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Ty-6, a major begomovirus resistance gene on chromosome 10, is effective against Tomato yellow leaf curl virus and Tomato mottle virus
Key message Ty - 6 is a major resistance gene on chromosome 10 of tomato that provides resistance against monopartite and bipartite begomoviruses and complements resistance conferred by the known Ty - 3 and ty - 5 genes. Resistance to monopartite and bipartite begomoviruses is an important breeding...
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Published in: | Theoretical and applied genetics 2019-05, Vol.132 (5), p.1543-1554 |
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container_title | Theoretical and applied genetics |
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creator | Gill, Upinder Scott, John W. Shekasteband, Reza Ogundiwin, Eben Schuit, Cees Francis, David M. Sim, Sung-Chur Smith, Hugh Hutton, Samuel F. |
description | Key message
Ty
-
6
is a major resistance gene on chromosome 10 of tomato that provides resistance against monopartite and bipartite begomoviruses and complements resistance conferred by the known
Ty
-
3
and
ty
-
5
genes.
Resistance to monopartite and bipartite begomoviruses is an important breeding objective for cultivated tomato. Several begomovirus resistance genes have been introgressed from related
Solanum
species and are available for breeding purposes. In the present study, we mapped an additional locus,
Ty
-
6
, to chromosome 10 of tomato.
Ty
-
6
is effective against both monopartite Tomato yellow leaf curl virus (TYLCV) and bipartite Tomato mottle virus (ToMoV). Gene action is incomplete dominance, with an intermediate resistance response when
Ty
-
6
is heterozygous. Analysis of populations segregating for
Ty
-
6
along with
Ty
-
3
or
ty
-
5
indicates that the highest level of resistance against TYLCV is attained when
Ty
-
6
is combined with an additional resistance allele. Our results also demonstrate that
ty
-
5
is ineffective against ToMoV. Although multiple SNPs linked to
Ty
-
6
were identified and can be used for breeding purposes, none of these were consistently polymorphic between
Ty
-
6
and
ty
-
6
breeding lines. Further research is underway to generate resequencing data for several
Ty
-
6
inbred lines for the discovery of additional sequence polymorphisms that can be used for fine mapping and characterizing the
Ty
-
6
locus. |
doi_str_mv | 10.1007/s00122-019-03298-0 |
format | article |
fullrecord | <record><control><sourceid>gale_pubme</sourceid><recordid>TN_cdi_pubmedcentral_primary_oai_pubmedcentral_nih_gov_6476845</recordid><sourceformat>XML</sourceformat><sourcesystem>PC</sourcesystem><galeid>A583293383</galeid><sourcerecordid>A583293383</sourcerecordid><originalsourceid>FETCH-LOGICAL-c641t-d83a4d3e7524afb81c28fdb46e87f868498067c375409c8fe2c93549ca9497a33</originalsourceid><addsrcrecordid>eNp9kt9qFDEUxoModl19AS8k4I1Cp-bvTOZGKKVqoSDoeh2ymTPTLDNJTWZW9wX63GadtnZFJJDAOb_zJV_4EHpJyQklpHqXCKGMFYTWBeGsVgV5hBZUcFYwJthjtCBEkEJWkh2hZyltCCFMEv4UHXFSSSU5XaCb1a4oj7HBg9mEiNfQhSFsXZwSjpBcGo23gDvwgIPH9irmdgoDYEqOsUsY2hbs6LaATWecTyNehcGMAe-g78MP3INpsZ1ij2dR45s7Ygjj2MNcf46etKZP8OL2XKJvH85XZ5-Ky88fL85OLwtbCjoWjeJGNByyJWHataKWqbZZixJU1apSiVqRsrK8koLUVrXAbM2lqK2pRV0Zzpfo_ax7Pa0HaCz4MZpeX0c3mLjTwTh92PHuSndhq0tRZXmZBd7cCsTwfYI06sElm70aD2FKmjHKqZSK79HXf6GbMEWf7WlGq5pWpSQPqM70oJ1vQ77X7kX1aZZhNedq_-6Tf1B5NTA4Gzy0LtcPBt4eDGRmhJ9jZ6aU9MXXL4csm1kbQ0oR2vv_oETvk6bnpOmcNP07aXlfolcPf_J-5C5aGeAzkHLLdxD_2P-P7C_0L90f</addsrcrecordid><sourcetype>Open Access Repository</sourcetype><iscdi>true</iscdi><recordtype>article</recordtype><pqid>2179176505</pqid></control><display><type>article</type><title>Ty-6, a major begomovirus resistance gene on chromosome 10, is effective against Tomato yellow leaf curl virus and Tomato mottle virus</title><source>Springer Nature</source><creator>Gill, Upinder ; Scott, John W. ; Shekasteband, Reza ; Ogundiwin, Eben ; Schuit, Cees ; Francis, David M. ; Sim, Sung-Chur ; Smith, Hugh ; Hutton, Samuel F.</creator><creatorcontrib>Gill, Upinder ; Scott, John W. ; Shekasteband, Reza ; Ogundiwin, Eben ; Schuit, Cees ; Francis, David M. ; Sim, Sung-Chur ; Smith, Hugh ; Hutton, Samuel F.</creatorcontrib><description>Key message
Ty
-
6
is a major resistance gene on chromosome 10 of tomato that provides resistance against monopartite and bipartite begomoviruses and complements resistance conferred by the known
Ty
-
3
and
ty
-
5
genes.
