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Lagercrantz, Ulf
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Kryvokhyzha, D., Salcedo, A., Eriksson, M. C., Duan, T., Tawari, N., Chen, J., . . . Lascoux, M. (2019). Parental legacy, demography, and admixture influenced the evolution of the two subgenomes of the tetraploid Capsella bursa-pastoris (Brassicaceae). PLoS Genetics, 15(2), Article ID e1007949.
Open this publication in new window or tab >>Parental legacy, demography, and admixture influenced the evolution of the two subgenomes of the tetraploid Capsella bursa-pastoris (Brassicaceae)
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2019 (English)In: PLoS Genetics, ISSN 1553-7390, E-ISSN 1553-7404, Vol. 15, no 2, article id e1007949Article in journal (Refereed) Published
Abstract [en]

Allopolyploidy is generally perceived as a major source of evolutionary novelties and as an instantaneous way to create isolation barriers. However, we do not have a clear understanding of how two subgenomes evolve and interact once they have fused in an allopolyploid species nor how isolated they are from their relatives. Here, we address these questions by analyzing genomic and transcriptomic data of allotetraploid Capsella bursa-pastoris in three differentiated populations, Asia, Europe, and the Middle East. We phased the two subgenomes, one descended from the outcrossing and highly diverse Capsella grandiflora (Cbp(Cg)) and the other one from the selfing and genetically depauperate Capsella orientalis (Cbp(Co)). For each subgenome, we assessed its relationship with the diploid relatives, temporal changes of effective population size (N-e), signatures of positive and negative selection, and gene expression patterns. In all three regions, N-e of the two subgenomes decreased gradually over time and the Cbp(Co) subgenome accumulated more deleterious changes than Cbp(Cg). There were signs of widespread admixture between C. bursa-pastoris and its diploid relatives. The two subgenomes were impacted differentially depending on geographic region suggesting either strong interploidy gene flow or multiple origins of C. bursa-pastoris. Selective sweeps were more common on the Cbp(Cg) subgenome in Europe and the Middle East, and on the Cbp(Co) subgenome in Asia. In contrast, differences in expression were limited with the Cbp(Cg) subgenome slightly more expressed than Cbp(Co) in Europe and the Middle-East. In summary, after more than 100,000 generations of co-existence, the two subgenomes of C. bursa-pastoris still retained a strong signature of parental legacy but their evolutionary trajectory strongly varied across geographic regions. Author summary Allopolyploid species have two or more sets of chromosomes that originate from hybridization of different species. It remains largely unknown how the two genomes evolve in the same organism and how strongly their evolutionary trajectory depends on the initial differences between the two parental species and the specific demographic history of the newly formed allopolyploid species. To address these questions, we analyzed the genomic and gene expression variation of the shepherd's purse, a recent allopolyploid species, in three regions of its natural range. After approximate to 100,000 generations of co-existence within the same species, the two subgenomes had still retained part of the initial difference between the two parental species in the number of deleterious mutations reflecting a history of mating system differences. This difference, as well as differences in patterns of positive selection and levels of gene expression, also strongly depended on the specific histories of the three regions considered. Most strikingly, and unexpectedly, the allopolyploid species showed signs of hybridization with different diploid relatives or multiple origins in different parts of its range. Regardless if it was hybridization or multiple origins, this profoundly altered the relationship between the two subgenomes in different regions. Hence, our study illustrates how both the genomic structure and ecological arena interact to determine the evolutionary trajectories of allopolyploid species.

National Category
Evolutionary Biology
Identifiers
urn:nbn:se:uu:diva-379946 (URN)10.1371/journal.pgen.1007949 (DOI)000459970100033 ()30768594 (PubMedID)
Funder
Swedish Research Council, 2012-04999Swedish Research Council, 2015-03797
Note

De 2 första författarna delar förstaförfattarskapet.

