Browsing by Author "He, Li"

Dioecy is rare in flowering plants (5–6% of species), but is often controlled genetically by sex-linked regions (SLRs). It has so far been unclear whether, polyploidy affects sex chromosome evolution, as it does in animals, though polyploidy is quite common in angiosperms, including in dioecious species. Plants could be different, as, unlike many animal systems, degenerated sex chromosomes, are uncommon in plants. Here we consider sex determination in plants and plant-specific factors, and propose that constraints created at the origin of polyploids limit successful polyploidization of species with SLRs. We consider the most likely case of a polyploid of a dioecious diploid with an established SLR, and discuss the outcome in autopolyploids and allopolyploids. The most stable system possibly has an SLR on just one chromosome, with a strongly dominant genetic factor in the heterogametic sex (e.g., xxxY male in a tetraploid). If recombination occurs with its homolog, this will prevent Y chromosome degeneration. Polyploidy may also allow for reversibility of multiplied Z or X chromosomes into autosomes. Otherwise, low dosage of Y-linked SLRs compared to their multiple homologous x copies may cause loss of reliable sex-determination at higher ploidy levels. We discuss some questions that can be studied using genome sequencing, chromosome level-assemblies, gene expression studies and analysis of loci under selection.

Summary In the Vetrix clade of Salix , a genus of woody flowering plants, sex determination involves chromosome 15, but an XY system has changed to a ZW system. We studied the detailed genetic changes involved. We used genome sequencing, with chromosome conformation capture (Hi‐C) and PacBio HiFi reads to assemble chromosome level gap‐free X and Y of Salix arbutifolia , and distinguished the haplotypes in the 15X‐ and 15Y‐linked regions, to study the evolutionary history of the sex‐linked regions (SLRs). Our sequencing revealed heteromorphism of the X and Y haplotypes of the SLR, with the X‐linked region being considerably larger than the corresponding Y region, mainly due to accumulated repetitive sequences and gene duplications. The phylogenies of single‐copy orthogroups within the SLRs indicate that S. arbutifolia and Salix purpurea share an ancestral SLR within a repeat‐rich region near the chromosome 15 centromere. During the change in heterogamety, the X‐linked region changed to a W‐linked one, while the Z was derived from the Y.

Polyploidy is common in the genus Salix. However, little is known about the origin, parentage and genomic composition of polyploid species because of a lack of suitable molecular markers and analysis tools. We established a phylogenomic framework including species of all described sections of Eurasian shrub willows. We analyzed the genomic composition of seven polyploid willow species in comparison to putative diploid parental species to draw conclusions on their origin and the effects of backcrossing and post-origin evolution. We applied recently developed programs like SNAPP, HyDe, and SNiPloid to establish a bioinformatic pipeline for unravelling the complexity of polyploid genomes. RAD sequencing revealed 23,393 loci and 320,010 high quality SNPs for the analysis of relationships of 35 species of Eurasian shrub willows (Salix subg. Chamaetia/Vetrix). Polyploid willow species appear to be predominantly of allopolyploid origin. More ancient allopolyploidization events were observed for two hexaploid and one octoploid species, while our data suggested a more recent allopolyploid origin for the included tetraploids and identified putative parental taxa. SNiPloid analyses disentangled the different genomic signatures resulting from hybrid origin, backcrossing, and secondary post-origin evolution in the polyploid species. Our RAD sequencing data demonstrate that willow genomes are shaped by ancient and recent reticulate evolution, polyploidization, and post-origin divergence of species.

