Genome analysis and avirulence gene cloning using a high-density RADseq linkage map of the flax rust fungus, Melampsora lini.
Identifieur interne : 000053 ( Main/Exploration ); précédent : 000052; suivant : 000054Genome analysis and avirulence gene cloning using a high-density RADseq linkage map of the flax rust fungus, Melampsora lini.
Auteurs : Claire Anderson [Australie] ; Muhammad Adil Khan [Australie] ; Ann-Maree Catanzariti [Australie] ; Cameron A. Jack [Australie] ; Adnane Nemri [Australie, Allemagne] ; Gregory J. Lawrence [Australie] ; Narayana M. Upadhyaya [Australie] ; Adrienne R. Hardham [Australie] ; Jeffrey G. Ellis [Australie] ; Peter N. Dodds [Australie] ; David A. Jones [Australie]Source :
- BMC genomics [ 1471-2164 ] ; 2016.
Descripteurs français
- KwdFr :
- Basidiomycota (génétique), Basidiomycota (pathogénicité), Biologie informatique (méthodes), Cartographie chromosomique (MeSH), Fréquence d'allèle (MeSH), Génome fongique (MeSH), Génomique (méthodes), Locus génétiques (MeSH), Mutation (MeSH), Perte d'hétérozygotie (MeSH), Phénotype (MeSH), Polymorphisme de nucléotide simple (MeSH), Recombinaison génétique (MeSH), Séquence d'acides aminés (MeSH), Séquençage nucléotidique à haut débit (MeSH), Virulence (génétique).
- MESH :
- génétique : Basidiomycota, Virulence.
- méthodes : Biologie informatique, Génomique.
- pathogénicité : Basidiomycota.
- Cartographie chromosomique, Fréquence d'allèle, Génome fongique, Locus génétiques, Mutation, Perte d'hétérozygotie, Phénotype, Polymorphisme de nucléotide simple, Recombinaison génétique, Séquence d'acides aminés, Séquençage nucléotidique à haut débit.
English descriptors
- KwdEn :
- Amino Acid Sequence (MeSH), Basidiomycota (genetics), Basidiomycota (pathogenicity), Chromosome Mapping (MeSH), Computational Biology (methods), Gene Frequency (MeSH), Genetic Loci (MeSH), Genome, Fungal (MeSH), Genomics (methods), High-Throughput Nucleotide Sequencing (MeSH), Loss of Heterozygosity (MeSH), Mutation (MeSH), Phenotype (MeSH), Polymorphism, Single Nucleotide (MeSH), Recombination, Genetic (MeSH), Virulence (genetics).
- MESH :
- genetics : Basidiomycota, Virulence.
- methods : Computational Biology, Genomics.
- pathogenicity : Basidiomycota.
- Amino Acid Sequence, Chromosome Mapping, Gene Frequency, Genetic Loci, Genome, Fungal, High-Throughput Nucleotide Sequencing, Loss of Heterozygosity, Mutation, Phenotype, Polymorphism, Single Nucleotide, Recombination, Genetic.
Abstract
BACKGROUND
Rust fungi are an important group of plant pathogens that cause devastating losses in agricultural, silvicultural and natural ecosystems. Plants can be protected from rust disease by resistance genes encoding receptors that trigger a highly effective defence response upon recognition of specific pathogen avirulence proteins. Identifying avirulence genes is crucial for understanding how virulence evolves in the field.
RESULTS
To facilitate avirulence gene cloning in the flax rust fungus, Melampsora lini, we constructed a high-density genetic linkage map using single nucleotide polymorphisms detected in restriction site-associated DNA sequencing (RADseq) data. The map comprises 13,412 RADseq markers in 27 linkage groups that together span 5860 cM and contain 2756 recombination bins. The marker sequences were used to anchor 68.9 % of the M. lini genome assembly onto the genetic map. The map and anchored assembly were then used to: 1) show that M. lini has a high overall meiotic recombination rate, but recombination distribution is uneven and large coldspots exist; 2) show that substantial genome rearrangements have occurred in spontaneous loss-of-avirulence mutants; and 3) identify the AvrL2 and AvrM14 avirulence genes by map-based cloning. AvrM14 is a dual-specificity avirulence gene that encodes a predicted nudix hydrolase. AvrL2 is located in the region of the M. lini genome with the lowest recombination rate and encodes a small, highly-charged proline-rich protein.
