Pierre Duchesne

AFLP utility for population assignment studies: analytical investigation and empirical comparison with microsatellites.

Individual‐based population assignment tests have thus far mainly relied on the use of microsatellite loci. However, the logistic difficulty of screening large numbers of loci required to reach sufficient statistical power hampers the usefulness of microsatellites in situations of weak population structuring. Amplified fragment length polymorphisms (AFLP) represents an alternative for overcoming this logistical issue as the technique allows the user to characterize a much larger number of loci with a comparable analytical effort. In this study, an assignment test based on maximum likelihood for dominant markers was used to investigate the potential usefulness of AFLP for population assignment. We also compared assignment success achieved with AFLP with that obtained using microsatellites in a case study of low population differentiation involving whitefish (Coregonus clupeaformis) sympatric ecotypes. The analytical investigation showed that the minimum number of AFLP loci required to reach an assignment success of 95% stood within values that are easily achievable in many situations. This also showed how assignment success varied according to the number of AFLP loci used, their absolute frequency and their frequency differential and sampling errors, as well as the number of putative source populations. The case study showed that given a comparable analytical effort in the laboratory, AFLP were much more efficient than the microsatellite loci in discriminating the source of an individual among putative populations. AFLP resulted in higher assignment success at all levels of stringency and the log‐likelihood differences between populations obtained with AFLP for each individual were much larger than those obtained with microsatellites. These results indicate that research involving individual‐based population assignment methods should benefit importantly from the use of AFLP markers, especially in systems characterized by weak population structuring.

Parallelism in gene transcription among sympatric lake whitefish ( Coregonus clupeaformis Mitchill) ecotypes

We tested the hypothesis that phenotypic parallelism between dwarf and normal whitefish ecotypes (Coregonus clupeaformis, Salmonidae) is accompanied by parallelism in gene transcription. The most striking phenotypic differences between these forms implied energetic metabolism and swimming activity. Therefore, we predicted that genes showing parallel expression should mainly belong to functional groups associated with these phenotypes. Transcriptome profiles were obtained from white muscle by using a 3557 cDNA gene microarray developed for the Atlantic salmon (Salmo salar). A total of 1181 genes expressed in both lake populations hybridized on the array. Significant differential expression between ecotypes was detected for 134 (11.3%) and 195 (16.5%) gene clones in Cliff Lake and Indian Pond, respectively. Fifty‐one genes (4.3%) showed parallel differential expression between lakes, among which 35 were expressed in opposite directions. Sixteen genes (1.35%) showed true parallelism of transcription, which mainly belonged to energetic metabolism and regulation of muscle contraction functional groups. Variance in expression was significantly reduced for these genes compared to those not showing directionality in parallelism of expression. Candidate genes associated with parallelism in swimming activity and energetic metabolism based on their level and variance in expression were identified. These results add to the growing evidence that parallel phenotypic evolution also involves parallelism at both the genotypic and regulatory level, which may at least partly be associated with genetic constraints. It also provides further evidence for the determinant role of divergent natural selection in driving phenotypic divergence, and perhaps reproductive isolation, in the adaptive radiation of lake whitefish. This study adds to a nascent field employing microarrays as powerful tools for investigating the evolutionary processes of adaptive divergence among natural populations.

Potential of microsatellites for individual assignment: the North Atlantic redfish (genus Sebastes) species complex as a case study

We used the four redfish taxa (genus Sebastes) from the North Atlantic to evaluate the potential of multilocus genotype information obtained from microsatellites in assigning individuals at two different levels of group divergence. We first tested the hypothesis that microsatellites can diagnostically discriminate individual redfish from different groups. Second, we compared two different methods to quantify the effect of number of loci and likelihood stringency levels on the power of microsatellites for redfish group membership. The potential of microsatellites to discriminate individuals from different taxa was illustrated by a shared allele distance tree in which four major clusters corresponding to each taxa were defined. Concomitant with this strong discrimination, microsatellites also proved to be powerful in reclassifying specimens to the taxon of origin, using either an empirical or simulated method of estimating assignment success. By testing for the effect of both the number of loci and the level of stringency on the assignment success, we found that 95% of all specimens were still correctly reclassified with only four loci at the most commonly used criterion of log0. In contrast, the results obtained at the population level within taxa highlighted several problems of assignment that may occur at low levels of divergence. Namely, a drastic decrease of success with increasing stringency illustrated the lack of power of our set of loci. Strong discrepancy was observed between results obtained from the empirical and simulated methods. Finally, the highest assignment success was obtained when reducing the number of loci used, an observation previously reported in studies of human populations.