Abstract Almost all species in the genus Salix (willow) are dioecious and willows have variable sex-determining systems, the role of this variation in maintaining species barriers is relatively untested. We first analyzed the sex determination systems (SDS) of two species, Salix cardiophylla and Salix interior, whose positions in the Salix phylogeny make them important for understanding a sex chromosome turnover that has been detected in their relatives, and that changed the system from male (XX/XY) to female (ZW/ZZ) heterogamety. We show that both species have male heterogamety, with sex-linked regions (SLRs) on chromosome 15 (termed a 15XY system). The SLRs occupy 21.3% and 22.8% of the entire reference chromosome, respectively. By constructing phylogenetic trees, we determined the phylogenetic positions of all the species with known SDSs. Reconstruction of ancestral SDS character states revealed that the 15XY system is likely the ancestral state in willows. Turnovers of 15XY to 15ZW and 15XY to 7XY likely contributed to early speciation in Salix and gave rise to major groups of the Vetrix and Salix clades. Finally, we tested introgression among species in the phylogenetic trees based on both autosomes and SLRs separately. Frequent introgression was observed among species with 15XY, 15ZW, and 7XY on autosomes, in contrast to the SLR datasets, which showed less introgression, and in particular no gene flow between 15ZW and 7XY species. We argue that, although SDS turnovers in willow speciation may not create complete reproductive barriers, the evolution of SLRs plays important roles in preventing introgression and maintaining species boundaries.

The genus Salix (willows), with 33 species, represents the most diverse genus of woody plants in the European Alps. Many species dominate subalpine and alpine types of vegetation. Despite a long history of research on willows, the evolutionary and ecological factors for this species richness are poorly known. Here we will review recent progress in research on phylogenetic relationships, evolution, ecology, and speciation in alpine willows. Phylogenomic reconstructions suggest multiple colonization of the Alps, probably from the late Miocene onward, and reject hypotheses of a single radiation. Relatives occur in the Arctic and in temperate Eurasia. Most species are widespread in the European mountain systems or in the European lowlands. Within the Alps, species differ ecologically according to different elevational zones and habitat preferences. Homoploid hybridization is a frequent process in willows and happens mostly after climatic fluctuations and secondary contact. Breakdown of the ecological crossing barriers of species is followed by introgressive hybridization. Polyploidy is an important speciation mechanism, as 40% of species are polyploid, including the four endemic species of the Alps. Phylogenomic data suggest an allopolyploid origin for all taxa analyzed so far. Further studies are needed to specifically analyze biogeographical history, character evolution, and genome evolution of polyploids.

Abstract Background and Aims The largest genus of Salicaceae sensu lato, Salix, has been shown to consist of two main clades: clade Salix, in which species have XY sex-determination systems (SDSs) on chromosome 7, and clade Vetrix including species with ZW SDSs on chromosome 15. Here, we test the utility of whole genome re-sequencing (WGR) for phylogenomic reconstructions of willows to infer changes between different SDSs. Methods We used more than 1 TB of WGR data from 70 Salix taxa to ascertain single nucleotide polymorphisms on the autosomes, the sex-linked regions (SLRs) and the chloroplast genomes, for phylogenetic and species tree analyses. To avoid bias, we chose reference genomes from both groups, Salix dunnii from clade Salix and S. purpurea from clade Vetrix. Key Results Two main largely congruent groups were recovered: the paraphyletic Salix grade and the Vetrix clade. The autosome dataset trees resolved four subclades (C1–C4) in Vetrix. C1 and C2 comprise species from the Hengduan Mountains and adjacent areas and from Eurasia, respectively. Section Longifoliae (C3) grouped within the Vetrix clade but fell into the Salix clade in trees based on the chloroplast dataset analysis. Salix triandra from Eurasia (C4) was revealed as sister to the remaining species of clade Vetrix. In Salix, the polyploid group C5 is paraphyletic to clade Vetrix and subclade C6 is consistent with Argus’s subgenus Protitea. Chloroplast datasets separated both Vetrix and Salix as monophyletic, and yielded C5 embedded within Salix. Using only diploid species, both the SLR and autosomal datasets yielded trees with Vetrix and Salix as well-supported clades. Conclusion WGR data are useful for phylogenomic analyses of willows. The different SDSs may contribute to the isolation of the two major groups, but the reproductive barrier between them needs to be studied.