Available from: 2019-03-25 Created: 2019-03-25 Last updated: 2019-03-25Bibliographically approved
Eklund, D. M., Kanei, M., Flores-Sandoval, E., Ishizaki, K., Nishihama, R., Kohchi, T., . . . Bowman, J. L. (2018). An Evolutionarily Conserved Abscisic Acid Signaling Pathway Regulates Dormancy in the Liverwort Marchantia polymorpha. Current Biology, 28(22), 3691-3699.e3
Open this publication in new window or tab >>An Evolutionarily Conserved Abscisic Acid Signaling Pathway Regulates Dormancy in the Liverwort Marchantia polymorpha
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2018 (English)In: Current Biology, ISSN 0960-9822, E-ISSN 1879-0445, Vol. 28, no 22, p. 3691-3699.e3Article in journal (Refereed) Published
Abstract [en]

Dormancy is a key process allowing land plants to adapt to changing conditions in the terrestrial habitat, allowing the cessation of growth in response to environmental or physiological cues, entrance into a temporary quiescent state, and subsequent reactivation of growth in more favorable environmental conditions [1-3]. Dormancy may be induced seasonally, sporadically (e.g., in response to drought), or developmentally (e.g., seeds and apical dominance). Asexual propagules, known as gemmae, derived via clonal reproduction in bryophytes, are often dormant until displaced from the parent plant. In the liverwort Marchantia polymorpha, gemmae are produced within specialized receptacles, gemma cups, located on the dorsal side of the vegetative thallus [4]. Mature gemmae are detached from the parent plant but may remain in the cup, with gemma growth suppressed as long as the gemmae remain in the gemma cup and the parental plant is alive [5]. Following dispersal of gemmae from gemma cups by rain, the gemmae germinate in the presence of light and moisture, producing clonal offspring [6]. In land plants, the plant hormone abscisic acid (ABA) regulates many aspects of dormancy and water balance [7]. Here, we demonstrate that ABA plays a central role in the control of gemma dormancy as transgenic M. polymorpha gemmae with reduced sensitivity to ABA fail to establish and/or maintain dormancy. Thus, the common ancestor of land plants used the ABA signaling module to regulate germination of progeny in response to environmental cues, with both gemmae and seeds being derived structures co-opting an ancestral response system.

Place, publisher, year, edition, pages
CELL PRESS, 2018
National Category
Botany
Identifiers
urn:nbn:se:uu:diva-371870 (URN)10.1016/j.cub.2018.10.018 (DOI)000450614200032 ()30416060 (PubMedID)
Funder
Australian Research Council, DP130100177Australian Research Council, DP160100892Swedish Research Council, 623-2010-6591Swedish Research Council, 2016-05180Swedish Research Council, 201604430
Available from: 2019-01-07 Created: 2019-01-07 Last updated: 2019-01-07Bibliographically approved
Linde, A.-M., Eklund, D. M., Kubota, A., Pederson, E. R. A., Holm, K., Gyllenstrand, N., . . . Lagercrantz, U. (2017). Early evolution of the land plant circadian clock. New Phytologist, 216(2), 576-590
Open this publication in new window or tab >>Early evolution of the land plant circadian clock
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2017 (English)In: New Phytologist, ISSN 0028-646X, E-ISSN 1469-8137, Vol. 216, no 2, p. 576-590Article in journal (Refereed) Published
Abstract [en]

While angiosperm clocks can be described as an intricate network of interlocked transcriptional feedback loops, clocks of green algae have been modelled as a loop of only two genes. To investigate the transition from a simple clock in algae to a complex one in angiosperms, we performed an inventory of circadian clock genes in bryophytes and charophytes. Additionally, we performed functional characterization of putative core clock genes in the liverwort Marchantia polymorpha and the hornwort Anthoceros agrestis. Phylogenetic construction was combined with studies of spatiotemporal expression patterns and analysis of M. polymorpha clock gene mutants. Homologues to core clock genes identified in Arabidopsis were found not only in bryophytes but also in charophytes, albeit in fewer copies. Circadian rhythms were detected for most identified genes in M. polymorpha and A. agrestis, and mutant analysis supports a role for putative clock genes in M. polymorpha. Our data are in line with a recent hypothesis that adaptation to terrestrial life occurred earlier than previously expected in the evolutionary history of charophyte algae. Both gene duplication and acquisition of new genes was important in the evolution of the plant circadian clock, but gene loss has also contributed to shaping the clock of bryophytes.