Resistance to monopartite and bipartite begomoviruses is an important breeding objective for cultivated tomato. Several begomovirus resistance genes have been introgressed from related
Solanum
species and are available for breeding purposes. In the present study, we mapped an additional locus,
Ty
-
6
, to chromosome 10 of tomato.
Ty
-
6
is effective against both monopartite Tomato yellow leaf curl virus (TYLCV) and bipartite Tomato mottle virus (ToMoV). Gene action is incomplete dominance, with an intermediate resistance response when
Ty
-
6
is heterozygous. Analysis of populations segregating for
Ty
-
6
along with
Ty
-
3
or
ty
-
5
indicates that the highest level of resistance against TYLCV is attained when
Ty
-
6
is combined with an additional resistance allele. Our results also demonstrate that
ty
-
5
is ineffective against ToMoV. Although multiple SNPs linked to
Ty
-
6
were identified and can be used for breeding purposes, none of these were consistently polymorphic between
Ty
-
6
and
ty
-
6
breeding lines. Further research is underway to generate resequencing data for several
Ty
-
6
inbred lines for the discovery of additional sequence polymorphisms that can be used for fine mapping and characterizing the
Ty
-
6
locus.</description><identifier>ISSN: 0040-5752</identifier><identifier>EISSN: 1432-2242</identifier><identifier>DOI: 10.1007/s00122-019-03298-0</identifier><identifier>PMID: 30758531</identifier><language>eng</language><publisher>Berlin/Heidelberg: Springer Berlin Heidelberg</publisher><subject>Agriculture ; Begomovirus ; Biochemistry ; Biomedical and Life Sciences ; Biotechnology ; Chromosome 10 ; Chromosome Mapping ; Chromosomes ; Chromosomes, Plant ; Disease resistance ; Disease Resistance - genetics ; DNA viruses ; Gene mapping ; Genes ; Genes, Plant ; Genetic aspects ; Inbreeding ; Leaf-curl ; Leaves ; Life Sciences ; Lycopersicon esculentum - genetics ; Lycopersicon esculentum - virology ; Original ; Original Article ; Plant Biochemistry ; Plant breeding ; Plant Breeding/Biotechnology ; Plant Diseases - genetics ; Plant Diseases - virology ; Plant Genetics and Genomics ; Plant viruses ; Quantitative Trait Loci ; Single-nucleotide polymorphism ; Solanaceae ; Tomatoes ; Yellow leaf</subject><ispartof>Theoretical and applied genetics, 2019-05, Vol.132 (5), p.1543-1554</ispartof><rights>The Author(s) 2019</rights><rights>COPYRIGHT 2019 Springer</rights><rights>Theoretical and Applied Genetics is a copyright of Springer, (2019). All Rights Reserved. © 2019. This work is published under http://creativecommons.org/licenses/by/4.0/ (the “License”). Notwithstanding the ProQuest Terms and Conditions, you may use this content in accordance with the terms of the License.</rights><lds50>peer_reviewed</lds50><oa>free_for_read</oa><woscitedreferencessubscribed>false</woscitedreferencessubscribed><citedby>FETCH-LOGICAL-c641t-d83a4d3e7524afb81c28fdb46e87f868498067c375409c8fe2c93549ca9497a33</citedby><cites>FETCH-LOGICAL-c641t-d83a4d3e7524afb81c28fdb46e87f868498067c375409c8fe2c93549ca9497a33</cites><orcidid>0000-0002-1635-4861</orcidid></display><links><openurl>$$Topenurl_article</openurl><openurlfulltext>$$Topenurlfull_article</openurlfulltext><thumbnail>$$Tsyndetics_thumb_exl</thumbnail><link.rule.ids>230,314,780,784,885,27923,27924</link.rule.ids><backlink>$$Uhttps://www.ncbi.nlm.nih.gov/pubmed/30758531$$D View this record in MEDLINE/PubMed$$Hfree_for_read</backlink></links><search><creatorcontrib>Gill, Upinder</creatorcontrib><creatorcontrib>Scott, John W.