CONCLUSIONS
The M. lini high-density linkage map has greatly advanced our understanding of virulence mechanisms in this pathogen by providing novel insights into genome variability and enabling identification of two new avirulence genes.
DOI: 10.1186/s12864-016-3011-9
PubMed: 27550217
PubMed Central: PMC4994203
Affiliations:
Links toward previous steps (curation, corpus...)
Le document en format XML
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Jack</name><affiliation wicri:level="1"><nlm:affiliation>ANU Bioinformatics Consulting Unit, The John Curtin School of Medical Research, The Australian National University, 131 Garran Road, Acton, ACT, 2601, Australia.</nlm:affiliation><country xml:lang="fr">Australie</country><wicri:regionArea>ANU Bioinformatics Consulting Unit, The John Curtin School of Medical Research, The Australian National University, 131 Garran Road, Acton, ACT, 2601</wicri:regionArea><wicri:noRegion>2601</wicri:noRegion></affiliation></author><author><name sortKey="Nemri, Adnane" sort="Nemri, Adnane" uniqKey="Nemri A" first="Adnane" last="Nemri">Adnane Nemri</name><affiliation wicri:level="1"><nlm:affiliation>CSIRO Agriculture, GPO Box 1600, Canberra, ACT, 2601, Australia.</nlm:affiliation><country xml:lang="fr">Australie</country><wicri:regionArea>CSIRO Agriculture, GPO Box 1600, Canberra, ACT, 2601</wicri:regionArea><wicri:noRegion>2601</wicri:noRegion></affiliation><affiliation wicri:level="1"><nlm:affiliation>Current address: KWS SAAT SE, Grimsehlstraße 31, Einbeck, 37574, Germany.</nlm:affiliation><country xml:lang="fr">Allemagne</country><wicri:regionArea>Current address: KWS SAAT SE, Grimsehlstraße 31, Einbeck, 37574</wicri:regionArea><wicri:noRegion>37574</wicri:noRegion><wicri:noRegion>37574</wicri:noRegion><wicri:noRegion>37574</wicri:noRegion></affiliation></author><author><name sortKey="Lawrence, Gregory J" sort="Lawrence, Gregory J" uniqKey="Lawrence G" first="Gregory J" last="Lawrence">Gregory J. Lawrence</name><affiliation wicri:level="1"><nlm:affiliation>CSIRO Agriculture, GPO Box 1600, Canberra, ACT, 2601, Australia.</nlm:affiliation><country xml:lang="fr">Australie</country><wicri:regionArea>CSIRO Agriculture, GPO Box 1600, Canberra, ACT, 2601</wicri:regionArea><wicri:noRegion>2601</wicri:noRegion></affiliation></author><author><name sortKey="Upadhyaya, Narayana M" sort="Upadhyaya, Narayana M" uniqKey="Upadhyaya N" first="Narayana M" last="Upadhyaya">Narayana M. Upadhyaya</name><affiliation wicri:level="1"><nlm:affiliation>CSIRO Agriculture, GPO Box 1600, Canberra, ACT, 2601, Australia.</nlm:affiliation><country xml:lang="fr">Australie</country><wicri:regionArea>CSIRO Agriculture, GPO Box 1600, Canberra, ACT, 2601</wicri:regionArea><wicri:noRegion>2601</wicri:noRegion></affiliation></author><author><name sortKey="Hardham, Adrienne R" sort="Hardham, Adrienne R" uniqKey="Hardham A" first="Adrienne R" last="Hardham">Adrienne R. Hardham</name><affiliation wicri:level="1"><nlm:affiliation>Research School of Biology, The Australian National University, 134 Linnaeus Way, Acton, ACT, 2601, Australia.</nlm:affiliation><country xml:lang="fr">Australie</country><wicri:regionArea>Research School of Biology, The Australian National University, 134 Linnaeus Way, Acton, ACT, 2601</wicri:regionArea><wicri:noRegion>2601</wicri:noRegion></affiliation></author><author><name sortKey="Ellis, Jeffrey G" sort="Ellis, Jeffrey G" uniqKey="Ellis J" first="Jeffrey G" last="Ellis">Jeffrey G. Ellis</name><affiliation wicri:level="1"><nlm:affiliation>CSIRO Agriculture, GPO Box 1600, Canberra, ACT, 2601, Australia.