Recent studies show that molecular convergence plays an unexpectedly common role in the evolution of convergent phenotypes. We exploited this phenomenon to find candidate loci underlying resistance to the emerald ash borer (EAB, Agrilus planipennis), the United States’ most costly invasive forest insect to date, within the pan-genome of ash trees (the genus Fraxinus). We show that EAB-resistant taxa occur within three independent phylogenetic lineages. In genomes from these resistant lineages, we detect 53 genes with evidence of convergent amino acid evolution. Gene-tree reconstruction indicates that, for 48 of these candidates, the convergent amino acids are more likely to have arisen via independent evolution than by another process such as hybridization or incomplete lineage sorting. Seven of the candidate genes have putative roles connected to the phenylpropanoid biosynthesis pathway and 17 relate to herbivore recognition, defence signalling or programmed cell death. Evidence for loss-of-function mutations among these candidates is more frequent in susceptible species than in resistant ones. Our results on evolutionary relationships, variability in resistance, and candidate genes for defence response within the ash genus could inform breeding for EAB resistance, facilitating ecological restoration in areas invaded by this beetle.
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The custom scripts used is this study have been deposited in GitHub: https://github.com/lkelly3/eab-ms-scripts.
Pautasso, M., Aas, G., Queloz, V. & Holdenrieder, O. European ash (Fraxinus excelsior) dieback – a conservation biology challenge. Biol. Conserv. 158, 37–49 (2013).
MacFarlane, D. W. & Meyer, S. P. Characteristics and distribution of potential ash tree hosts for emerald ash borer. For. Ecol. Manage. 213, 15–24 (2005).
Boyd, I. L., Freer-Smith, P. H., Gilligan, C. A. & Godfray, H. C. J. The consequence of tree pests and diseases for ecosystem services. Science 342, 1235773 (2013).
Herms, D. A. & McCullough, D. G. Emerald ash borer invasion of North America: history, biology, ecology, impacts, and management. Annu. Rev. Entomol. 59, 13–30 (2014).
Orlova-Bienkowskaja, M. J. Ashes in Europe are in danger: the invasive range of Agrilus planipennis in European Russia is expanding. Biol. Invasions 16, 1345–1349 (2014).
McCullough, D. G. Challenges, tactics and integrated management of emerald ash borer in North America. Forestry 93, 197–211 (2019).
Drogvalenko, A. N., Orlova-Bienkowskaja, M. J. & Bieńkowski, A. O. Record of the emerald ash borer (Agrilus planipennis) in Ukraine is confirmed. Insects 10, 338 (2019).
Semizer-Cuming, D., Krutovsky, K. V., Baranchikov, Y. N., Kjӕr, E. D. & Williams, C. G. Saving the world’s ash forests calls for international cooperation now. Nat. Ecol. Evol. 3, 141–144 (2019).
Evans, H. F., Williams, D., Hoch, G., Loomans, A. & Marzano, M. Developing a European toolbox to manage potential invasion by emerald ash borer (Agrilus planipennis) and bronze birch borer (Agrilus anxius), important pests of ash and birch. Forestry 93, 187–196 (2020).
Baranchikov, Y., Mozolevskaya, E., Yurchenko, G. & Kenis, M. Occurrence of the emerald ash borer, Agrilus planipennis in Russia and its potential impact on European forestry. Bull. OEPP 38, 233–238 (2008).
Zhao, T. et al. Induced outbreaks of indigenous insect species by exotic tree species. Acta Entomol. Sin. 50, 826–833 (2007).
Liu, H. et al. Exploratory survey for the emerald ash borer, Agrilus planipennis (Coleoptera: Buprestidae), and its natural enemies in China. Great Lakes Entomol. 36, 191–204 (2003).
Wei, X. et al. Emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), in China: a review and distribution survey. Acta Entomol. Sin. 47, 679–685 (2004).
Orlova-Bienkowskaja, M. J. & Volkovitsh, M. G. Are native ranges of the most destructive invasive pests well known? A case study of the native range of the emerald ash borer, Agrilus planipennis (Coleoptera: Buprestidae). Biol. Invasions 20, 1275–1286 (2018).
Showalter, D. N., Villari, C., Herms, D. A. & Bonello, P. Drought stress increased survival and development of emerald ash borer larvae on coevolved Manchurian ash and implicates phloem-based traits in resistance. Agric. For. Entomol. 20, 170–179 (2018).