</creatorcontrib><creatorcontrib>Shekasteband, Reza</creatorcontrib><creatorcontrib>Ogundiwin, Eben</creatorcontrib><creatorcontrib>Schuit, Cees</creatorcontrib><creatorcontrib>Francis, David M.</creatorcontrib><creatorcontrib>Sim, Sung-Chur</creatorcontrib><creatorcontrib>Smith, Hugh</creatorcontrib><creatorcontrib>Hutton, Samuel F.</creatorcontrib><title>Ty-6, a major begomovirus resistance gene on chromosome 10, is effective against Tomato yellow leaf curl virus and Tomato mottle virus</title><title>Theoretical and applied genetics</title><addtitle>Theor Appl Genet</addtitle><addtitle>Theor Appl Genet</addtitle><description>Key message
Ty
-
6
is a major resistance gene on chromosome 10 of tomato that provides resistance against monopartite and bipartite begomoviruses and complements resistance conferred by the known
Ty
-
3
and
ty
-
5
genes.
Resistance to monopartite and bipartite begomoviruses is an important breeding objective for cultivated tomato. Several begomovirus resistance genes have been introgressed from related
Solanum
species and are available for breeding purposes. In the present study, we mapped an additional locus,
Ty
-
6
, to chromosome 10 of tomato.
Ty
-
6
is effective against both monopartite Tomato yellow leaf curl virus (TYLCV) and bipartite Tomato mottle virus (ToMoV). Gene action is incomplete dominance, with an intermediate resistance response when
Ty
-
6
is heterozygous. Analysis of populations segregating for
Ty
-
6
along with
Ty
-
3
or
ty
-
5
indicates that the highest level of resistance against TYLCV is attained when
Ty
-
6
is combined with an additional resistance allele. Our results also demonstrate that
ty
-
5
is ineffective against ToMoV. Although multiple SNPs linked to
Ty
-
6
were identified and can be used for breeding purposes, none of these were consistently polymorphic between
Ty
-
6
and
ty
-
6
breeding lines. Further research is underway to generate resequencing data for several
Ty
-
6
inbred lines for the discovery of additional sequence polymorphisms that can be used for fine mapping and characterizing the
Ty
-
6
locus.</description><subject>Agriculture</subject><subject>Begomovirus</subject><subject>Biochemistry</subject><subject>Biomedical and Life Sciences</subject><subject>Biotechnology</subject><subject>Chromosome 10</subject><subject>Chromosome Mapping</subject><subject>Chromosomes</subject><subject>Chromosomes, Plant</subject><subject>Disease resistance</subject><subject>Disease Resistance - genetics</subject><subject>DNA viruses</subject><subject>Gene mapping</subject><subject>Genes</subject><subject>Genes, Plant</subject><subject>Genetic aspects</subject><subject>Inbreeding</subject><subject>Leaf-curl</subject><subject>Leaves</subject><subject>Life Sciences</subject><subject>Lycopersicon esculentum - genetics</subject><subject>Lycopersicon esculentum - virology</subject><subject>Original</subject><subject>Original Article</subject><subject>Plant Biochemistry</subject><subject>Plant breeding</subject><subject>Plant Breeding/Biotechnology</subject><subject>Plant Diseases - genetics</subject><subject>Plant Diseases - virology</subject><subject>Plant Genetics and Genomics</subject><subject>Plant viruses</subject><subject>Quantitative Trait Loci</subject><subject>Single-nucleotide polymorphism</subject><subject>Solanaceae</subject><subject>Tomatoes</subject><subject>Yellow