</nlm:affiliation><country xml:lang="fr">Australie</country><wicri:regionArea>CSIRO Agriculture, GPO Box 1600, Canberra, ACT, 2601</wicri:regionArea><wicri:noRegion>2601</wicri:noRegion></affiliation></author><author><name sortKey="Dodds, Peter N" sort="Dodds, Peter N" uniqKey="Dodds P" first="Peter N" last="Dodds">Peter N. Dodds</name><affiliation wicri:level="1"><nlm:affiliation>CSIRO Agriculture, GPO Box 1600, Canberra, ACT, 2601, Australia.</nlm:affiliation><country xml:lang="fr">Australie</country><wicri:regionArea>CSIRO Agriculture, GPO Box 1600, Canberra, ACT, 2601</wicri:regionArea><wicri:noRegion>2601</wicri:noRegion></affiliation></author><author><name sortKey="Jones, David A" sort="Jones, David A" uniqKey="Jones D" first="David A" last="Jones">David A. Jones</name><affiliation wicri:level="1"><nlm:affiliation>Research School of Biology, The Australian National University, 134 Linnaeus Way, Acton, ACT, 2601, Australia. david.jones@anu.edu.au.</nlm:affiliation><country xml:lang="fr">Australie</country><wicri:regionArea>Research School of Biology, The Australian National University, 134 Linnaeus Way, Acton, ACT, 2601</wicri:regionArea><wicri:noRegion>2601</wicri:noRegion></affiliation></author></analytic><series><title level="j">BMC genomics</title><idno type="eISSN">1471-2164</idno><imprint><date when="2016" type="published">2016</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass><keywords scheme="KwdEn" xml:lang="en"><term>Amino Acid Sequence (MeSH)</term><term>Basidiomycota (genetics)</term><term>Basidiomycota (pathogenicity)</term><term>Chromosome Mapping (MeSH)</term><term>Computational Biology (methods)</term><term>Gene Frequency (MeSH)</term><term>Genetic Loci (MeSH)</term><term>Genome, Fungal (MeSH)</term><term>Genomics (methods)</term><term>High-Throughput Nucleotide Sequencing (MeSH)</term><term>Loss of Heterozygosity (MeSH)</term><term>Mutation (MeSH)</term><term>Phenotype (MeSH)</term><term>Polymorphism, Single Nucleotide (MeSH)</term><term>Recombination, Genetic (MeSH)</term><term>Virulence (genetics)</term></keywords><keywords scheme="KwdFr" xml:lang="fr"><term>Basidiomycota (génétique)</term><term>Basidiomycota (pathogénicité)</term><term>Biologie informatique (méthodes)</term><term>Cartographie chromosomique (MeSH)</term><term>Fréquence d'allèle (MeSH)</term><term>Génome fongique (MeSH)</term><term>Génomique (méthodes)</term><term>Locus génétiques (MeSH)</term><term>Mutation (MeSH)</term><term>Perte d'hétérozygotie (MeSH)</term><term>Phénotype (MeSH)</term><term>Polymorphisme de nucléotide simple (MeSH)</term><term>Recombinaison génétique (MeSH)</term><term>Séquence d'acides aminés (MeSH)</term><term>Séquençage nucléotidique à haut débit (MeSH)</term><term>Virulence (génétique)</term></keywords><keywords scheme="MESH" qualifier="genetics" xml:lang="en"><term>Basidiomycota</term><term>Virulence</term></keywords><keywords scheme="MESH" qualifier="génétique" xml:lang="fr"><term>Basidiomycota</term><term>Virulence</term></keywords><keywords scheme="MESH" qualifier="methods" xml:lang="en"><term>Computational Biology</term><term>Genomics</term></keywords><keywords scheme="MESH" qualifier="méthodes" xml:lang="fr"><term>Biologie informatique</term><term>Génomique</term></keywords><keywords scheme="MESH" qualifier="pathogenicity" xml:lang="en"><term>Basidiomycota</term></keywords><keywords scheme="MESH" qualifier="pathogénicité" xml:lang="fr"><term>Basidiomycota</term></keywords><keywords scheme="MESH" xml:lang="en"><term>Amino Acid Sequence</term><term>Chromosome Mapping</term><term>Gene Frequency</term><term>Genetic Loci</term><term>Genome, Fungal</term><term>High-Throughput Nucleotide Sequencing</term><term>Loss of Heterozygosity</term><term>Mutation</term><term>Phenotype</term><term>Polymorphism, Single Nucleotide</term><term>Recombination, Genetic</term></keywords><keywords