Genetic evidence for kin aggregation in the intertidal acorn barnacle ( Semibalanus balanoides )

It is generally assumed that larvae of benthic species are thoroughly mixed in the plankton and distributed randomly at settlement. Yet, it has also been hypothesized that a combination of larval gregarious behaviour coupled with particular oceanographic conditions may prevent larvae from mixing completely, and result in nonrandom spatial distributions following settlement. Using microsatellite markers, the main objective of this study was to investigate the occurrence of statistical connections between relatedness and settlement in the intertidal acorn barnacle from the Gulf of St Lawrence, Canada. A second objective was to test the hypothesis that patches of kin‐related individuals came from a common parental site. Our results indicated that a significant number of barnacles within a given sample were more closely related than expected by chance despite the enormous potential for admixture during the planktonic phase. Thus, eight out of 37 samples analysed had relatedness values significantly higher than expected from random settlement. Moreover, analyses of sibship network construction and network complexity tests provided evidence for the occurrence of networks within several samples that were characterized by strong connections among individuals. Thus, nonrandom planktonic dispersal associated with relatively stable oceanic currents, as well as additional ecological factors to be rigorously investigated (e.g. behavioural mechanisms), may be more important in determining patterns of genetic structure in marine benthic invertebrates than generally assumed. Therefore, documenting genetic patterns associated with kin aggregation should be a fruitful and an important avenue for future studies in marine invertebrates.

Surviving with low genetic diversity: the case of albatrosses.

Low genetic diversity is predicted to negatively impact species viability and has been a central concern for conservation. In contrast, the possibility that some species may thrive in spite of a relatively poor diversity has received little attention. The wandering and Amsterdam albatrosses (Diomedea exulans and Diomedea amsterdamensis) are long-lived seabirds standing at an extreme along the gradient of life strategies, having traits that may favour inbreeding and low genetic diversity. Divergence time of the two species is estimated at 0.84 Myr ago from cytochrome b data. We tested the hypothesis that both albatrosses inherited poor genetic diversity from their common ancestor. Within the wandering albatross, per cent polymorphic loci and expected heterozygosity at amplified fragment length polymorphisms were approximately one-third of the minimal values reported in other vertebrates. Genetic diversity in the Amsterdam albatross, which is recovering from a severe bottleneck, was about twice as low as in the wandering albatross. Simulations supported the hypothesis that genetic diversity in albatrosses was already depleted prior to their divergence. Given the generally high breeding success of these species, it is likely that they are not suffering much from their impoverished diversity. Whether albatrosses are unique in this regard is unknown, but they appear to challenge the classical view about the negative consequences of genetic depletion on species survival.

Migratory charr schools exhibit population and kin associations beyond juvenile stages

Few studies have critically investigated the genetic composition of wild fish schools. Yet, such investigations may have profound implications for the understanding of social organization and population differentiation in both fundamental and applied research. Using 20 microsatellite loci, we investigated the composition of 53 schools (total n = 211) of adult and subadult migratory brook charr (Salvelinus fontinalis) sampled from the known feeding areas of two populations inhabiting Mistassini Lake (Québec, Canada). We specifically tested whether (i) school members originated from the same population, (ii) individuals from the same population within schools were kin (half‐ or full‐siblings), and (iii) kin schooling relationships differed between sexes. Randomization tests revealed a tendency for most schools to be population specific, although some schools were population mixtures. Significantly more kin were found within schools than expected at random for both populations (≈ 21–34% of the total number of school members). This result, combined with the observed size range of individuals, indicated that stable associations between kin may occur beyond juvenile stages for up to 4 years. Nevertheless, a high proportion of school members were non‐kin (≈ 66–79%). No differences were detected between sexes in the propensity to school with kin. We discuss the hypothesis that the stable kin groups, rather than arising from kin selection, may instead be a by‐product of familiarity based on individual selection for the maintenance of local adaptations related to migration (natal and feeding area philopatry). Our results are noteworthy because they suggest that there is some degree of permanence in the composition of wild fish schools. Additionally, they support the hypothesis that schools can be hierarchically structured (from population members down to family groups) and are thus nonrandom genetic entities.