Whitehill, J. G. A. et al. Interspecific proteomic comparisons reveal ash phloem genes potentially involved in constitutive resistance to the emerald ash borer. PLoS ONE 6, e24863 (2011).
Whitehill, J. G. A. et al. Interspecific comparison of constitutive ash phloem phenolic chemistry reveals compounds unique to Manchurian ash, a species resistant to emerald ash borer. J. Chem. Ecol. 38, 499–511 (2012).
Lane, T. et al. The green ash transcriptome and identification of genes responding to abiotic and biotic stresses. BMC Genomics 17, 702 (2016).
Sackton, T. B. et al. Convergent regulatory evolution and loss of flight in paleognathous birds. Science 364, 74–78 (2019).
Arnold, B. J. et al. Borrowed alleles and convergence in serpentine adaptation. Proc. Natl Acad. Sci. USA 113, 8320–8325 (2016).
Hu, Y. et al. Comparative genomics reveals convergent evolution between the bamboo-eating giant and red pandas. Proc. Natl Acad. Sci. USA 114, 1081–1086 (2017).
Yang, X. et al. The Kalanchoë genome provides insights into convergent evolution and building blocks of crassulacean acid metabolism. Nat. Commun. 8, 1899 (2017).
Hill, J. et al. Recurrent convergent evolution at amino acid residue 261 in fish rhodopsin. Proc. Natl Acad. Sci. USA 116, 18473–18478 (2019).
Zhen, Y., Aardema, M. L., Medina, E. M., Schumer, M. & Andolfatto, P. Parallel molecular evolution in an herbivore community. Science 337, 1634–1637 (2012).
Wallander, E. Systematics and floral evolution in Fraxinus (Oleaceae). Belg. Dendrol. Belg. 2012, 39–58 (2012).
Koch, J. L., Carey, D. W., Mason, M. E., Poland, T. M. & Knight, K. S. Intraspecific variation in Fraxinus pennsylvanica responses to emerald ash borer (Agrilus planipennis). New For. (Dordr.) 46, 995–1011 (2015).
Sollars, E. S. A. et al. Genome sequence and genetic diversity of European ash trees. Nature 541, 212–216 (2017).
Cruz, F. et al. Genome sequence of the olive tree, Olea europaea. Gigascience 5, 29 (2016).
Hellsten, U. et al. Fine-scale variation in meiotic recombination in Mimulus inferred from population shotgun sequencing. Proc. Natl Acad. Sci. USA 110, 19478–19482 (2013).
Tomato Genome Consortium. The tomato genome sequence provides insights into fleshy fruit evolution. Nature 485, 635–641 (2012).
Wright, J. W. New chromosome counts in Acer and Fraxinus. Morris Arb. Bull. 8, 33–34 (1957).
Bernards, M. A. & Båstrup-Spohr, L. in Induced Plant Resistance to Herbivory (ed. Schaller, A.) 189–211 (Springer, 2008).
Stahl, E., Hilfiker, O. & Reymond, P. Plant–arthropod interactions: who is the winner? Plant J. 93, 703–728 (2018).
Abdulrazzak, N. et al. A coumaroyl-ester-3-hydroxylase insertion mutant reveals the existence of nonredundant meta-hydroxylation pathways and essential roles for phenolic precursors in cell expansion and plant growth. Plant Physiol. 140, 30–48 (2006).
Rupasinghe, S., Baudry, J. & Schuler, M. A. Common active site architecture and binding strategy of four phenylpropanoid P450s from Arabidopsis thaliana as revealed by molecular modeling. Protein Eng. 16, 721–731 (2003).
Dolan, W. L. & Chapple, C. Conservation and divergence of mediator structure and function: insights from plants. Plant Cell Physiol. 58, 4–21 (2017).
Bonawitz, N. D. et al. Disruption of mediator rescues the stunted growth of a lignin-deficient Arabidopsis mutant. Nature 509, 376–380 (2014).
Dolan, W. L. & Chapple, C. Transcriptome analysis of four Arabidopsis thaliana mediator tail mutants reveals overlapping and unique functions in gene regulation. G3 (Bethesda) 8, 3093–3108 (2018).
Xu, Z. et al. Functional genomic analysis of Arabidopsis thaliana glycoside hydrolase family 1. Plant Mol. Biol. 55, 343–367 (2004).
Rigsby, C. M., Herms, D. A., Bonello, P. & Cipollini, D. Higher activities of defense-associated enzymes may contribute to greater resistance of Manchurian ash to emerald ash borer than a closely related and susceptible congener. J. Chem. Ecol. 42, 782–792 (2016).