scheme="MESH" xml:lang="fr"><term>Cartographie chromosomique</term><term>Fréquence d'allèle</term><term>Génome fongique</term><term>Locus génétiques</term><term>Mutation</term><term>Perte d'hétérozygotie</term><term>Phénotype</term><term>Polymorphisme de nucléotide simple</term><term>Recombinaison génétique</term><term>Séquence d'acides aminés</term><term>Séquençage nucléotidique à haut débit</term></keywords></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p><b>BACKGROUND</b></p><p>Rust fungi are an important group of plant pathogens that cause devastating losses in agricultural, silvicultural and natural ecosystems. Plants can be protected from rust disease by resistance genes encoding receptors that trigger a highly effective defence response upon recognition of specific pathogen avirulence proteins. Identifying avirulence genes is crucial for understanding how virulence evolves in the field.</p></div><div type="abstract" xml:lang="en"><p><b>RESULTS</b></p><p>To facilitate avirulence gene cloning in the flax rust fungus, Melampsora lini, we constructed a high-density genetic linkage map using single nucleotide polymorphisms detected in restriction site-associated DNA sequencing (RADseq) data. The map comprises 13,412 RADseq markers in 27 linkage groups that together span 5860 cM and contain 2756 recombination bins. The marker sequences were used to anchor 68.9 % of the M. lini genome assembly onto the genetic map. The map and anchored assembly were then used to: 1) show that M. lini has a high overall meiotic recombination rate, but recombination distribution is uneven and large coldspots exist; 2) show that substantial genome rearrangements have occurred in spontaneous loss-of-avirulence mutants; and 3) identify the AvrL2 and AvrM14 avirulence genes by map-based cloning. AvrM14 is a dual-specificity avirulence gene that encodes a predicted nudix hydrolase. AvrL2 is located in the region of the M. lini genome with the lowest recombination rate and encodes a small, highly-charged proline-rich protein.</p></div><div type="abstract" xml:lang="en"><p><b>CONCLUSIONS</b></p><p>The M. lini high-density linkage map has greatly advanced our understanding of virulence mechanisms in this pathogen by providing novel insights into genome variability and enabling identification of two new avirulence genes.</p></div></front></TEI><pubmed><MedlineCitation Status="MEDLINE" Owner="NLM"><PMID Version="1">27550217</PMID><DateCompleted><Year>2017</Year><Month>09</Month><Day>11</Day></DateCompleted><DateRevised><Year>2018</Year><Month>11</Month><Day>13</Day></DateRevised><Article PubModel="Electronic"><Journal><ISSN IssnType="Electronic">1471-2164</ISSN><JournalIssue CitedMedium="Internet"><Volume>17</Volume><PubDate><Year>2016</Year><Month>08</Month><Day>22</Day></PubDate></JournalIssue><Title>BMC genomics</Title><ISOAbbreviation>BMC Genomics</ISOAbbreviation></Journal><ArticleTitle>Genome analysis and avirulence gene cloning using a high-density RADseq linkage map of the flax rust fungus, Melampsora lini.</ArticleTitle><Pagination><MedlinePgn>667</MedlinePgn></Pagination><ELocationID EIdType="doi" ValidYN="Y">10.1186/s12864-016-3011-9</ELocationID><Abstract><AbstractText Label="BACKGROUND">Rust fungi are an important group of plant pathogens that cause devastating losses in agricultural, silvicultural and natural ecosystems. Plants can be protected from rust disease by resistance genes encoding receptors that trigger a highly effective defence response upon recognition of specific pathogen avirulence proteins. Identifying avirulence genes is crucial for understanding how virulence evolves in the field.