolutionary relationship between ST and SR varies among individual markers, but genetic differentiation between SR and ST is chromosome‐wide and likely due to large chromosomal inversions that suppress recombination. However, SR does not consist of a single multilocus haplotype: we find evidence for gene flow between ST and SR at every locus assayed. Furthermore, we do not find long‐distance linkage disequilibrium within SR chromosomes, suggesting that recombination occurs in females homozygous for SR. Finally, polymorphism on SR is reduced compared to that on ST, and loci displaying signatures of selection on ST do not show similar patterns on SR. Thus, even if selection is less effective on SR, our results suggest that gene flow with ST and recombination between SR chromosomes may prevent the accumulation of deleterious mutations and allow its long‐term persistence at relatively high frequencies.     

Isolation of a Loma salmonae variant: biological characteristics and host range

A Salvelinus -infecting variant of Loma salmonae , derived from naturally-infected Chinook salmon Oncorhynchus tshawytscha by serial passage through brook trout Salvelinus fontinalis , has been isolated and amplified. Loma salmonae SV ( Salvelinus -variant) has a high preference for species of Salvelinus (brook trout and Arctic charr S. alpinus ) and low virulence and preference for species of Oncorhynchus (rainbow trout O. mykiss , Chinook salmon, cohoSalmon O. kisutch ) or Salmo (Atlantic salmon Salmo salar ). Although this variant of L. salmonae was different from the original, the differences do not justify describing it as a new species, although definitive determination is pending.     

Continuous linkage map of human chromosome 14 short tandem repeat polymorphisms.

Nine moderately to highly informative short tandem repeat polymorphisms were assigned to chromosome 14 using somatic cell hybrids and were mapped using linkage analysis. The nine markers formed a continuous linkage map covering almost the entire long arm from 14q11.2 to q32. The markers filled a large gap within previously reported linkage maps for this chromosome. Best order of the new loci from q11.2 to q32 was D14S50, D14S54, D14S49, D14S47, D14S52, D14S53, D14S55, D14S48, and D14S51. The order shown for all adjacent pairs of loci was very strongly favored with the exception of loci pair D14S55 and D14S48, for which the order was moderately favored. Map lengths for the nine loci were 142 cM in females and 72 cM in males. Female recombination frequencies exceeded male recombination frequencies in the middle and distal portions of the map.     

Cytogenetic analyses of a Salvelinus hybrid reveal an evolutionary relationship between the parental species

Mitotic analyses of the brook trout (Salvelinus fontinalis) x arctic char (S. alpinus) hybrids (sparctic trout) revealed a mode of 2n = 82 with 18 metacentric and 64 acrocentric chromosomes. The brook trout had 2n = 84 with 16 metacentric chromosomes and the arctic char had 2n = 80 with 20 metacentric chromosomes; both species are derivatives of a single tetraploid event. Variable multivalent-like configurations that may be centromeric associations of bivalents were observed in C-banded pachytene figures of female sparctic trout. Metaphase I analyses of sparctic trout males indicated that two fusions of nonhomologous acrocentric chromosomes representing two duplicated chromosome sets must have occurred in the arctic char after its evolutionary divergence from the brook trout. A mode of seven tetravalent rods per cell suggests that preferential multivalent pairing occurs in the sparctic hybrid; metaphase I analyses of S. alpinus males revealed a mode of only five tetravalent rods per cell. The presence of multivalents implies that the arctic char, like the brook trout, is still undergoing diploidization. Cytochemical detection of the nucleolar organizer region (NOR) revealed intra- and interspecific as well as intraindividual variability in the numbers and types of chromosomes (metacentric or acrocentric) on which NORs appeared in arctic char and sparctic trout. Brook trout only had NORs on acrocentric chromosomes. This may indicate that different chromosomal fusions occurred in the evolution of brook trout from arctic char.     

Genetic variation for tolerance to acidic water in salmonids

Water pH is an important factor affecting the general water quality as well as quality traits in fishes, and the magnitude of the effect varies among species. The massive and negative effect of acidification of rivers and lakes became evident during the 1960s and 1970s and caused the depletion of fish stocks in several countries in the northern hemisphere. Significant variation in tolerance to acidic water has been documented among salmonid species, and large genetic variation has been identified among strains of brown trout Salmo trutta, brook trout Salvelinus fontinalis and Atlantic salmon Salmo salar. For S. trutta, S. fontinalis and S. salar, there is considerable additive genetic variation in tolerance to acidic water, with heritabilities (h(2)) ranging from 0·09 to 0·27 for dead eyed-eggs (the period most sensitive to low pH). The main reasons for depletion of freshwater fish stocks are discussed.     

