ALBERT

Notes: Biochemical methods can detect variation at individual genetic loci, making possible the direct assessment of natural hybridization and introgression between fish populations. Protein electro-phoresis has been used to confirm and extend knowledge of many situations where species hybrids have been detected by morphological analyses. New cases of natural hybridization, including some at the subspecies level, have also been identified. Biochemical studies have provided the first conclusive evidence of natural post F1 hybrids and of introgression between fish taxa. The strongest cases for introgression have used a combined analysis of nuclear protein genes and taxaspecific maternally inherited mitochondrial DNA variation. Information on the significance of introgression as a source of gene flow between taxa, particularly below the species level where sympatric subspecies and sibling species are involved, should expand in the future as the numbers and types of nuclear and mitochondrial DNA loci which can be assayed for variation increase. The full importance of introgressive hybridization in speciation may then be understood.

Notes: Previous studies of the ESTD* isozyme locus in the Atlantic salmon show the *80 allele to be absent across the species’ European range, with the exception of northern Russia, whereas the allele is nearly fixed in North American populations. The allele was found in samples from 15 out of 18 rivers on the Kola Peninsula and White sea coast and had frequencies that ranged from 0·017 to 0·363. Typing of fish in nine of these rivers for mtDNA variation in the ND1 gene region found variation characteristic of North American salmon in three Kola Peninsula populations and is the only part of Europe where such variants have been detected. The study area was completely glaciated during the late Pleistocene period and the restriction of the European distribution of these ESTD* and mtDNA variants to this area suggests that salmon that colonized the Russian Arctic coast rivers included fish of North American origin after the Pleistocene glacier had retreated.

Notes: Electrophoretic studies of proteins remain a primary source of insight into genetic diversity in many species including the Atlantic salmon Salmo salar, one of the most culturally and economically important fish species of the North Atlantic region. Since 1966, 〉350 scientific papers on protein variation have been published encompassing 25 000+ salmon from over 400 locations in 〉200 river systems across the species’ distribution. Variation has been detected at 30% of the 110 protein loci screened, though most studies examine 〈40. The method has been applied largely to the investigation of population structure and differentiation, but work has also led to the systematic revision of the genus Salmo and remains the primary source of insight into hybridization in the wild with brown trout Salmo trutta. Spatial patterns of differentiation show temporal stability, both within and among river systems, and strongly support structuring of the species into river and tributary specific populations and the designation of European and North American populations as distinct sub-species. They also show widespread regional differentiation within both continents, beyond the marked subcontinental differences between Baltic Sea and Atlantic Ocean populations in Europe. Most of the differentiation probably reflects gene flow and founder events associated with colonization following the retreat of the glaciers from much of the species’ modern range. However, variation at MEP-2* shows strong correlations with environmental temperature, both within and among rivers, and associations with phenotypic performance. This suggests selection is acting on the locus and provides compelling evidence for the local adaptation of populations. Protein studies have led to more population centred management of the species and have been exploited in the discrimination of regional stocks in mixed stock analysis in high seas fisheries, particularly in the Baltic Sea, and as markers for the assessment of stocking success. They have also advanced insight into how the genetic character of populations can be changed in cultivation and the potential impact of salmon aquaculture and stocking on wild populations. The method has been largely superseded by DNA based analyses, but the results remain highly relevant to Atlantic salmon management and conservation and are an irreplaceable data set for studying genetic stability of populations over time.

Notes: A previous analysis of proportions of stocked and wild Atlantic salmon among angled fish in the Rivers Asón and Nansa in northern Spain based on the analysis of MEP-2* genotypes is extended. The results reinforce the initial conclusion that returns of stocked Scottish salmon are significantly lower than returns for wild fish.

Notes: The effect of natural selection on the mMEP-2* locus on measures of genetic divergence among Atlantic salmon populations was investigated by examining the pattern of change in the level of genetic differentiation (FST) averaged over loci when data on the mMEP-2* locus were either included or excluded. The level of FST among populations at various geographic scales was estimated from allele frequencies at up to four loci (sAAT-4*, IDDH-1*, IDHP-3*, and mMEP-2*). At smaller geographic scales (within river systems or limited geographic regions) levels of variance in mMEP-2* allele frequencies were reduced relative to mean levels. At larger geographic scales (across continents or the species range) variation in mMEP-2* allele frequencies was greater than mean levels. These results suggest an a priori hypothesis for the effect of selection on the mMEP-2* locus which may be applied in future studies on variation in protein coding or other (e.g. mini- and microsatellite) loci in the Atlantic salmon. It is recommended that estimates of gene flow among populations of the Atlantic salmon based on mean FST estimates which include data on the mMEP-2* locus should be viewed with caution.

Notes: Spatial variation at the diallelic Me-2 locus in the Atlantic salmon, Salmo salar L., was analysed using data from 95 river basins. Gene diversity was apportioned as follows: 63% within samples, 20% between North America and Europe, 14% between regions within continents and 3% within and among rivers within regions. On both continents the variation between rivers was clinal with latitude and highly correlated with summer temperatures. The correlation was detectable within and between rivers. These correlations strongly suggest that variation at the locus is subject to the direct or indirect effects of natural selection, and that caution is required when interpreting between-location differentiation at the locus as evidence for distinct stocks.

Notes: From 1983 to 1986 the Rivers Ason and Nansa (northern Spain) were stocked with over 100 000 eyed ova year−1 from a Scottish hatchery in an effort to increase angling catches of Atlantic salmon. An inherent genetic marker was used to estimate the contributions of stocked and wild fish from these year-classes to the 1988 rod-and-line fishery. Stocking with foreign ova yielded 0.9 fish per 10 000 ova in the R. Asón and zero fish in the R. Nansa. Natural egg deposition yielded 4.1 and 4.9 fish per 10000 ova, respectively. Reasons for the difference in performance between stocked and wild fish are examined.

Notes: PCR primers were successfully designed to amplify small ND1 gene fragments for RFLP genotyping of degraded Atlantic salmon Salmo salar mtDNA. Analysis of archival scales with these primers, when existing primer sets failed, show Atlantic salmon from the George River, Quebec, to include European haplotypes and those from the Kapisidlit River, West Greenland, to be fixed for a European haplotype characteristic of Baltic populations.

Notes: The distribution of genetic variation for NAD+ malic enzyme (ME*) polymorphism in the Atlantic salmon was assessed in both anadromous and resident populations. The analysis revealed a major allelic divergence between North America and Europe. The *80 variant occurred in 39 of 40 North American samples, ranging in frequency from 0 to 0.515. In contrast, it was detected in only three fish from two of the 35 European locations analysed. Eleven fish from two rivers in north-west France had the *110 variant. While heterogeneity among North American populations was significant, no regional differentiation was apparent and anadromous and resident salmon were not found to differ.