</AbstractText><AbstractText Label="RESULTS">To facilitate avirulence gene cloning in the flax rust fungus, Melampsora lini, we constructed a high-density genetic linkage map using single nucleotide polymorphisms detected in restriction site-associated DNA sequencing (RADseq) data. The map comprises 13,412 RADseq markers in 27 linkage groups that together span 5860 cM and contain 2756 recombination bins. The marker sequences were used to anchor 68.9 % of the M. lini genome assembly onto the genetic map. The map and anchored assembly were then used to: 1) show that M. lini has a high overall meiotic recombination rate, but recombination distribution is uneven and large coldspots exist; 2) show that substantial genome rearrangements have occurred in spontaneous loss-of-avirulence mutants; and 3) identify the AvrL2 and AvrM14 avirulence genes by map-based cloning. AvrM14 is a dual-specificity avirulence gene that encodes a predicted nudix hydrolase. AvrL2 is located in the region of the M. lini genome with the lowest recombination rate and encodes a small, highly-charged proline-rich protein.</AbstractText><AbstractText Label="CONCLUSIONS">The M. lini high-density linkage map has greatly advanced our understanding of virulence mechanisms in this pathogen by providing novel insights into genome variability and enabling identification of two new avirulence genes.</AbstractText></Abstract><AuthorList CompleteYN="Y"><Author ValidYN="Y"><LastName>Anderson</LastName><ForeName>Claire</ForeName><Initials>C</Initials><AffiliationInfo><Affiliation>Research School of Biology, The Australian National University, 134 Linnaeus Way, Acton, ACT, 2601, Australia.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Khan</LastName><ForeName>Muhammad Adil</ForeName><Initials>MA</Initials><AffiliationInfo><Affiliation>Research School of Biology, The Australian National University, 134 Linnaeus Way, Acton, ACT, 2601, Australia.</Affiliation></AffiliationInfo><AffiliationInfo><Affiliation>Current address: ARC Centre of Excellence in Plant Energy Biology, The University of Western Australia, 35 Stirling Highway, Crawley, WA, 6009, Australia.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Catanzariti</LastName><ForeName>Ann-Maree</ForeName><Initials>AM</Initials><AffiliationInfo><Affiliation>Research School of Biology, The Australian National University, 134 Linnaeus Way, Acton, ACT, 2601, Australia.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Jack</LastName><ForeName>Cameron A</ForeName><Initials>CA</Initials><AffiliationInfo><Affiliation>ANU Bioinformatics Consulting Unit, The John Curtin School of Medical Research, The Australian National University, 131 Garran Road, Acton, ACT, 2601, Australia.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Nemri</LastName><ForeName>Adnane</ForeName><Initials>A</Initials><AffiliationInfo><Affiliation>CSIRO Agriculture, GPO Box 1600, Canberra, ACT, 2601, Australia.</Affiliation></AffiliationInfo><AffiliationInfo><Affiliation>Current address: KWS SAAT SE, Grimsehlstraße 31, Einbeck, 37574, Germany.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Lawrence</LastName><ForeName>Gregory J</ForeName><Initials>GJ</Initials><AffiliationInfo><Affiliation>CSIRO Agriculture, GPO Box 1600, Canberra, ACT, 2601, Australia.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Upadhyaya</LastName><ForeName>Narayana M</ForeName><Initials>NM</Initials><AffiliationInfo><Affiliation>CSIRO Agriculture, GPO Box 1600, Canberra, ACT, 2601, Australia.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Hardham</LastName><ForeName>Adrienne R</ForeName><Initials>AR</Initials><AffiliationInfo><Affiliation>Research School of Biology, The Australian National University, 134 Linnaeus Way, Acton, ACT, 2601, Australia.