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Wine yeast strains of S. cerevisiae do not fully metabolize urea during grape must fermentation. Urea is secreted by yeast cells and it reacts spontaneously with ethanol in wine to form ethyl carbamate, a potential carcinogenic agent for humans. The lack of urea catabolism by yeast in wine may be ascribed to the transcriptional repression of the DUR1,2 gene by good nitrogen sources present in the grape must. We expressed the DUR1,2 gene under control of the S. cerevisiae PGK1 promoter and terminator signals and integrated this DUR1,2 expression cassette, flanked by ura3 sequences, into the URA3 -locus of the industrial wine yeast UC Davis 522. In vivo assays showed that the metabolically engineered industrial strain reduced ethyl carbamate in Chardonnay wine by 89.1%. Analyses of the genotype, phenotype, and transcriptome revealed that the engineered yeast 522EC− is substantially equivalent to the parental 522 strain." /> <meta name="DC.Contributor" content="Joana Coulon" /> <meta name="DC.Contributor" content="John I. Husnik" /> <meta name="DC.Contributor" content="Debra L. Inglis" /> <meta name="DC.Contributor" content="George K. van der Merwe" /> <meta name="DC.Contributor" content="Aline Lonvaud" /> <meta name="DC.Contributor" content="Daniel J. Erasmus" /> <meta name="DC.Contributor" content="Hennie J.J. van Vuuren" /> <meta name="article:published_time" content="2006-06-01" /> <meta name="article:section" content="Article" /> <meta name="citation_title" content="Metabolic Engineering of Saccharomyces cerevisiae to Minimize the Production of Ethyl Carbamate in Wine" /> <meta name="citation_abstract" lang="en" content="<p><i>Saccharomyces cerevisiae</i> metabolizes arginine, one of the major amino acids in grape musts, to ornithine and urea during wine fermentations. Wine yeast strains of <i>S. cerevisiae</i> do not fully metabolize urea during grape must fermentation. Urea is secreted by yeast cells and it reacts spontaneously with ethanol in wine to form ethyl carbamate, a potential carcinogenic agent for humans. The lack of urea catabolism by yeast in wine may be ascribed to the transcriptional repression of the <i>DUR1,2</i> gene by good nitrogen sources present in the grape must. We expressed the <i>DUR1,2</i> gene under control of the <i>S. cerevisiae PGK1</i> promoter and terminator signals and integrated this <i>DUR1,2</i> expression cassette, flanked by ura3 sequences, into the <i>URA3</i>-locus of the industrial wine yeast UC Davis 522. In vivo assays showed that the metabolically engineered industrial strain reduced ethyl carbamate in Chardonnay wine by 89.1%. Analyses of the genotype, phenotype, and transcriptome revealed that the engineered yeast 522<sup>EC−</sup> is substantially equivalent to the parental 522 strain.</p>" /> <meta name="citation_journal_title" content="American Journal of Enology and Viticulture" /> <meta name="citation_publisher" content="American Journal of Enology and Viticulture" /> <meta name="citation_publication_date" content="2006/06/01" /> <meta name="citation_mjid" content="ajev;57/2/113" /> <meta name="citation_id" content="57/2/113" /> <meta name="citation_public_url" content="https://www.ajevonline.org/content/57/2/113" /> <meta name="citation_abstract_html_url" content="https://www.ajevonline.org/content/57/2/113.abstract" /> <meta name="citation_full_html_url" content="https://www.ajevonline.org/content/57/2/113.full" /> <meta name="citation_pdf_url" content="https://www.ajevonline.org/content/ajev/57/2/113.full.pdf" /> <meta name="citation_issn" content="0002-9254" /> <meta name="citation_journal_abbrev" content="Am J Enol Vitic." /> <meta name="citation_doi" content="10.5344/ajev.2006.57.2.113" /> <meta name="citation_volume" content="57" /> <meta name="citation_issue" content="2" /> <meta name="citation_article_type" content="Research Article" /> <meta name="citation_section" content="Article" /> <meta name="citation_firstpage" content="113" /> <meta name="citation_lastpage" content="124" /> <meta name="citation_author" content="Joana Coulon" /> <meta name="citation_author_institution" content="Ph.D. student, Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; Ph.D. students, Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada." /> <meta name="citation_author" content="John I. Husnik" /> <meta name="citation_author_institution" content="Ph.D. student, Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; Ph.D. students, Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada." /> <meta name="citation_author" content="Debra L. Inglis" /> <meta