RAD-seq reveals genetic structure of the F2-generation of natural willow hybrids (Salix L.) and a great potential for interspecific introgression

Hybridization of species with porous genomes can eventually lead to introgression via repeated backcrossing. The potential for introgression between species is reflected by the extent of segregation distortion in later generation hybrids.
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RAD-seq reveals genetic structure of the F2-generation of natural willow hybrids (Salix L.) and a great potential for interspecific introgressionGramlich et al. BMC Plant Biology (2018) 18:317https://doi.org/10.1186/s12870-018-1552-6 RESEARCH ARTICLE Open AccessRAD-seq reveals genetic structure ofthe F2-generation of natural willowhybrids (Salix L.) and a great potentialfor interspecific introgressionSusanne Gramlich* , Natascha Dorothea Wagner and Elvira Hörandl Abstract Background: Hybridization of species with porous genomes can eventually lead to introgression via repeated backcrossing. The potential for introgression between species is reflected by the extent of segregation distortion in later generation hybrids. Here we studied a population of hybrids between Salix purpurea and S. helvetica that has emerged within the last 30 years on a glacier forefield in the European Alps due to secondary contact of the parental species. We used 5758 biallelic SNPs produced by RAD sequencing with the aim to ascertain the predominance of backcrosses (F1 hybrid x parent) or F2 hybrids (F1 hybrid x F1 hybrid) among hybrid offspring. Further, the SNPs were used to study segregation distortion in the second hybrid generation. Results: The analyses in STRUCTURE and NewHybrids revealed that the population consisted of parents and F1 hybrids, whereas hybrid offspring consisted mainly of backcrosses to either parental species, but also some F2 hybrids. Although there was a clear genetic differentiation between S. purpurea and S. helvetica (FST = 0.24), there was no significant segregation distortion in the backcrosses or the F2 hybrids. Plant height of the backcrosses resembled the respective parental species, whereas F2 hybrids were more similar to the subalpine S. helvetica. Conclusions: The co-occurrence of the parental species and the hybrids on the glacier forefield, the high frequency of backcrossing, and the low resistance to gene flow via backcrossing make a scenario of introgression in this young hybrid population highly likely, potentially leading to the transfer of adaptive traits. We further suggest that this willow hybrid population may serve as a model for the evolutionary processes initiated by recent global warming. Keywords: Population genomics, Hybrid evolution, Population structure, Sex chromosomes, Climate changeBackground of previously allopatric species [6]. The absence ofNatural hybridization due to secondary contact has been strong pre- or postzygotic reproductive barriers mayobserved in many plant and animal species. Especially in then lead to hybridization.North America and Northern and Central Europe, a Many studies have investigated the evolutionary rele-major driving force for secondary contact is the ongoing vance of hybridization. Although there are some docu-recolonization after the retreat of glaciers [1]. This mented cases of homoploid hybrid speciation [7–9],process is amplified by human-induced global warming speciation seems to be a rather rare outcome comparedthat also causes rapid range shifts of species [2–5], espe- to the plenty of reported incidents of hybridization [10].cially in mountain regions, leading to secondary contact Important requirements for homoploid hybrid speciation seem to be strong ecological or geographical barriers that restrict gene flow between hybrids and the parental* Correspondence: susanne.gramlich@biologie.uni-goettingen.deDepartment of Systematics, Biodiversity and Evolution of Plants (with species [7, 11–13]. Thus, hybrid speciation is closelyHerbarium), University Goettingen, Untere Karspüle 2, 37073 Goettingen, connected to the availability of novel or extreme habitatsGermany © The Author(s). 2018 Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver (http://creativecommons.org/publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated.Gramlich et al. BMC Plant Biology (2018) 18:317 Page 2 of 12[14, 15]. Chromosomal rearrangements can also rapidly common phenomenon in Salix [42, 43]. Although S. pur-establish crossing barriers between parents and hybrids purea and S. helvetica belong to different sections of the[9]. Generally, interspecific hybridization seems more genus [44], they form natural hybrid zones in thelikely to result in introgression than in speciation [7, 14]. European Alps [39]. The composition of such hybridIn extreme cases, introgression can lead to ge ...