leaf</subject><issn>0040-5752</issn><issn>1432-2242</issn><fulltext>true</fulltext><rsrctype>article</rsrctype><creationdate>2019</creationdate><recordtype>article</recordtype><recordid>eNp9kt9qFDEUxoModl19AS8k4I1Cp-bvTOZGKKVqoSDoeh2ymTPTLDNJTWZW9wX63GadtnZFJJDAOb_zJV_4EHpJyQklpHqXCKGMFYTWBeGsVgV5hBZUcFYwJthjtCBEkEJWkh2hZyltCCFMEv4UHXFSSSU5XaCb1a4oj7HBg9mEiNfQhSFsXZwSjpBcGo23gDvwgIPH9irmdgoDYEqOsUsY2hbs6LaATWecTyNehcGMAe-g78MP3INpsZ1ij2dR45s7Ygjj2MNcf46etKZP8OL2XKJvH85XZ5-Ky88fL85OLwtbCjoWjeJGNByyJWHataKWqbZZixJU1apSiVqRsrK8koLUVrXAbM2lqK2pRV0Zzpfo_ax7Pa0HaCz4MZpeX0c3mLjTwTh92PHuSndhq0tRZXmZBd7cCsTwfYI06sElm70aD2FKmjHKqZSK79HXf6GbMEWf7WlGq5pWpSQPqM70oJ1vQ77X7kX1aZZhNedq_-6Tf1B5NTA4Gzy0LtcPBt4eDGRmhJ9jZ6aU9MXXL4csm1kbQ0oR2vv_oETvk6bnpOmcNP07aXlfolcPf_J-5C5aGeAzkHLLdxD_2P-P7C_0L90f</recordid><startdate>20190501</startdate><enddate>20190501</enddate><creator>Gill, Upinder</creator><creator>Scott, John W.</creator><creator>Shekasteband, Reza</creator><creator>Ogundiwin, Eben</creator><creator>Schuit, Cees</creator><creator>Francis, David M.</creator><creator>Sim, Sung-Chur</creator><creator>Smith, Hugh</creator><creator>Hutton, Samuel F.</creator><general>Springer Berlin Heidelberg</general><general>Springer</general><general>Springer Nature B.V</general><scope>C6C</scope><scope>CGR</scope><scope>CUY</scope><scope>CVF</scope><scope>ECM</scope><scope>EIF</scope><scope>NPM</scope><scope>AAYXX</scope><scope>CITATION</scope><scope>ISR</scope><scope>3V.</scope><scope>7SS</scope><scope>7TK</scope><scope>7X7</scope><scope>7XB</scope><scope>88A</scope><scope>88E</scope><scope>8AO</scope><scope>8FD</scope><scope>8FE</scope><scope>8FH</scope><scope>8FI</scope><scope>8FJ</scope><scope>8FK</scope><scope>ABUWG</scope><scope>AFKRA</scope><scope>AZQEC</scope><scope>BBNVY</scope><scope>BENPR</scope><scope>BHPHI</scope><scope>CCPQU</scope><scope>DWQXO</scope><scope>FR3</scope><scope>FYUFA</scope><scope>GHDGH</scope><scope>GNUQQ</scope><scope>HCIFZ</scope><scope>K9.</scope><scope>LK8</scope><scope>M0S</scope><scope>M1P</scope><scope>M7P</scope><scope>P64</scope><scope>PQEST</scope><scope>PQQKQ</scope><scope>PQUKI</scope><scope>RC3</scope><scope>7X8</scope><scope>5PM</scope><orcidid>https://orcid.org/0000-0002-1635-4861</orcidid></search><sort><creationdate>20190501</creationdate><title>Ty-6, a major begomovirus resistance gene on chromosome 10, is effective against Tomato yellow leaf curl virus and Tomato mottle virus</title><author>Gill, Upinder ; Scott, John W. ; Shekasteband, Reza ; Ogundiwin, Eben ; Schuit, Cees ; Francis, David M. ; Sim, Sung-Chur ; Smith, Hugh ; Hutton, Samuel F.</author></sort><facets><frbrtype>5</frbrtype><frbrgroupid>cdi_FETCH-LOGICAL-c641t-d83a4d3e7524afb81c28fdb46e87f868498067c375409c8fe2c93549ca9497a33</frbrgroupid><rsrctype>articles</rsrctype><prefilter>articles</prefilter><language>eng</language><creationdate>2019</creationdate><topic>Agriculture</topic><topic>Begomovirus</topic><topic>Biochemistry</topic><topic>Biomedical and Life Sciences</topic><topic>Biotechnology</topic><topic>Chromosome 10</topic><topic>Chromosome Mapping</topic><topic>Chromosomes</topic><topic>Chromosomes, Plant</topic><topic>Disease resistance</topic><topic>Disease Resistance - genetics</topic><topic>DNA viruses</topic><topic>Gene mapping</topic><topic>Genes</topic><topic>Genes, Plant</topic><topic>Genetic aspects</topic><topic>Inbreeding</topic><topic>Leaf-curl</topic><topic>Leaves</topic><topic>Life Sciences</topic><topic>Lycopersicon esculentum - genetics</topic><topic>Lycopersicon esculentum - virology</topic><topic>Original</topic><topic>Original Article</topic><topic>Plant Biochemistry</topic><topic>Plant breeding</topic><topic>Plant Breeding/Biotechnology</topic><topic>Plant Diseases - genetics</topic><topic>Plant Diseases - virology</topic><topic>Plant Genetics and Genomics</topic><topic>Plant viruses</topic><topic>Quantitative Trait Loci</topic><topic>Single-nucleotide polymorphism</topic><topic>Solanaceae</topic><topic>Tomatoes</topic><topic>Yellow leaf</topic><toplevel>peer_reviewed</toplevel><toplevel>online_resources</toplevel><creatorcontrib>Gill, Upinder</creatorcontrib><creatorcontrib>Scott, John W.