An analytical investigation of the dynamics of inbreeding in multi-generation supportive breeding

Supportive breeding is being increasingly usedas a measure to reduce the short-termprobability of extinction of populations withhighly reduced abundance relative to historicallevels. In this paper, we provide a conceptualframework and analytical tools to computechanges in inbreeding coefficient (F) in thecase of supportive breeding over any number ofgenerations. The dynamics of inbreedingcoefficients were investigated by means of asystem of recurrence equations. We focussed onquantifying the dynamics of F for specificcombinations of parameter values in terms ofthe effects of captive population census size,refreshment rate of breeders in captivity,scale of supplementation program, and migrationrate. We observed that supplementation did notalways result in substantial inbreedingincrement and several conditions loweredoverall inbreeding relative to controlsituations without supplementation. The censussize of captive populations was the single mostimportant controllable parameter determiningthe genetic consequences of supportivebreeding. While the proportion of captivebreeders brought into captivity from the wildbore a complex relationship to inbreedingcoefficient dynamics, the results indicatedthat managers should generally aim at highrefreshment rates (that is, large proportionsof their captive stock originating from thewild). This is especially important when asmall captive population is expected tocontribute large numbers of breeders to thesupplemented population. The analysis alsoshowed how supplemented populations connectedto a large metapopulation through gene flowrecover from the genetic risks of inbreedingdue to supportive breeding program more quicklythan isolated populations. The results of thisstudy join those of an increasing number ofinvestigations showing that supportive breedingdoes not always increase inbreeding, and mayeven decrease it in several circumstances.However, supportive breeding systems arecomplex, and results such as presented hereshould not be used in isolation, but inconsideration of other issues such as theconsequences on long-term fitness of wildindividuals.

FLOCK Provides Reliable Solutions to the “Number of Populations” Problem

Identifying groups of individuals forming coherent genetic clusters is relevant to many fields of biology. This paper addresses the K-partition problem: given a collection of genotypes, partition those genotypes into K groups, each group being a sample of the K source populations that are represented in the collection of genotypes. This problem involves allocating genotypes to genetic groups while building those groups at the same time without the use of any other a priori information. FLOCK is a non-Markov chain Monte Carlo (MCMC) algorithm that uses an iterative method to partition a collection of genotypes into k groups. Rules to estimate K are formulated and their validity firmly established by running simulations under several migration rates, migration regimes, number of loci, and values of K. FLOCK tended to build clusters largely consistent with the source samples. The performance of FLOCK was also compared with that of STRUCTURE and BAPS. FLOCK provided more accurate allocations to clusters and more reliable estimates of K; it also ran much faster than STRUCTURE. FLOCK is based on an entirely novel approach and provides a true alternative to the existing, MCMC based, algorithms. FLOCK v.2.0 for microsatellites or for AFLP markers can be downloaded from http://www.bio.ulaval.ca/no_cache/departement/professeurs/fiche_des_professeurs/professeur/11/13/.

Stable genetic polymorphism in heterogeneous environments: balance between asymmetrical dispersal and selection in the acorn barnacle

Elucidating the processes responsible for maintaining polymorphism at ecologically relevant genes is intimately related to understanding the interplay between selection imposed by habitat heterogeneity and a species’ capacity for dispersal in the face of environmental constraints. In this paper, we used a model‐based approach to solve equilibria of balanced polymorphism, given values of fitness and larval dispersal among different habitats in the acorn barnacle Semibalanus balanoides from the Gulf of St Lawrence. Our results showed that allele frequencies observed at both MPI* and GPI* loci represented stable equilibria, given empirical estimates of fitness values, and that considerably more larvae dispersed from one region (north) to the other (south) than vice versa. Dispersal conditions were predicted to be similar for the maintenance of polymorphism at both loci. Moreover, the values of asymmetrical dispersal required by the model to reach stable equilibria were compatible with empirical estimates of larval dispersal and oceanic circulation documented in this system. Overall, this study illustrated the usefulness of a modified and computable version of Bulmers model (1972) in order to test hypotheses of balanced polymorphism resulting from interactions between spatial selection and asymmetrical dispersal.

Groups of related belugas (Delphinapterus leucas) travel together during their seasonal migrations in and around Hudson Bay

Social structure involving long-term associations with relatives should facilitate the learning of complex behaviours such as long-distance migration. In and around Hudson Bay (Canada), three stocks of beluga whales form a panmictic unit, but have different migratory behaviours associated with different summering areas. We analysed genetic variation at 13 microsatellite loci among 1524 belugas, to test hypotheses about social structure in belugas. We found significant proportions of mother–offspring pairs throughout the migratory cycle, but average relatedness extended beyond close kinship only during migration. Average relatedness was significantly above random expectations for pairs caught at the same site but on different days or months of a year, suggesting that belugas maintain associations with a network of relatives during migration. Pairs involving a female (female–female or male–female) were on average more related than pairs of males, and males seemed to disperse from their matrilineal group to associate with other mature males. Altogether, our results indicate that relatives other than strictly parents, and especially females, play a role in maintaining a social structure that could facilitate the learning of migration routes. Cultural conservatism may limit contributions from nearby summer stocks to endangered stocks such as the Eastern Hudson Bay beluga.