Male and female recombination landscapes of diploid Arabidopsis arenosa

The number and placement of meiotic crossover events during meiosis have important implications for the fidelity of chromosome segregation as well as patterns of inheritance. Despite the functional importance of recombination, recombination landscapes vary widely among and within species, and this can have a strong impact on evolutionary processes. A good knowledge of recombination landscapes is important for model systems in evolutionary and ecological genetics, since it can improve interpretation of genomic patterns of differentiation and genome evolution, and provides an important starting point for understanding the causes and consequences of recombination rate variation. Arabidopsis arenosa is a powerful evolutionary genetic model for studying the molecular basis of adaptation and recombination rate evolution. Here, we generate genetic maps for 2 diploid A. arenosa individuals from distinct genetic lineages where we have prior knowledge that meiotic genes show evidence of selection. We complement the genetic maps with cytological approaches to map and quantify recombination rates, and test the idea that these populations might have distinct patterns of recombination. We explore how recombination differs at the level of populations, individuals, sexes and genomic regions. We show that the positioning of crossovers along a chromosome correlates with their number, presumably a consequence of crossover interference, and discuss how this effect can cause differences in recombination landscape among sexes or species. We identify several instances of female segregation distortion. We found that averaged genome-wide recombination rate is lower and sex differences subtler in A. arenosa than in Arabidopsis thaliana.ISSN:1943-263

Male meiotic recombination rate varies with seasonal temperature fluctuations in wild populations of autotetraploid Arabidopsis arenosa

Meiosis, the cell division by which eukaryotes produce haploid gametes, is essential for fertility in sexually reproducing species. This process is sensitive to temperature, and can fail outright at temperature extremes. At less extreme values, temperature affects the genome-wide rate of homologous recombination, which has important implications for evolution and population genetics. Numerous studies in laboratory conditions have shown that recombination rate plasticity is common, perhaps nearly universal, among eukaryotes. These studies have also shown that variation in the length or timing of stresses can strongly affect results, raising the important question whether these findings translate to more variable field conditions. Moreover, lower or higher recombination rate could cause certain kinds of meiotic aberrations, especially in polyploid species—raising the additional question whether temperature fluctuations in field conditions cause problems. Here, we tested whether (1) recombination rate varies across a season in the wild in two natural populations of autotetraploid Arabidopsis arenosa, (2) whether recombination rate correlates with temperature fluctuations in nature, and (3) whether natural temperature fluctuations might cause meiotic aberrations. We found that plants in two genetically distinct populations showed a similar plastic response with recombination rate increases correlated with both high and low temperatures. In addition, increased recombination rate correlated with increased multivalent formation, especially at lower temperature, hinting that polyploids in particular may suffer meiotic problems in conditions they encounter in nature. Our results show that studies of recombination rate plasticity done in laboratory settings inform our understanding of what happens in nature

Sanger_fasta_seqs

Fasta sequences of re-sequenced RAD-markers (Sanger sequencing). First line (">") denotes the sequenced individual and the primer used. "F" stands for forward and "R" for reverse prime

Consensus sequences

Consensus sequences with assigned genotype for each individual. 48 text files in total - for each of the four lines, 10 daughter individual files (marked with "d") and files for two mother samples (marked with "m"). _consensus.txt file structure: contig 3'position coverage(sum) haplotype(X-OR-Y) diploid_homozygote(N)-OR-diploid_heterozygote(Y)-OR-haploid_low_coverage(L)-OR-hemizygote(H) consensus_sequenc

Male meiotic recombination rate varies with seasonal temperature fluctuations in wild populations of autotetraploid Arabidopsis arenosa