</creatorcontrib><creatorcontrib>Shekasteband, Reza</creatorcontrib><creatorcontrib>Ogundiwin, Eben</creatorcontrib><creatorcontrib>Schuit, Cees</creatorcontrib><creatorcontrib>Francis, David M.</creatorcontrib><creatorcontrib>Sim, Sung-Chur</creatorcontrib><creatorcontrib>Smith, Hugh</creatorcontrib><creatorcontrib>Hutton, Samuel F.</creatorcontrib><collection>Springer Open Access</collection><collection>Medline</collection><collection>MEDLINE</collection><collection>MEDLINE (Ovid)</collection><collection>MEDLINE</collection><collection>MEDLINE</collection><collection>PubMed</collection><collection>CrossRef</collection><collection>Gale In Context: Science</collection><collection>ProQuest Central (Corporate)</collection><collection>Entomology Abstracts (Full archive)</collection><collection>Neurosciences Abstracts</collection><collection>ProQuest_Health & Medical Collection</collection><collection>ProQuest Central (purchase pre-March 2016)</collection><collection>Biology Database (Alumni Edition)</collection><collection>Medical Database (Alumni Edition)</collection><collection>ProQuest Pharma Collection</collection><collection>Technology Research Database</collection><collection>ProQuest SciTech Collection</collection><collection>ProQuest Natural Science Collection</collection><collection>Hospital Premium Collection</collection><collection>Hospital Premium Collection (Alumni Edition)</collection><collection>ProQuest Central (Alumni) (purchase pre-March 2016)</collection><collection>ProQuest Central (Alumni)</collection><collection>ProQuest Central</collection><collection>ProQuest Central Essentials</collection><collection>Biological Science Collection</collection><collection>AUTh Library subscriptions: ProQuest Central</collection><collection>ProQuest Natural Science Collection</collection><collection>ProQuest One Community College</collection><collection>ProQuest Central</collection><collection>Engineering Research Database</collection><collection>Health Research Premium Collection</collection><collection>Health Research Premium Collection (Alumni)</collection><collection>ProQuest Central Student</collection><collection>SciTech Premium Collection</collection><collection>ProQuest Health & Medical Complete (Alumni)</collection><collection>Biological Sciences</collection><collection>Health & Medical Collection (Alumni Edition)</collection><collection>PML(ProQuest Medical Library)</collection><collection>Biological Science Database</collection><collection>Biotechnology and BioEngineering Abstracts</collection><collection>ProQuest One Academic Eastern Edition (DO NOT USE)</collection><collection>ProQuest One Academic</collection><collection>ProQuest One Academic UKI Edition</collection><collection>Genetics Abstracts</collection><collection>MEDLINE - Academic</collection><collection>PubMed Central (Full Participant titles)</collection><jtitle>Theoretical and applied genetics</jtitle></facets><delivery><delcategory>Remote Search Resource</delcategory><fulltext>fulltext</fulltext></delivery><addata><au>Gill, Upinder</au><au>Scott, John W.