Villari, C., Herms, D. A., Whitehill, J. G. A., Cipollini, D. & Bonello, P. Progress and gaps in understanding mechanisms of ash tree resistance to emerald ash borer, a model for wood-boring insects that kill angiosperms. New Phytol. 209, 63–79 (2016).
Erb, M. & Reymond, P. Molecular interactions between plants and insect herbivores. Annu. Rev. Plant Biol. 70, 527–557 (2019).
Huang, J., Zhu, C. & Li, X. SCFSNIPER4 controls the turnover of two redundant TRAF proteins in plant immunity. Plant J. 95, 504–515 (2018).
Hua, Z. & Vierstra, R. D. The cullin-RING ubiquitin-protein ligases. Annu. Rev. Plant Biol. 62, 299–334 (2011).
Erb, M., Meldau, S. & Howe, G. A. Role of phytohormones in insect-specific plant reactions. Trends Plant Sci. 17, 250–259 (2012).
Berens, M. L., Berry, H. M., Mine, A., Argueso, C. T. & Tsuda, K. Evolution of hormone signaling networks in plant defense. Annu. Rev. Phytopathol. 55, 401–425 (2017).
Lin, S.-H. et al. Mutation of the Arabidopsis NRT1.5 nitrate transporter causes defective root-to-shoot nitrate transport. Plant Cell 20, 2514–2528 (2008).
Huysmans, M., Lema, A. S., Coll, N. S. & Nowack, M. K. Dying two deaths – programmed cell death regulation in development and disease. Curr. Opin. Plant Biol. 35, 37–44 (2017).
Bellin, D., Asai, S., Delledonne, M. & Yoshioka, H. Nitric oxide as a mediator for defense responses. Mol. Plant Microbe Interact. 26, 271–277 (2013).
Zebelo, S. A. & Maffei, M. E. Role of early signalling events in plant–insect interactions. J. Exp. Bot. 66, 435–448 (2015).
Seifi, H. S. & Shelp, B. J. Spermine differentially refines plant defense responses against biotic and abiotic stresses. Front. Plant Sci. 10, 117 (2019).
Whitehill, J. G. A., Rigsby, C., Cipollini, D., Herms, D. A. & Bonello, P. Decreased emergence of emerald ash borer from ash treated with methyl jasmonate is associated with induction of general defense traits and the toxic phenolic compound verbascoside. Oecologia 176, 1047–1059 (2014).
Nelson, R., Wiesner-Hanks, T., Wisser, R. & Balint-Kurti, P. Navigating complexity to breed disease-resistant crops. Nat. Rev. Genet. 19, 21–33 (2018).
Radville, L., Chaves, A. & Preisser, E. L. Variation in plant defense against invasive herbivores: evidence for a hypersensitive response in eastern hemlocks (Tsuga canadensis). J. Chem. Ecol. 37, 592–597 (2011).
Hilker, M. & Fatouros, N. E. Resisting the onset of herbivore attack: plants perceive and respond to insect eggs. Curr. Opin. Plant Biol. 32, 9–16 (2016).
Kim, C. Y., Bove, J. & Assmann, S. M. Overexpression of wound-responsive RNA-binding proteins induces leaf senescence and hypersensitive-like cell death. New Phytol. 180, 57–70 (2008).
Bollhöner, B. et al. The function of two type II metacaspases in woody tissues of Populus trees. New Phytol. 217, 1551–1565 (2018).
Altmann, S. et al. Transcriptomic basis for reinforcement of elm antiherbivore defence mediated by insect egg deposition. Mol. Ecol. 27, 4901–4915 (2018).
Rebek, E. J., Herms, D. A. & Smitley, D. R. Interspecific variation in resistance to emerald ash borer (Coleoptera: Buprestidae) among North American and Asian ash (Fraxinus spp.). Environ. Entomol. 37, 242–246 (2008).
Wei, Z. & Green, P. S. Fraxinus. Flora China 15, 273–279 (1996).
Davidson, C. G. ‘Northern Treasure’ and ‘Northern Gem’ hybrid ash. HortScience 34, 151–152 (1999).
Koch, J. L. et al. Strategies for selecting and breeding EAB-resistant ash. In Proc. 22nd US Department of Agriculture Interagency Research Symposium on Invasive Species (eds McManus, K. A. & Gottschalk, K. W.) 33–35 (US Department of Agriculture, Forest Service, Northern Research Station, 2011).