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Ellis</LastName><ForeName>Jeffrey G</ForeName><Initials>JG</Initials><AffiliationInfo><Affiliation>CSIRO Agriculture, GPO Box 1600, Canberra, ACT, 2601, Australia.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Dodds</LastName><ForeName>Peter N</ForeName><Initials>PN</Initials><AffiliationInfo><Affiliation>CSIRO Agriculture, GPO Box 1600, Canberra, ACT, 2601, Australia.</Affiliation></AffiliationInfo></Author><Author ValidYN="Y"><LastName>Jones</LastName><ForeName>David A</ForeName><Initials>DA</Initials><AffiliationInfo><Affiliation>Research School of Biology, The Australian National University, 134 Linnaeus Way, Acton, ACT, 2601, Australia. david.jones@anu.edu.au.</Affiliation></AffiliationInfo></Author></AuthorList><Language>eng</Language><PublicationTypeList><PublicationType UI="D016428">Journal Article</PublicationType><PublicationType UI="D013485">Research Support, Non-U.S. Gov't</PublicationType></PublicationTypeList><ArticleDate DateType="Electronic"><Year>2016</Year><Month>08</Month><Day>22</Day></ArticleDate></Article><MedlineJournalInfo><Country>England</Country><MedlineTA>BMC Genomics</MedlineTA><NlmUniqueID>100965258</NlmUniqueID><ISSNLinking>1471-2164</ISSNLinking></MedlineJournalInfo><CitationSubset>IM</CitationSubset><MeshHeadingList><MeshHeading><DescriptorName UI="D000595" MajorTopicYN="N">Amino Acid Sequence</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D001487" MajorTopicYN="N">Basidiomycota</DescriptorName><QualifierName UI="Q000235" MajorTopicYN="Y">genetics</QualifierName><QualifierName UI="Q000472" MajorTopicYN="N">pathogenicity</QualifierName></MeshHeading><MeshHeading><DescriptorName UI="D002874" MajorTopicYN="Y">Chromosome Mapping</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D019295" MajorTopicYN="N">Computational Biology</DescriptorName><QualifierName UI="Q000379" MajorTopicYN="N">methods</QualifierName></MeshHeading><MeshHeading><DescriptorName UI="D005787" MajorTopicYN="N">Gene Frequency</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D056426" MajorTopicYN="N">Genetic Loci</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D016681" MajorTopicYN="Y">Genome, Fungal</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D023281" MajorTopicYN="Y">Genomics</DescriptorName><QualifierName UI="Q000379" MajorTopicYN="N">methods</QualifierName></MeshHeading><MeshHeading><DescriptorName UI="D059014" MajorTopicYN="N">High-Throughput Nucleotide Sequencing</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D019656" MajorTopicYN="N">Loss of Heterozygosity</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D009154" MajorTopicYN="N">Mutation</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D010641" MajorTopicYN="N">Phenotype</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D020641" MajorTopicYN="N">Polymorphism, Single Nucleotide</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D011995" MajorTopicYN="N">Recombination, Genetic</DescriptorName></MeshHeading><MeshHeading><DescriptorName UI="D014774" MajorTopicYN="N">Virulence</DescriptorName><QualifierName UI="Q000235" MajorTopicYN="Y">genetics</QualifierName></MeshHeading></MeshHeadingList><KeywordList Owner="NOTNLM"><Keyword MajorTopicYN="Y">Avirulence gene</Keyword><Keyword MajorTopicYN="Y">Genetic linkage map</Keyword><Keyword MajorTopicYN="Y">Loss of heterozygosity (LOH)</Keyword><Keyword MajorTopicYN="Y">Map-based cloning</Keyword><Keyword MajorTopicYN="Y">Melampsora lini</Keyword><Keyword MajorTopicYN="Y">Recombination</Keyword><Keyword MajorTopicYN="Y">Restriction-site associated DNA sequencing (RADseq)</Keyword><Keyword MajorTopicYN="Y">Rust fungus</Keyword><Keyword MajorTopicYN="Y">Scaffold 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