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      AvrPm2 encodes an RNase‐like avirulence effector which is conserved in the two different specialized forms of wheat and rye powdery mildew fungus

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          Summary

          • There is a large diversity of genetically defined resistance genes in bread wheat against the powdery mildew pathogen Blumeria graminis ( B. g.) f. sp.  tritici. Many confer race‐specific resistance to this pathogen, but until now only the mildew avirulence gene AvrPm3 a2/f2 that is recognized by Pm3a/f was known molecularly.

          • We performed map‐based cloning and genome‐wide association studies to isolate a candidate for the mildew avirulence gene AvrPm2. We then used transient expression assays in Nicotiana benthamiana to demonstrate specific and strong recognition of AvrPm2 by Pm2.

          • The virulent AvrPm2 allele arose from a conserved 12 kb deletion, while there is no protein sequence diversity in the gene pool of avirulent B. g. tritici isolates. We found one polymorphic AvrPm2 allele in B. g. triticale and one orthologue in B. g. secalis and both are recognized by Pm2. AvrPm2 belongs to a small gene family encoding structurally conserved RNase‐like effectors, including Avr a13 from B. g. hordei, the cognate Avr of the barley resistance gene Mla13.

          • These results demonstrate the conservation of functional avirulence genes in two cereal powdery mildews specialized on different hosts, thus providing a possible explanation for successful introgression of resistance genes from rye or other grass relatives to wheat.

          Abstract

          See also the Commentary on this article by Spanu, 213: 969–971 .

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          Most cited references39

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          TranslatorX: multiple alignment of nucleotide sequences guided by amino acid translations

          We present TranslatorX, a web server designed to align protein-coding nucleotide sequences based on their corresponding amino acid translations. Many comparisons between biological sequences (nucleic acids and proteins) involve the construction of multiple alignments. Alignments represent a statement regarding the homology between individual nucleotides or amino acids within homologous genes. As protein-coding DNA sequences evolve as triplets of nucleotides (codons) and it is known that sequence similarity degrades more rapidly at the DNA than at the amino acid level, alignments are generally more accurate when based on amino acids than on their corresponding nucleotides. TranslatorX novelties include: (i) use of all documented genetic codes and the possibility of assigning different genetic codes for each sequence; (ii) a battery of different multiple alignment programs; (iii) translation of ambiguous codons when possible; (iv) an innovative criterion to clean nucleotide alignments with GBlocks based on protein information; and (v) a rich output, including Jalview-powered graphical visualization of the alignments, codon-based alignments coloured according to the corresponding amino acids, measures of compositional bias and first, second and third codon position specific alignments. The TranslatorX server is freely available at http://translatorx.co.uk.
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            Codon-substitution models for heterogeneous selection pressure at amino acid sites.

            Comparison of relative fixation rates of synonymous (silent) and nonsynonymous (amino acid-altering) mutations provides a means for understanding the mechanisms of molecular sequence evolution. The nonsynonymous/synonymous rate ratio (omega = d(N)d(S)) is an important indicator of selective pressure at the protein level, with omega = 1 meaning neutral mutations, omega 1 diversifying positive selection. Amino acid sites in a protein are expected to be under different selective pressures and have different underlying omega ratios. We develop models that account for heterogeneous omega ratios among amino acid sites and apply them to phylogenetic analyses of protein-coding DNA sequences. These models are useful for testing for adaptive molecular evolution and identifying amino acid sites under diversifying selection. Ten data sets of genes from nuclear, mitochondrial, and viral genomes are analyzed to estimate the distributions of omega among sites. In all data sets analyzed, the selective pressure indicated by the omega ratio is found to be highly heterogeneous among sites. Previously unsuspected Darwinian selection is detected in several genes in which the average omega ratio across sites is 1. Genes undergoing positive selection include the beta-globin gene from vertebrates, mitochondrial protein-coding genes from hominoids, the hemagglutinin (HA) gene from human influenza virus A, and HIV-1 env, vif, and pol genes. Tests for the presence of positively selected sites and their subsequent identification appear quite robust to the specific distributional form assumed for omega and can be achieved using any of several models we implement. However, we encountered difficulties in estimating the precise distribution of omega among sites from real data sets.
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              Genome expansion and gene loss in powdery mildew fungi reveal tradeoffs in extreme parasitism.

              Powdery mildews are phytopathogens whose growth and reproduction are entirely dependent on living plant cells. The molecular basis of this life-style, obligate biotrophy, remains unknown. We present the genome analysis of barley powdery mildew, Blumeria graminis f.sp. hordei (Blumeria), as well as a comparison with the analysis of two powdery mildews pathogenic on dicotyledonous plants. These genomes display massive retrotransposon proliferation, genome-size expansion, and gene losses. The missing genes encode enzymes of primary and secondary metabolism, carbohydrate-active enzymes, and transporters, probably reflecting their redundancy in an exclusively biotrophic life-style. Among the 248 candidate effectors of pathogenesis identified in the Blumeria genome, very few (less than 10) define a core set conserved in all three mildews, suggesting that most effectors represent species-specific adaptations.
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                Author and article information

                Contributors
                dazhaoyu@china.com
                bkeller@botinst.uzh.ch
                Journal
                New Phytol
                New Phytol
                10.1111/(ISSN)1469-8137
                NPH
                The New Phytologist
                John Wiley and Sons Inc. (Hoboken )
                0028-646X
                1469-8137
                09 December 2016
                February 2017
                : 213
                : 3 ( doiID: 10.1111/nph.2017.213.issue-3 )
                : 1301-1314
                Affiliations
                [ 1 ] Department of Plant and Microbial BiologyUniversity of Zürich Zürich 8008Switzerland
                [ 2 ] Institute of Plant Protection and Soil ScienceHubei Academy of Agricultural Sciences Wuhan 430064China
                [ 3 ]Ministry of Agriculture Key Laboratory of Integrated Pest Management in Crops in Central China Wuhan 430064China
                [ 4 ] College of Life ScienceWuhan University Wuhan 430072China
                [ 5 ] Institute of Plant ScienceARO‐Volcani Center Bet Dagan 50250Israel
                Author notes
                [*] [* ] Authors for correspondence:

                Beat Keller

                Tel: +41(0)446348230

                Email: bkeller@ 123456botinst.uzh.ch

                Dazhao Yu

                Tel: +86 13871126229

                Email: dazhaoyu@ 123456china.com

                [†]

                These authors contributed equally to this work.

                Article
                NPH14372 2016-22987
                10.1111/nph.14372
                5347869
                27935041
                c166a88e-80b2-4d52-9b21-133440de30a6
                © 2016 The Authors. New Phytologist © 2016 New Phytologist Trust

                This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

                History
                : 10 October 2016
                : 02 November 2016
                Page count
                Figures: 7, Tables: 0, Pages: 14, Words: 10673
                Funding
                Funded by: Swiss National Science Foundation
                Award ID: grant 310030‐163260
                Funded by: National Basic Research Program of China
                Award ID: 2013CB127700
                Funded by: China Agriculture Research System
                Award ID: CARS0304B
                Categories
                Full Paper
                Research
                Full Papers
                Custom metadata
                2.0
                nph14372
                February 2017
                Converter:WILEY_ML3GV2_TO_NLMPMC version:5.0.8 mode:remove_FC converted:13.03.2017

                Plant science & Botany
                avirulence gene,blumeria graminis,pm2,powdery mildew,rnase‐like,wheat
                Plant science & Botany
                avirulence gene, blumeria graminis, pm2, powdery mildew, rnase‐like, wheat

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