name="citation_author_institution" content="Ph.D. student, Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; Ph.D. students, Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada." /> <meta name="citation_author" content="George K. van der Merwe" /> <meta name="citation_author_institution" content="Ph.D. student, Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; Ph.D. students, Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada." /> <meta name="citation_author" content="Aline Lonvaud" /> <meta name="citation_author_institution" content="Ph.D. student, Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; Ph.D. students, Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada." /> <meta name="citation_author" content="Daniel J. Erasmus" /> <meta name="citation_author_institution" content="Ph.D. student, Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; Ph.D. students, Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada." /> <meta name="citation_author" content="Hennie J.J. van Vuuren" /> <meta name="citation_author_institution" content="Ph.D. student, Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; Ph.D. students, Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada." /> <meta name="citation_author_email" content="hjjvv@interchange.ubc.ca" /> <meta name="citation_reference" content="An, D., and C.S. 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The lack of urea catabolism by yeast in wine may be ascribed to the transcriptional repression of the DUR1,2 gene by good nitrogen sources present in the grape must. We expressed the DUR1,2 gene under control of the S. cerevisiae PGK1 promoter and terminator signals and integrated this DUR1,2 expression cassette, flanked by ura3 sequences, into the URA3 -locus of the industrial wine yeast UC Davis 522. In vivo assays showed that the metabolically engineered industrial strain reduced ethyl carbamate in Chardonnay wine by 89.1%. Analyses of the genotype, phenotype, and transcriptome revealed that the engineered yeast 522EC− is substantially equivalent to the parental 522 strain." /> <meta name="og-title" property="og:title" content="Metabolic Engineering of Saccharomyces cerevisiae to Minimize the Production of Ethyl Carbamate in Wine" /> <meta name="og-url" property="og:url" content="https://www.ajevonline.org/content/57/2/113" /> <meta name="og-site-name" property="og:site_name" content="American Journal of Enology and Viticulture" /> <meta name="og-description" property="og:description" content="Saccharomyces cerevisiae metabolizes arginine, one of the major amino acids in grape musts, to ornithine and urea during wine fermentations. Wine yeast strains of S. cerevisiae do not fully metabolize urea during grape must fermentation. Urea is secreted by yeast cells and it reacts spontaneously with ethanol in wine to form ethyl carbamate, a potential carcinogenic agent for humans. The lack of urea catabolism by yeast in wine may be ascribed to the transcriptional repression of the DUR1,2 gene by good nitrogen sources present in the grape must. We expressed the DUR1,2 gene under control of the S. cerevisiae PGK1 promoter and terminator signals and integrated this DUR1,2 expression cassette, flanked by ura3 sequences, into the URA3 -locus of the industrial wine yeast UC Davis 522. In vivo assays showed that the metabolically engineered industrial strain reduced ethyl carbamate in Chardonnay wine by 89.1%. Analyses of the genotype, phenotype, and transcriptome revealed that the engineered yeast 522EC− is substantially equivalent to the parental 522 strain." /> <meta name="og-type" property="og:type" content="article" /> <meta name="og-image" property="og:image" content="" /> <meta name="viewport" content="width=device-width, initial-scale=1" /> <link rel="alternate" type="application/vnd.ms-powerpoint" title="Powerpoint" href="/content/57/2/113.ppt" /> <link rel="alternate" type="text/plain" title="Full Text (Plain)" href="/content/57/2/113.full.txt" /> <link rel="alternate" type="application/pdf" title="Full Text (PDF)" href="/content/57/2/113.full.pdf" /> <link rel="canonical" href="https://www.ajevonline.org/content/57/2/113" /> <link rel="shortlink" href="/node/5278" /> <title>Metabolic Engineering of Saccharomyces cerevisiae to Minimize the Production of Ethyl Carbamate in Wine | American Journal of Enology and Viticulture</title> <link type="text/css" rel="stylesheet" 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</header> <section id="section-content" class="section section-content"> <div id="zone-content" class="zone zone-content clearfix container-30"> <div class="grid-28 suffix-1 prefix-1 region region-content" id="region-content"> <div class="region-inner region-content-inner"> <a