Keywords
Marchantia polymorpha, bryophyte, circadian clock, evolution, transcription factor
National Category
Botany Evolutionary Biology Biochemistry and Molecular Biology Developmental Biology
Research subject
Biology with specialization in Evolutionary Functional Genomics
Identifiers
urn:nbn:se:uu:diva-317912 (URN)10.1111/nph.14487 (DOI)000427294000024 ()28244104 (PubMedID)
Funder
Swedish Research Council, 2011-5609Swedish Research Council, 2014-522
Available from: 2017-03-22 Created: 2017-03-22 Last updated: 2019-09-05Bibliographically approved
Bowman, J. L., Kohchi, T., Yamato, K. T., Jenkins, J., Shu, S., Ishizaki, K., . . . Schmutz, J. (2017). Insights into Land Plant Evolution Garnered from the Marchantia polymorpha Genome. Cell, 171(2), 287-304.15
Open this publication in new window or tab >>Insights into Land Plant Evolution Garnered from the Marchantia polymorpha Genome
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2017 (English)In: Cell, ISSN 0092-8674, E-ISSN 1097-4172, Vol. 171, no 2, p. 287-304.15Article in journal (Refereed) Published
Abstract [en]

The evolution of land flora transformed the terrestrial environment. Land plants evolved from an ancestral charophycean alga from which they inherited developmental, biochemical, and cell biological attributes. Additional biochemical and physiological adaptations to land, and a life cycle with an alternation between multicellular haploid and diploid generations that facilitated efficient dispersal of desiccation tolerant spores, evolved in the ancestral land plant. We analyzed the genome of the liverwort Marchantia polymorpha, a member of a basal land plant lineage. Relative to charophycean algae, land plant genomes are characterized by genes encoding novel biochemical pathways, new phytohormone signaling pathways (notably auxin), expanded repertoires of signaling pathways, and increased diversity in some transcription factor families. Compared with other sequenced land plants, M. polymorpha exhibits low genetic redundancy in most regulatory pathways, with this portion of its genome resembling that predicted for the ancestral land plant. PAPERCLIP.

Keywords
Marchantia polymorpha, auxin, charophycean algae, land plant evolution, sex chromosome
National Category
Evolutionary Biology
Identifiers
urn:nbn:se:uu:diva-333225 (URN)10.1016/j.cell.2017.09.030 (DOI)000412346100008 ()28985561 (PubMedID)
Available from: 2017-11-09 Created: 2017-11-09 Last updated: 2018-01-10Bibliographically approved
Hallingback, H. R., Fogelqvist, J., Powers, S. J., Turrion-Gomez, J., Rossiter, R., Amey, J., . . . Roennberg-Wastljung, A.-C. (2016). Association mapping in Salix viminalis L. (Salicaceae) - identification of candidate genes associated with growth and phenology. Global Change Biology Bioenergy, 8(3), 670-685
Open this publication in new window or tab >>Association mapping in Salix viminalis L. (Salicaceae) - identification of candidate genes associated with growth and phenology
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2016 (English)In: Global Change Biology Bioenergy, ISSN 1757-1693, E-ISSN 1757-1707, Vol. 8, no 3, p. 670-685Article in journal (Refereed) Published
Abstract [en]

Willow species (Salix) are important as short-rotation biomass crops for bioenergy, which creates a demand for faster genetic improvement and breeding through deployment of molecular marker-assisted selection (MAS). To find markers associated with important adaptive traits, such as growth and phenology, for use in MAS, we genetically dissected the trait variation of a Salix viminalis (L.) population of 323 accessions. The accessions were sampled throughout northern Europe and were established at two field sites in Pustnas, Sweden, and at Woburn, UK, offering the opportunity to assess the impact of genotype-by-environment interactions (GxE) on trait-marker associations. Field measurements were recorded for growth and phenology traits. The accessions were genotyped using 1536 SNP markers developed from phenology candidate genes and from genes previously observed to be differentially expressed in contrasting environments. Association mapping between 1233 of these SNPs and the measured traits was performed taking into account population structure and threshold selection bias. At a false discovery rate (FDR) of 0.2, 29 SNPs were associated with bud burst, leaf senescence, number of shoots or shoot diameter. The percentage of accession variation explained by these associations ranged from 0.3% to 4.4%, suggesting that the studied traits are controlled by many loci of limited individual impact. Despite this, a SNP in the EARLY FLOWERING 3 gene was repeatedly associated (FDR<0.2) with bud burst. The rare homozygous genotype exhibited 0.4-1.0 lower bud burst scores than the other genotype classes on a five-grade scale. Consequently, this marker could be promising for use in MAS and the gene deserves further study. Otherwise, associations were less consistent across sites, likely due to their small estimates and to considerable GxE interactions indicated by multivariate association analyses and modest trait accession correlations across sites (0.32-0.61).