Duan, J. J., Larson, K., Watt, T., Gould, J. & Lelito, J. P. Effects of host plant and larval density on intraspecific competition in larvae of the emerald ash borer (Coleoptera: Buprestidae). Environ. Entomol. 42, 1193–1200 (2013).
Cappaert, D., McCullough, D. G., Poland, T. M. & Siegert, N. W. Emerald ash borer in North America: a research and regulatory challenge. Am. Entomol. 51, 152–165 (2005).
Chamorro, M. L., Volkovitsh, M. G., Poland, T. M., Haack, R. A. & Lingafelter, S. W. Preimaginal stages of the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae): an invasive pest on ash trees (Fraxinus). PLoS ONE 7, e33185 (2012).
Pellicer, J., Kelly, L. J., Leitch, I. J., Zomlefer, W. B. & Fay, M. F. A universe of dwarfs and giants: genome size and chromosome evolution in the monocot family Melanthiaceae. New Phytol. 201, 1484–1497 (2014).
Loureiro, J., Rodriguez, E., Dolezel, J. & Santos, C. Two new nuclear isolation buffers for plant DNA flow cytometry: a test with 37 species. Ann. Bot. 100, 875–888 (2007).
Doležel, J., Binarová, P. & Lucretti, S. Analysis of nuclear DNA content in plant cells by flow cytometry. Biol. Plant. 31, 113–120 (1989).
Bennett Michael, D. & Smith, J. B. Nuclear DNA amounts in angiosperms. Philos. Trans. R. Soc. Lond. B 334, 309–345 (1991).
Whittemore, A. T. & Xia, Z.-L. Genome size variation in elms (Ulmus spp.) and related genera. HortScience 52, 547–553 (2017).
Doležel, J. et al. Plant genome size estimation by flow cytometry: inter-laboratory comparison. Ann. Bot. 82, 17–26 (1998).
Greilhuber, J. & Obermayer, R. Genome size and maturity group in Glycine max (soybean). Heredity 78, 547–551 (1997).
Doyle, J. J. & Doyle, J. L. A rapid DNA isolation procedure for small quantities of fresh leaf tissue. Phytochem. Bull. 19, 11–15 (1987).
Martin, M. Cutadapt removes adapter sequences from high-throughput sequencing reads. EMBnet J. 17, 10–12 (2011).
Joshi, N. A. & Fass, J. N. Sickle: a sliding-window, adaptive, quality-based trimming tool for fastq files (2011); https://github.com/najoshi/sickle
Schmieder, R. & Edwards, R. Quality control and preprocessing of metagenomic datasets. Bioinformatics 27, 863–864 (2011).
Boetzer, M., Henkel, C. V., Jansen, H. J., Butler, D. & Pirovano, W. Scaffolding pre-assembled contigs using SSPACE. Bioinformatics 27, 578–579 (2011).
Luo, R. et al. SOAPdenovo2: an empirically improved memory-efficient short-read de novo assembler. Gigascience 1, 18 (2012).
Camacho, C. et al. BLAST+: architecture and applications. BMC Bioinform. 10, 421 (2009).
Simão, F. A., Waterhouse, R. M., Ioannidis, P., Kriventseva, E. V. & Zdobnov, E. M. BUSCO: assessing genome assembly and annotation completeness with single-copy orthologs. Bioinformatics 31, 3210–3212 (2015).
Keilwagen, J. et al. Using intron position conservation for homology-based gene prediction. Nucleic Acids Res. 44, e89 (2016).
Trapnell, C. et al. Transcript assembly and quantification by RNA-Seq reveals unannotated transcripts and isoform switching during cell differentiation. Nat. Biotechnol. 28, 511–515 (2010).
Quinlan, A. R. & Hall, I. M. BEDTools: a flexible suite of utilities for comparing genomic features. Bioinformatics 26, 841–842 (2010).
Altenhoff, A. M., Gil, M., Gonnet, G. H. & Dessimoz, C. Inferring hierarchical orthologous groups from orthologous gene pairs. PLoS ONE 8, e53786 (2013).
Altenhoff, A. M. et al. The OMA orthology database in 2015: function predictions, better plant support, synteny view and other improvements. Nucleic Acids Res. 43, D240–D249 (2015).
Goodstein, D. M. et al. Phytozome: a comparative platform for green plant genomics. Nucleic Acids Res. 40, D1178–D1186 (2012).