id="main-content"></a> <div class="block block-system block-main block-system-main odd block-without-title" id="block-system-main"> <div class="block-inner clearfix"> <div class="content clearfix"> <div class="panel-display panels-960-layout jcore-2col-layout" > <div class="panel-row-wrapper panel-row-first clearfix"> <div class="top-wrapper"> <div class="panel-panel panel-region-top"> <div class="inside"><div class="panel-pane pane-highwire-article-citation" > <div class="pane-content"> <div class="highwire-article-citation highwire-citation-type-highwire-article node5278--3" data-node-nid="5278" id="top-node-5278--41439355453" data-pisa="ajev;57/2/113" data-pisa-master="ajev;57/2/113" data-apath="/ajev/57/2/113.atom" data-hw-author-tooltip-instance="highwire_author_tooltip"><div class="highwire-cite highwire-cite-highwire-article highwire-citation-jnl-asev-styles-article-title-complete clearfix has-author-tooltip"> <div class="highwire-cite-overline"><span class="highwire-cite-metadata-art-cat highwire-cite-metadata">Article</span></div> <h1 class="highwire-cite-title" id="page-title">Metabolic Engineering of <em>Saccharomyces cerevisiae</em> to Minimize the Production of Ethyl Carbamate in Wine</h1> <div class="highwire-cite-authors"><span class="highwire-citation-authors"><span class="highwire-citation-author first" data-delta="0"><span class="nlm-given-names">Joana</span> <span class="nlm-surname">Coulon</span></span>, <span class="highwire-citation-author" data-delta="1"><span class="nlm-given-names">John I.</span> <span class="nlm-surname">Husnik</span></span>, <span class="highwire-citation-author" data-delta="2"><span class="nlm-given-names">Debra L.</span> <span class="nlm-surname">Inglis</span></span>, <span class="highwire-citation-author" data-delta="3"><span class="nlm-given-names">George K.</span> <span class="nlm-surname">van der Merwe</span></span>, <span class="highwire-citation-author" data-delta="4"><span class="nlm-given-names">Aline</span> <span class="nlm-surname">Lonvaud</span></span>, <span class="highwire-citation-author" data-delta="5"><span class="nlm-given-names">Daniel J.</span> <span class="nlm-surname">Erasmus</span></span>, <span class="highwire-citation-author" data-delta="6"><span class="nlm-given-names">Hennie J.J.</span> <span class="nlm-surname">van Vuuren</span></span></span></div> <div class="highwire-cite-metadata"><span class="highwire-cite-metadata-journal highwire-cite-metadata">Am J Enol Vitic.  </span><span class="highwire-cite-metadata-date highwire-cite-metadata">2006  </span><span class="highwire-cite-metadata-volume highwire-cite-metadata">57: </span><span class="highwire-cite-metadata-pages highwire-cite-metadata">113-124  </span><span class="highwire-cite-metadata-doi highwire-cite-metadata">;<span class="label"> DOI:</span> 10.5344/ajev.2006.57.2.113 </span></div> </div> <div id="hw-article-author-popups-top-node-5278--41439355453" style="display: none;"><div class="author-tooltip-0"><div class="author-tooltip-name">Joana Coulon </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span>Ph.D. student, <span class='nlm-sup'>5</span>Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; <span class='nlm-sup'>2, 6</span>Ph.D. students, <span class='nlm-sup'>7</span>Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; <span class='nlm-sup'>3</span>Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and <span class='nlm-sup'>4</span>Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada.</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Joana%2BCoulon%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Coulon%20J&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="ads-search-link"><a href="http://adsabs.harvard.edu/cgi-bin/nph-abs_connect?return_req=no_params&author=Coulon%2C%20J.&db_key=PRE&arxiv_sel=gr-qc" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on ADS search</a></li><li class="author-tooltip-agricola-link"><a href="/lookup/external-ref?access_num=Coulon&link_type=AGRICOLA" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Agricola</a></li><li class="author-site-search-link last"><a href="/search/author1%3AJoana%2BCoulon%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li></ul></div><div class="author-tooltip-1"><div class="author-tooltip-name">John I. Husnik </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span>Ph.D. student, <span class='nlm-sup'>5</span>Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; <span class='nlm-sup'>2, 6</span>Ph.D. students, <span class='nlm-sup'>7</span>Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; <span class='nlm-sup'>3</span>Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and <span class='nlm-sup'>4</span>Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada.</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=John%2BI.%2BHusnik%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Husnik%20JI&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="ads-search-link"><a href="http://adsabs.harvard.edu/cgi-bin/nph-abs_connect?return_req=no_params&author=Husnik%2C%20J.%20I.&db_key=PRE&arxiv_sel=gr-qc" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on ADS search</a></li><li class="author-tooltip-agricola-link"><a href="/lookup/external-ref?access_num=Husnik&link_type=AGRICOLA" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Agricola</a></li><li class="author-site-search-link last"><a href="/search/author1%3AJohn%2BI.%2BHusnik%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li></ul></div><div class="author-tooltip-2"><div class="author-tooltip-name">Debra L. Inglis </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span>Ph.D. student, <span class='nlm-sup'>5</span>Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; <span class='nlm-sup'>2, 6</span>Ph.D. students, <span class='nlm-sup'>7</span>Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; <span class='nlm-sup'>3</span>Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and <span class='nlm-sup'>4</span>Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada.</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Debra%2BL.%2BInglis%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Inglis%20DL&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="ads-search-link"><a href="http://adsabs.harvard.edu/cgi-bin/nph-abs_connect?return_req=no_params&author=Inglis%2C%20D.%20L.&db_key=PRE&arxiv_sel=gr-qc" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on ADS search</a></li><li class="author-tooltip-agricola-link"><a href="/lookup/external-ref?access_num=Inglis&link_type=AGRICOLA" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Agricola</a></li><li class="author-site-search-link last"><a href="/search/author1%3ADebra%2BL.%2BInglis%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li></ul></div><div class="author-tooltip-3"><div class="author-tooltip-name">George K. van der Merwe </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span>Ph.D. student, <span class='nlm-sup'>5</span>Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; <span class='nlm-sup'>2, 6</span>Ph.D. students, <span class='nlm-sup'>7</span>Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; <span class='nlm-sup'>3</span>Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and <span class='nlm-sup'>4</span>Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada.</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=George%2BK.%2Bvan%2Bder%2BMerwe%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=van%20der%20Merwe%20GK&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="ads-search-link"><a href="http://adsabs.harvard.edu/cgi-bin/nph-abs_connect?return_req=no_params&author=van%20der%20Merwe%2C%20G.%20K.&db_key=PRE&arxiv_sel=gr-qc" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on ADS search</a></li><li class="author-tooltip-agricola-link"><a href="/lookup/external-ref?access_num=van%20der%20Merwe&link_type=AGRICOLA" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Agricola</a></li><li class="author-site-search-link last"><a href="/search/author1%3AGeorge%2BK.%2Bvan%2Bder%2BMerwe%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li></ul></div><div class="author-tooltip-4"><div class="author-tooltip-name">Aline Lonvaud </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span>Ph.D. student, <span class='nlm-sup'>5</span>Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; <span class='nlm-sup'>2, 6</span>Ph.D. students, <span class='nlm-sup'>7</span>Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; <span class='nlm-sup'>3</span>Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and <span class='nlm-sup'>4</span>Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada.</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Aline%2BLonvaud%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Lonvaud%20A&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="ads-search-link"><a href="http://adsabs.harvard.edu/cgi-bin/nph-abs_connect?return_req=no_params&author=Lonvaud%2C%20A.