Keywords
adaptation, association mapping, candidate gene, growth, marker-assisted selection, phenology, Salix, short-rotation coppice, willow
National Category
Wood Science Bioenergy
Identifiers
urn:nbn:se:uu:diva-300297 (URN)10.1111/gcbb.12280 (DOI)000374060500013 ()
Funder
Swedish Research CouncilKnut and Alice Wallenberg FoundationSwedish Energy Agency, 31468-1;30599-2
Available from: 2016-08-08 Created: 2016-08-08 Last updated: 2017-11-28Bibliographically approved
Kryvokhyzha, D., Holm, K., Chen, J., Cornille, A., Glemin, S., Wright, S. I., . . . Lascoux, M. (2016). The influence of population structure on gene expression and flowering time variation in the ubiquitous weed Capsella bursa-pastoris (Brassicaceae). Molecular Ecology, 25(5), 1106-1121
Open this publication in new window or tab >>The influence of population structure on gene expression and flowering time variation in the ubiquitous weed Capsella bursa-pastoris (Brassicaceae)
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2016 (English)In: Molecular Ecology, ISSN 0962-1083, E-ISSN 1365-294X, Vol. 25, no 5, p. 1106-1121Article in journal (Refereed) Published
Abstract [en]

Population structure is a potential problem when testing for adaptive phenotypic differences among populations. The observed phenotypic differences among populations can simply be due to genetic drift, and if the genetic distance between them is not considered, the differentiation may be falsely interpreted as adaptive. Conversely, adaptive and demographic processes might have been tightly associated and correcting for the population structure may lead to false negatives. Here, we evaluated this problem in the cosmopolitan weed Capsella bursa-pastoris. We used RNA-Seq to analyse gene expression differences among 24 accessions, which belonged to a much larger group that had been previously characterized for flowering time and circadian rhythm and were genotyped using genotyping-by-sequencing (GBS) technique. We found that clustering of accessions for gene expression retrieved the same three clusters that were obtained with GBS data previously, namely Europe, the Middle East and Asia. Moreover, the three groups were also differentiated for both flowering time and circadian rhythm variation. Correction for population genetic structure when analysing differential gene expression analysis removed all differences among the three groups. This may suggest that most differences are neutral and simply reflect population history. However, geographical variation in flowering time and circadian rhythm indicated that the distribution of adaptive traits might be confounded by population structure. To bypass this confounding effect, we compared gene expression differentiation between flowering ecotypes within the genetic groups. Among the differentially expressed genes, FLOWERING LOCUS C was the strongest candidate for local adaptation in regulation of flowering time.

Keywords
adaptation, circadian rhythm, genetic drift, neutrality, population structure, RNA-Seq, transcriptome
National Category
Biochemistry and Molecular Biology Evolutionary Biology
Identifiers
urn:nbn:se:uu:diva-282478 (URN)10.1111/mec.13537 (DOI)000371433400007 ()26797895 (PubMedID)
Funder
Swedish Research Council
Available from: 2016-04-05 Created: 2016-04-05 Last updated: 2018-02-13Bibliographically approved
Eklund, D. M., Ishizaki, K., Flores-Sandoval, E., Kikuchi, S., Takebayashi, Y., Tsukamoto, S., . . . Bowman, J. L. (2015). Auxin Produced by the Indole-3-Pyruvic Acid Pathway Regulates Development and Gemmae Dormancy in the Liverwort Marchantia polymorph. The Plant Cell, 27(6), 1650-1669
Open this publication in new window or tab >>Auxin Produced by the Indole-3-Pyruvic Acid Pathway Regulates Development and Gemmae Dormancy in the Liverwort Marchantia polymorph
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2015 (English)In: The Plant Cell, ISSN 1040-4651, E-ISSN 1532-298X, Vol. 27, no 6, p. 1650-1669Article in journal (Refereed) Published
Abstract [en]