Meiosis, the cell division by which eukaryotes produce haploid gametes, is essential for fertility in sexually reproducing species. This process is sensitive to temperature, and can fail outright at temperature extremes. At less extreme values, temperature affects the genome-wide rate of homologous recombination, which has important implications for evolution and population genetics. Numerous studies in laboratory conditions have shown that recombination rate plasticity is common, perhaps nearly universal, among eukaryotes. These studies have also shown that variation in the length or timing of stresses can strongly affect results, raising the important question whether these findings translate to more variable field conditions. Moreover, lower or higher recombination rate could cause certain kinds of meiotic aberrations, especially in polyploid species-raising the additional question whether temperature fluctuations in field conditions cause problems. Here, we tested whether (1) recombination rate varies across a season in the wild in two natural populations of autotetraploid Arabidopsis arenosa, (2) whether recombination rate correlates with temperature fluctuations in nature, and (3) whether natural temperature fluctuations might cause meiotic aberrations. We found that plants in two genetically distinct populations showed a similar plastic response with recombination rate increases correlated with both high and low temperatures. In addition, increased recombination rate correlated with increased multivalent formation, especially at lower temperature, hinting that polyploids in particular may suffer meiotic problems in conditions they encounter in nature. Our results show that studies of recombination rate plasticity done in laboratory settings inform our understanding of what happens in nature.ISSN:0962-1083ISSN:1365-294

Data from: How clonal are clones? A quest for loss of heterozygosity during asexual reproduction in Daphnia magna

Due to the lack of recombination, asexual organisms are predicted to accumulate mutations and show high levels of within-individual allelic divergence (heterozygosity) however, empirical evidence for this prediction is largely missing. Instead, evidence of genome homogenization during asexual reproduction is accumulating. Ameiotic crossover recombination is a mechanism that could lead to long genomic stretches of loss of heterozygosity (LOH) and unmasking of mutations that have little or no effect in heterozygous state. Therefore, LOH might be an important force for inducing variation among asexual offspring and may contribute to the limited longevity of asexual lineages. To investigate the genetic consequences of asexuality, here we used high-throughput sequencing of Daphnia magna for assessing the rate of LOH over a single generation of asexual reproduction. Comparing parthenogenetic daughters with their mothers at several thousand genetic markers generated by Restriction site Associated DNA (RAD) sequencing resulted in high LOH rate estimation that largely overlapped with our estimates for the error rate. To distinguish these two, we Sanger re-sequenced the top 18 candidate RAD-loci for LOH, and all of them proved to be false positives. Hence, even though we cannot exclude the possibility that short stretches of LOH occur in genomic regions not covered by our markers, we conclude that LOH does not occur frequently during asexual reproduction in D. magna and ameiotic crossovers are very rare or absent. This finding suggests that clonal lineages of D. magna will remain genetically homogeneous at least over time periods typically relevant for experimental work

Transition from Environmental to Partial Genetic Sex Determination in Daphnia through the Evolution of a Female-Determining Incipient W Chromosome

Sex chromosomes can evolve during the evolution of genetic sex determination (GSD) from environmental sex determination (ESD). Despite theoretical attention, early mechanisms involved in the transition from ESD to GSD have yet to be studied in nature. No mixed ESD-GSD animal species have been reported, except for some species of Daphnia, small freshwater crustaceans in which sex is usually determined solely by the environment, but in which a dominant female sex-determining locus is present in some populations. This locus follows Mendelian single-locus inheritance, but has otherwise not been characterized genetically. We now show that the sex-determining genomic region maps to the same low-recombining peri-centromeric region of linkage group 3 (LG3) in three highly divergent populations of D. magna, and spans 3.6 Mb. Despite low levels of recombination, the associated region contains signs of historical recombination, suggesting a role for selection acting on several genes thereby maintaining linkage disequilibrium among the 36 associated SNPs. The region carries numerous genes involved in sex differentiation in other taxa, including transformer2 and sox9. Taken together, the region determining the genetic females shows characteristics of a sex-related supergene, suggesting that LG3 is potentially an incipient W chromosome despite the lack of significant additional restriction of recombination between Z and W. The occurrence of the female-determining locus in a pre-existing low recombining region illustrates one possible form of recombination suppression in sex chromosomes. D. magna is a promising model for studying the evolutionary transitions from ESD to GSD and early sex chromosome evolution