</au><au>Shekasteband, Reza</au><au>Ogundiwin, Eben</au><au>Schuit, Cees</au><au>Francis, David M.</au><au>Sim, Sung-Chur</au><au>Smith, Hugh</au><au>Hutton, Samuel F.</au><format>journal</format><genre>article</genre><ristype>JOUR</ristype><atitle>Ty-6, a major begomovirus resistance gene on chromosome 10, is effective against Tomato yellow leaf curl virus and Tomato mottle virus</atitle><jtitle>Theoretical and applied genetics</jtitle><stitle>Theor Appl Genet</stitle><addtitle>Theor Appl Genet</addtitle><date>2019-05-01</date><risdate>2019</risdate><volume>132</volume><issue>5</issue><spage>1543</spage><epage>1554</epage><pages>1543-1554</pages><issn>0040-5752</issn><eissn>1432-2242</eissn><abstract>Key message
Ty
-
6
is a major resistance gene on chromosome 10 of tomato that provides resistance against monopartite and bipartite begomoviruses and complements resistance conferred by the known
Ty
-
3
and
ty
-
5
genes.
Resistance to monopartite and bipartite begomoviruses is an important breeding objective for cultivated tomato. Several begomovirus resistance genes have been introgressed from related
Solanum
species and are available for breeding purposes. In the present study, we mapped an additional locus,
Ty
-
6
, to chromosome 10 of tomato.
Ty
-
6
is effective against both monopartite Tomato yellow leaf curl virus (TYLCV) and bipartite Tomato mottle virus (ToMoV). Gene action is incomplete dominance, with an intermediate resistance response when
Ty
-
6
is heterozygous. Analysis of populations segregating for
Ty
-
6
along with
Ty
-
3
or
ty
-
5
indicates that the highest level of resistance against TYLCV is attained when
Ty
-
6
is combined with an additional resistance allele. Our results also demonstrate that
ty
-
5
is ineffective against ToMoV. Although multiple SNPs linked to
Ty
-
6
were identified and can be used for breeding purposes, none of these were consistently polymorphic between
Ty
-
6
and
ty
-
6
breeding lines. Further research is underway to generate resequencing data for several
Ty
-
6
inbred lines for the discovery of additional sequence polymorphisms that can be used for fine mapping and characterizing the
Ty
-
6
locus.</abstract><cop>Berlin/Heidelberg</cop><pub>Springer Berlin Heidelberg</pub><pmid>30758531</pmid><doi>10.1007/s00122-019-03298-0</doi><tpages>12</tpages><orcidid>https://orcid.org/0000-0002-1635-4861</orcidid><oa>free_for_read</oa></addata></record> |
fulltext | fulltext |
identifier | ISSN: 0040-5752 |
ispartof | Theoretical and applied genetics, 2019-05, Vol.132 (5), p.1543-1554 |
issn | 0040-5752 1432-2242 |
language | eng |
recordid | cdi_pubmedcentral_primary_oai_pubmedcentral_nih_gov_6476845 |
source | Springer Nature |
subjects | Agriculture Begomovirus Biochemistry Biomedical and Life Sciences Biotechnology Chromosome 10 Chromosome Mapping Chromosomes Chromosomes, Plant Disease resistance Disease Resistance - genetics DNA viruses Gene mapping Genes Genes, Plant Genetic aspects Inbreeding Leaf-curl Leaves Life Sciences Lycopersicon esculentum - genetics Lycopersicon esculentum - virology Original Original Article Plant Biochemistry Plant breeding Plant Breeding/Biotechnology Plant Diseases - genetics Plant Diseases - virology Plant Genetics and Genomics Plant viruses Quantitative Trait Loci Single-nucleotide polymorphism Solanaceae Tomatoes Yellow leaf |
title | Ty-6, a major begomovirus resistance gene on chromosome 10, is effective against Tomato yellow leaf curl virus and Tomato mottle virus |
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