The plant hormone auxin (indole-3-acetic acid [IAA]) has previously been suggested to regulate diverse forms of dormancy in both seed plants and liverworts. Here, we use loss-and gain-of-function alleles for auxin synthesis-and signaling-related genes, as well as pharmacological approaches, to study how auxin regulates development and dormancy in the gametophyte generation of the liverwort Marchantia polymorpha. We found that M. polymorpha possess the smallest known toolkit for the indole-3-pyruvic acid (IPyA) pathway in any land plant and that this auxin synthesis pathway mainly is active in meristematic regions of the thallus. Previously a Trp-independent auxin synthesis pathway has been suggested to produce a majority of IAA in bryophytes. Our results indicate that the Trp-dependent IPyA pathway produces IAA that is essential for proper development of the gametophyte thallus of M. polymorpha. Furthermore, we show that dormancy of gemmae is positively regulated by auxin synthesized by the IPyA pathway in the apex of the thallus. Our results indicate that auxin synthesis, transport, and signaling, in addition to its role in growth and development, have a critical role in regulation of gemmae dormancy in M. polymorpha.

National Category
Biochemistry and Molecular Biology
Identifiers
urn:nbn:se:uu:diva-260155 (URN)10.1105/tpc.15.00065 (DOI)000357718800009 ()26036256 (PubMedID)
Available from: 2015-08-17 Created: 2015-08-17 Last updated: 2017-12-04Bibliographically approved
Avia, K., Karkkainen, K., Lagercrantz, U. & Savolainen, O. (2014). Association of FLOWERING LOCUS T/TERMINAL FLOWER 1-like gene FTL2 expression with growth rhythm in Scots pine (Pinus sylvestris). New Phytologist, 204(1), 159-170
Open this publication in new window or tab >>Association of FLOWERING LOCUS T/TERMINAL FLOWER 1-like gene FTL2 expression with growth rhythm in Scots pine (Pinus sylvestris)
2014 (English)In: New Phytologist, ISSN 0028-646X, E-ISSN 1469-8137, Vol. 204, no 1, p. 159-170Article in journal (Refereed) Published
Abstract [en]

Understanding the genetic basis of the timing of bud set, an important trait in conifers, is relevant for adaptation and forestry practice. In common garden experiments, both Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) show a latitudinal cline in the trait. We compared the regulation of their bud set biology by examining the expression of PsFTL2, a Pinus sylvestris homolog to PaFTL2, a FLOWERING LOCUS T/TERMINAL FLOWER 1 (FT/TFL1)-like gene, the expression levels of which have been found previously to be associated with the timing of bud set in Norway spruce. In a common garden study, we analyzed the relationship of bud phenology under natural and artificial photoperiods and the expression of PsFTL2 in a set of Scots pine populations from different latitudes. The expression of PsFTL2 increased in the needles preceding bud set and decreased during bud burst. In the northernmost population, even short night periods were efficient to trigger this expression, which also increased earlier under all photoperiodic regimes compared with the southern populations. Despite the different biology, with few limitations, the two conifers that diverged 140 million yr ago probably share an association of FTL2 with bud set, pointing to a common mechanism for the timing of growth cessation in conifers.

Keywords
bud burst, bud set, CENTRORADIALIS/TERMINAL FLOWER 1/SELF-PRUNING (CETS), conifers, FLOWERING LOCUS T/TERMINAL FLOWER 1-like 2 (FTL2), gene expression, Pinus sylvestris
National Category
Ecology
Identifiers
urn:nbn:se:uu:diva-232983 (URN)10.1111/nph.12901 (DOI)000341193500018 ()
Available from: 2014-10-14 Created: 2014-09-29 Last updated: 2017-12-05Bibliographically approved
Ghelardini, L., Berlin, S., Weih, M., Lagercrantz, U., Gyllenstrand, N. & Ronnberg-Wastljung, A. C. (2014). Genetic architecture of spring and autumn phenology in Salix. BMC Plant Biology, 14, 31
Open this publication in new window or tab >>Genetic architecture of spring and autumn phenology in Salix
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2014 (English)In: BMC Plant Biology, ISSN 1471-2229, E-ISSN 1471-2229, Vol. 14, p. 31-Article in journal (Refereed) Published
Abstract [en]