Wallander, E. Systematics of Fraxinus (Oleaceae) and evolution of dioecy. Plant Syst. Evol. 273, 25–49 (2008).
Hinsinger, D. D. et al. The phylogeny and biogeographic history of ashes (Fraxinus, Oleaceae) highlight the roles of migration and vicariance in the diversification of temperate trees. PLoS ONE 8, e80431 (2013).
Edgar, R. C. MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Res. 32, 1792–1797 (2004).
Sela, I., Ashkenazy, H., Katoh, K. & Pupko, T. GUIDANCE2: accurate detection of unreliable alignment regions accounting for the uncertainty of multiple parameters. Nucleic Acids Res. 43, W7–W14 (2015).
Rice, P., Longden, I. & Bleasby, A. EMBOSS: the European Molecular Biology Open Software Suite. Trends Genet. 16, 276–277 (2000).
Ronquist, F. et al. MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Syst. Biol. 61, 539–542 (2012).
Ané, C., Larget, B., Baum, D. A., Smith, S. D. & Rokas, A. Bayesian estimation of concordance among gene trees. Mol. Biol. Evol. 24, 412–426 (2007).
Larget, B. R., Kotha, S. K., Dewey, C. N. & Ané, C. BUCKy: gene tree/species tree reconciliation with Bayesian concordance analysis. Bioinformatics 26, 2910–2911 (2010).
Castoe, T. A. et al. Evidence for an ancient adaptive episode of convergent molecular evolution. Proc. Natl Acad. Sci. USA 106, 8986–8991 (2009).
Yang, Z. PAML 4: phylogenetic analysis by maximum likelihood. Mol. Biol. Evol. 24, 1586–1591 (2007).
Langmead, B. & Salzberg, S. L. Fast gapped-read alignment with Bowtie 2. Nat. Methods 9, 357–359 (2012).
Li, H. et al. The sequence alignment/map format and SAMtools. Bioinformatics 25, 2078–2079 (2009).
McKenna, A. et al. The genome analysis toolkit: a MapReduce framework for analyzing next-generation DNA sequencing data. Genome Res. 20, 1297–1303 (2010).
Cingolani, P. et al. A program for annotating and predicting the effects of single nucleotide polymorphisms, SnpEff: SNPs in the genome of Drosophila melanogaster strain w1118; iso-2; iso-3. Fly 6, 80–92 (2012).
Martin, M. et al. WhatsHap: fast and accurate read-based phasing. Preprint at bioRxiv https://doi.org/10.1101/085050 (2016).
Milne, I. et al. Using Tablet for visual exploration of second-generation sequencing data. Brief. Bioinform. 14, 193–202 (2013).
Katoh, K. & Standley, D. M. MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Mol. Biol. Evol. 30, 772–780 (2013).
Pond, S. L. K., Frost, S. D. W. & Muse, S. V. HyPhy: hypothesis testing using phylogenies. Bioinformatics 21, 676–679 (2005).
Kosakovsky Pond, S. L., Posada, D., Gravenor, M. B., Woelk, C. H. & Frost, S. D. W. Automated phylogenetic detection of recombination using a genetic algorithm. Mol. Biol. Evol. 23, 1891–1901 (2006).
Benson, D. A. et al. GenBank. Nucleic Acids Res. 41, D36–D42 (2013).
Liu, Z. et al. Evolutionary interplay between sister cytochrome P450 genes shapes plasticity in plant metabolism. Nat. Commun. 7, 13026 (2016).
Altenhoff, A. M. et al. The OMA orthology database in 2018: retrieving evolutionary relationships among all domains of life through richer web and programmatic interfaces. Nucleic Acids Res. 46, D477–D485 (2018).
Crooks, G. E., Hon, G., Chandonia, J.-M. & Brenner, S. E. WebLogo: a sequence logo generator. Genome Res. 14, 1188–1190 (2004).
Alexa, A., Rahnenführer, J. & Lengauer, T. Improved scoring of functional groups from gene expression data by decorrelating GO graph structure. Bioinformatics 22, 1600–1607 (2006).
Alexa, A. & Rahnenfuhrer, J. topGO: enrichment analysis for gene ontology R Package v.2.32.0 (2016).
R Core Team et al. R: a language and environment for statistical computing (R Foundation for Statistical Computing, 2013).