Gene mapping of isozyme loci in chum salmon

Recombination values were used to calculate the gene-centromere map distances for four electrophoretically detected loci, Aat3, Idh1, Idh4, and Mpi, in chum salmon (Oncorhynchus keta). We also report the results from 39 pairwise examinations for joint segregation for 10 loci in nine testcross families. Only two loci assorted nonrandomly--either Aat1 or Aat2 with Gpt. Gene-centromere distances for Aat3 and Mpi differed significantly from those reported previously for rainbow trout (Salmo gairdneri), a closely related species. This difference indicates either the presence of chromosome rearrangements or a different rate of recombination between the species. These results contrast with the conservation of linkage distances previously reported within and between other salmonid genera.     

Joint Segregation of Biochemical Loci in Salmonidae. II. Linkage Associations from a Hybridized Salvelinus Genome (S. NAMAYCUSHxS. FONTINALIS)

The results of more than 300 pairwise examinations of biochemical loci for joint segregation in brook trout (Salvelinus fontinalis) and in the hybridized genome of lake trout (S. namaycush) x brook trout are summarized. Nineteen loci have been assigned to the following eight linkage groupings on the basis of nonrandom assortment, including cases of both classical linkage and pseudolinkage: ODH with PMI with PGI-3, PGI-2 with SDH, ADA-1 with AGP-2, AAT-(1,2) with AGP-1 with MDH-1, MDH-3 with MDH-4, LDH-3 with LDH-4, IDH-3 with ME-2 and GUS with CPK-1. Pseudolinkage (an excess of nonparental progeny types) was observed only for male testcross parents. The results suggest that this phenomenon involves homeologous chromosome arms as evidenced by the de novo association of presumed duplicate loci in each case. Classical linkage has not been found for the five pairs of duplicate loci examined in Salvelinus, suggesting that not all of the eight metacentrics in the haploid complement involve fusions of homeologous chromosomes. Females consistently showed a greater degree of recombination.     

A microsatellite linkage map of rainbow trout (Oncorhynchus mykiss) characterized by large sex-specific differences in recombination rates

We constructed a genetic linkage map for a tetraploid derivative species, the rainbow trout (Oncorhynchus mykiss), using 191 microsatellite, 3 RAPD, 7 ESMP, and 7 allozyme markers in three backcross families. The linkage map consists of 29 linkage groups with potential arm displacements in the female map due to male-specific pseudolinkage arrangements. Synteny of duplicated microsatellite markers was used to identify and confirm some previously reported pseudolinkage arrangements based upon allozyme markers. Fifteen centromeric regions (20 chromosome arms) were identified with a half-tetrad analysis using gynogenetic diploids. Female map length is approximately 10 M, but this is a large underestimate as many genotyped segments remain unassigned at a LOD threshold of 3.0. Extreme differences in female:male map distances were observed (ratio F:M, 3.25:1). Females had much lower recombination rates (0.14:1) in telomeric regions than males, while recombination rates were much higher in females within regions proximal to the centromere (F:M, 10:1). Quadrivalent formations that appear almost exclusively in males are postulated to account for the observed differences.     

Post-stocking movements and recapture of hatchery-reared trout released into flowing waters—a review

The success of stocking with hatchery-reared trout has been the subject of varied investigations for the past half-century. Percentage returns are summarised, and literature on the post-stocking movements of hatchery-reared trout is reviewed. Factors affecting the poststocking movements are considered, special attention being paid to studies on industrial rivers. Highest returns are obtained from stockings, with trout of a size suitable for angling, made during or shortly before the angling season. The majority of stocked brown trout, Salmo trutta tend to remain close to the area of stocking, but brook trout, Salvelinus fontinalis and rainbow trout, Salmo gairdneri show greater movement, usually in a down-stream direction. Greater dispersion of all species occurs if they have overwintered prior to capture or have been stocked in'cold water'or in small upstream stretches of river.