Background: In woody plants from temperate regions, adaptation to the local climate results in annual cycles of growth and dormancy, and optimal regulation of these cycles are critical for growth, long-term survival, and competitive success. In this study we have investigated the genetic background to growth phenology in a Salix pedigree by assessing genetic and phenotypic variation in growth cessation, leaf senescence and bud burst in different years and environments. A previously constructed linkage map using the same pedigree and anchored to the annotated genome of P. trichocarpa was improved in target regions and used for QTL analysis of the traits. The major aims in this study were to map QTLs for phenology traits in Salix, and to identify candidate genes in QTL hot spots through comparative mapping with the closely related Populus trichocarpa. Results: All traits varied significantly among genotypes and the broad-sense heritabilities ranged between 0.5 and 0.9, with the highest for leaf senescence. In total across experiment and years, 80 QTLs were detected. For individual traits, the QTLs explained together from 21.5 to 56.5% of the variation. Generally each individual QTL explained a low amount of the variation but three QTLs explained above 15% of the variation with one QTL for leaf senescence explaining 34% of the variation. The majority of the QTLs were recurrently identified across traits, years and environments. Two hotspots were identified on linkage group (LG) II and X where narrow QTLs for all traits co-localized. Conclusions: This study provides the most detailed analysis of QTL detection for phenology in Salix conducted so far. Several hotspot regions were found where QTLs for different traits and QTLs for the same trait but identified during different years co-localised. Many QTLs co-localised with QTLs found in poplar for similar traits that could indicate common pathways for these traits in Salicaceae. This study is an important first step in identifying QTLs and candidate genes for phenology traits in Salix.

Keywords
Phenology, Adaptation, Salix, QTL, Candidate genes
National Category
Ecology
Identifiers
urn:nbn:se:uu:diva-224929 (URN)10.1186/1471-2229-14-31 (DOI)000334471800001 ()
Available from: 2014-05-28 Created: 2014-05-23 Last updated: 2017-12-05Bibliographically approved
Berlin, S., Trybush, S. O., Fogelqvist, J., Gyllenstrand, N., Hallingbaeck, H. R., Ahman, I., . . . Hanley, S. J. (2014). Genetic diversity, population structure and phenotypic variation in European Salix viminalis L. (Salicaceae). Tree Genetics & Genomes, 10(6), 1595-1610
Open this publication in new window or tab >>Genetic diversity, population structure and phenotypic variation in European Salix viminalis L. (Salicaceae)
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2014 (English)In: Tree Genetics & Genomes, ISSN 1614-2942, E-ISSN 1614-2950, Vol. 10, no 6, p. 1595-1610Article in journal (Refereed) Published
Abstract [en]

To investigate the potential of association genetics for willow breeding, Salix viminalis germplasm was assembled from UK and Swedish collections (comprising accessions from several European countries) and new samples collected from nature. A subset of the germplasm was planted at two sites (UK and Sweden), genotyped using 38 SSR markers and assessed for phenological and biomass traits. Population structure, genetic differentiation (F-ST) and quantitative trait differentiation (Q(ST)) were investigated. The extent and patterns of trait adaptation were assessed by comparing F-ST and Q(ST) parameters. Of the 505 genotyped diploid accessions, 27 % were not unique. Genetic diversity was high: 471 alleles was amplified; the mean number of alleles per locus was 13.46, mean observed heterozygosity was 0.55 and mean expected heterozygosity was 0.62. Bayesian clustering identified four subpopulations which generally corresponded to Western Russia, Western Europe, Eastern Europe and Sweden. All pairwise F-ST values were highly significant (p<0.001) with the greatest genetic differentiation detected between the Western Russian and the Western European subpopulations (F-ST = 0.12), and the smallest between the Swedish and Eastern European populations (F-ST = 0.04). The Swedish population also had the highest number of identical accessions, supporting the view that S. viminalis was introduced into this country and has been heavily influenced by humans. Q(ST) values were high for growth cessation and leaf senescence, and to some extent stem diameter, but low for bud burst time and shoot number. Overall negative clines between longitudinal coordinates and leaf senescence, bud burst and stem diameter were also found.

Keywords
Salix, Population structure, Adaptation, Breeding, SSRs
National Category
Ecology Botany Evolutionary Biology
Identifiers
urn:nbn:se:uu:diva-237890 (URN)10.1007/s11295-014-0782-5 (DOI)000343924600009 ()
Available from: 2014-12-11 Created: 2014-12-08 Last updated: 2017-12-05Bibliographically approved
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