&db_key=PRE&arxiv_sel=gr-qc" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on ADS search</a></li><li class="author-tooltip-agricola-link"><a href="/lookup/external-ref?access_num=Lonvaud&link_type=AGRICOLA" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Agricola</a></li><li class="author-site-search-link last"><a href="/search/author1%3AAline%2BLonvaud%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li></ul></div><div class="author-tooltip-5"><div class="author-tooltip-name">Daniel J. Erasmus </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span>Ph.D. student, <span class='nlm-sup'>5</span>Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; <span class='nlm-sup'>2, 6</span>Ph.D. students, <span class='nlm-sup'>7</span>Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; <span class='nlm-sup'>3</span>Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and <span class='nlm-sup'>4</span>Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada.</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Daniel%2BJ.%2BErasmus%2B" 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data-hide-link-title="0">Search for this author on this site</a></li></ul></div><div class="author-tooltip-6"><div class="author-tooltip-name">Hennie J.J. van Vuuren </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span>Ph.D. student, <span class='nlm-sup'>5</span>Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; <span class='nlm-sup'>2, 6</span>Ph.D. students, <span class='nlm-sup'>7</span>Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; <span class='nlm-sup'>3</span>Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and <span class='nlm-sup'>4</span>Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada.</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Hennie%2BJ.J.%2Bvan%2BVuuren%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=van%20Vuuren%20HJ&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="ads-search-link"><a href="http://adsabs.harvard.edu/cgi-bin/nph-abs_connect?return_req=no_params&author=van%20Vuuren%2C%20H.%20J.&db_key=PRE&arxiv_sel=gr-qc" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on ADS search</a></li><li class="author-tooltip-agricola-link"><a href="/lookup/external-ref?access_num=van%20Vuuren&link_type=AGRICOLA" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on 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Wine yeast strains of <em>S. cerevisiae</em> do not fully metabolize urea during grape must fermentation. Urea is secreted by yeast cells and it reacts spontaneously with ethanol in wine to form ethyl carbamate, a potential carcinogenic agent for humans. The lack of urea catabolism by yeast in wine may be ascribed to the transcriptional repression of the <em>DUR1,2</em> gene by good nitrogen sources present in the grape must. We expressed the <em>DUR1,2</em> gene under control of the <em>S. cerevisiae PGK1</em> promoter and terminator signals and integrated this <em>DUR1,2</em> expression cassette, flanked by ura3 sequences, into the <em>URA3</em>-locus of the industrial wine yeast UC Davis 522. In vivo assays showed that the metabolically engineered industrial strain reduced ethyl carbamate in Chardonnay wine by 89.1%. Analyses of the genotype, phenotype, and transcriptome revealed that the engineered yeast 522<sup>EC−</sup> is substantially equivalent to the parental 522 strain.</p></div><ul class="kwd-group KWD"><li class="kwd"><a href="/keyword/wine" class="hw-term hw-article-keyword hw-article-keyword-wine" rel="nofollow">wine</a></li><li class="kwd"><a href="/keyword/ethyl-carbamate" class="hw-term hw-article-keyword hw-article-keyword-ethyl-carbamate" rel="nofollow">ethyl carbamate</a></li><li class="kwd"><a xmlns:default="http://www.w3.org/1999/xhtml" href="/keyword/dur12" class="hw-term hw-article-keyword hw-article-keyword-dur12" rel="nofollow"> <em xmlns:default="http://www.w3.org/1999/xhtml">DUR1,2</em> </a></li><li class="kwd"><a href="/keyword/carcinogen" class="hw-term hw-article-keyword hw-article-keyword-carcinogen" rel="nofollow">carcinogen</a></li></ul><ul class="history-list"><li xmlns:hwp="http://schema.highwire.org/Journal" class="received" hwp:start="2005-10"><span class="received-label">Received </span>October 2005.</li><li xmlns:hwp="http://schema.highwire.org/Journal" class="rev-recd" hwp:start="2006-01"><span class="rev-recd-label">Revision received </span>January 2006.