Petersen, T. N., Brunak, S., von Heijne, G. & Nielsen, H. SignalP 4.0: discriminating signal peptides from transmembrane regions. Nat. Methods 8, 785–786 (2011).
Käll, L., Krogh, A. & Sonnhammer, E. L. L. Advantages of combined transmembrane topology and signal peptide prediction—the Phobius web server. Nucleic Acids Res. 35, W429–W432 (2007).
Källberg, M. et al. Template-based protein structure modeling using the RaptorX web server. Nat. Protoc. 7, 1511–1522 (2012).
Waterhouse, A. et al. SWISS-MODEL: homology modelling of protein structures and complexes. Nucleic Acids Res. 46, W296–W303 (2018).
Kelley, L. A., Mezulis, S., Yates, C. M., Wass, M. N. & Sternberg, M. J. E. The Phyre2 web portal for protein modeling, prediction and analysis. Nat. Protoc. 10, 845–858 (2015).
Trott, O. & Olson, A. J. AutoDock Vina: improving the speed and accuracy of docking with a new scoring function, efficient optimization, and multithreading. J. Comput. Chem. 31, 455–461 (2010).
Dallakyan, S. & Olson, A. J. Small-molecule library screening by docking with PyRx. Methods Mol. Biol. 1263, 243–250 (2015).
This research used Queen Mary’s Apocrita HPC facility, supported by QMUL Research-IT (https://doi.org/10.5281/zenodo.438045). We thank J. Carlson for providing F. pennsylvanica DNA; T. Baxter, S. Brockington, P. Brownless, D. Crowley, S. Honey, R. Irvine, R. Jinks, P. Jones, T. Kirkham, H. McAllister, I. Parkinson and S. Redstone for help with obtaining Fraxinus materials from UK collections; T. Poland for providing EAB eggs; M. Miller for propagating trees for the bioassays; R. Matko for preparation of voucher specimens; J. Pellicer for advice on flow cytometry; P. Howard and M. Struebig for advice on DNA extractions; J. Keilwagen for help with GeMoMa; K. Davies and J. Parker for help with convergence analysis software; the Evolution Labchat group and Rossiter Lab at QMUL for discussions; and R. Rose and J. Sayers for advice on protein-modelling analyses. This project was funded by the Living with Environmental Change Tree Health and Plant Biosecurity Initiative – Phase 2 (grant no. BB/L012162/1), funded jointly by BBSRC, Defra, ESRC, Forestry Commission, NERC and the Scottish Government. R.J.A.B. acknowledges additional support from the DEFRA Future Proofing Plant Health scheme. R.J.A.B. and L.J.K. acknowledge additional support from the Erica Waltraud Albrecht Endowment Fund. W.J.P. was funded by the Walsh Scholarship Programme of the Department of Agriculture, Food and the Marine, Ireland. E.D.C. was supported by the Marie Skłodowska-Curie Individual Fellowship ‘FraxiFam’ (grant agreement no. 660003).
The authors declare no competing interests.
Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
a, Predicted protein structure for OG36502, modelled using the protein sequence for Fraxinus platypoda. The serine/asparagine variant at the site where convergence was detected is highlighted; the serine is a putative phosphorylation site. b, Predicted protein structure for OG40061, modelled using the protein sequence for F. mandshurica. The asparagine/serine variant at the site where convergence was detected is highlighted; the serine is a putative phosphorylation site. The putative substrate, NADP, is shown docked within the predicted active site. c, Predicted protein structure for OG38407, modelled using the protein sequence for F. mandshurica. The aspartic acid/asparagine variant at the site where convergence was detected is highlighted; the site falls within a leucine rich repeat region (LRR; shaded blue) which is predicted to span from position 111–237 within the protein sequence (detected using the GenomeNet MOTIF tool (www.genome.jp/tools/motif/), searching against the NCBI-CDD and Pfam databases with default parameters; the LRR region was identified as positions 111–237 with an e-value of 1e-05). d, Predicted protein structure for OG21033, modelled using the protein sequence for F. platypoda. The lysine/glutamine at the site where convergence was detected is highlighted. The putative substrate, β-D-Glcp-(1 → 3)-β-D-GlcpA-(1 → 4)-β-D-Glcp, is shown docked within the predicted active site.
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Kelly, L.J., Plumb, W.J., Carey, D.W. et al. Convergent molecular evolution among ash species resistant to the emerald ash borer. Nat Ecol Evol 4, 1116–1128 (2020). https://doi.org/10.1038/s41559-020-1209-3
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