</li></ul><ul class="copyright-statement"><li class="fn" id="copyright-statement-1">Copyright © 2006 by the American Society for Enology and Viticulture</li></ul><span class="highwire-journal-article-marker-end"></span></div><span class="related-urls"></span></div><a href="https://www.ajevonline.org/content/57/2/113.full" class="hw-link hw-link-article-full-text" data-icon-position="" data-hide-link-title="0">View Full Text</a></div> </div> </div> <div class="panel-separator"></div><div class="panel-pane pane-panels-mini pane-jnl-asev-challenge" > <div class="pane-content"> <div class="panel-display highwire-1col-stacked-layout" id="mini-panel-jnl_asev_challenge"> <div class="panel-row-wrapper clearfix"> <div class="content-wrapper"> <div class="panel-panel panel-region-content"> <div class="inside"><div class="panel-pane pane-snippet pane-asev-sign-in-link" > <div class="pane-content"> <div class="snippet asev-sign-in-link" id="asev-sign-in-link"> <div class="snippet-content"> <h3>Sign in for ASEV members</h3> <p>ASEV Members, please <a href="https://www.asev.org/ajev-online-access?ReturnUrl=https://www.ajevonline.org/content/57/2/113">sign in at ASEV </a> to access the journal online.</p> </div> </div> </div> </div> <div class="panel-separator"></div><div class="panel-pane pane-custom pane-1 login-form-heading" > <div class="pane-content"> <h3>Sign in for Institutional and Non-member Subscribers</h3> </div> </div> <div class="panel-separator"></div><div class="panel-pane pane-highwire-opportunity-challenge" > <div class="pane-content"> <div class='opportunity-challenge-login'><h3>Log in using your username and password</h3><div class="drupal-login compact"><form action="/content/57/2/113" method="post" id="highwire-user-login" accept-charset="UTF-8"><div><div class="form-item form-item-label-invisible form-type-textfield form-item-name"> <label class="element-invisible" for="edit-name">Username <span class="form-required" title="This field is required.">*</span></label> <input placeholder="Username" type="text" id="edit-name" name="name" value="" size="30" maxlength="60" class="form-text required" /> </div> <div class="form-item form-item-label-invisible form-type-password form-item-pass"> <label class="element-invisible" for="edit-pass">Password <span class="form-required" title="This field is required.">*</span></label> <input placeholder="Password" type="password" id="edit-pass" name="pass" size="30" maxlength="128" class="form-text required" /> </div> <input type="hidden" name="form_build_id" value="form-VNnL_-ercKILf5LuVUztZJzh0wHX-4P8_9FnoYUGuO4" /> <input type="hidden" name="form_id" value="highwire_user_login" /> <div class="form-actions form-wrapper" id="edit-actions"><input type="submit" id="edit-submit" name="op" value="Log in" class="form-submit" /></div><div class="reset-password"><a href="/user/password" class="" data-icon-position="" data-hide-link-title="0">Forgot</a> your user name or password?</div></div></form></div></div> </div> </div> <div class="panel-separator"></div><div class="panel-pane pane-highwire-purchase-link" > <div class="pane-content"> <p><a href="/highwire/payment/ppv/5278">Pay Per Article</a> - 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class="highwire-cite-authors"><span class="highwire-citation-authors"><span class="highwire-citation-author first" data-delta="0"><span class="nlm-given-names">Joana</span> <span class="nlm-surname">Coulon</span></span>, <span class="highwire-citation-author" data-delta="1"><span class="nlm-given-names">John I.</span> <span class="nlm-surname">Husnik</span></span>, <span class="highwire-citation-author" data-delta="2"><span class="nlm-given-names">Debra L.</span> <span class="nlm-surname">Inglis</span></span>, <span class="highwire-citation-author" data-delta="3"><span class="nlm-given-names">George K.</span> <span class="nlm-surname">van der Merwe</span></span>, <span class="highwire-citation-author" data-delta="4"><span class="nlm-given-names">Aline</span> <span class="nlm-surname">Lonvaud</span></span>, <span class="highwire-citation-author" data-delta="5"><span class="nlm-given-names">Daniel J.</span> <span class="nlm-surname">Erasmus</span></span>, <span class="highwire-citation-author" data-delta="6"><span class="nlm-given-names">Hennie J.J.</span> <span class="nlm-surname">van Vuuren</span></span></span></div> <div class="highwire-cite-metadata"><span class="highwire-cite-metadata-journal highwire-cite-metadata">Am J Enol Vitic.  </span><span class="highwire-cite-metadata-date highwire-cite-metadata">2006  </span><span class="highwire-cite-metadata-volume highwire-cite-metadata">57: </span><span class="highwire-cite-metadata-pages highwire-cite-metadata">113-124  </span><span class="highwire-cite-metadata-doi highwire-cite-metadata">;<span class="label"> DOI:</span> 10.5344/ajev.2006.57.2.113 </span></div> </div> <div id="hw-article-author-popups-citation-node-5278--2222522498" style="display: none;"><div class="author-tooltip-0"><div class="author-tooltip-name">Joana Coulon </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span>Ph.D. student, <span class='nlm-sup'>5</span>Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; <span class='nlm-sup'>2, 6</span>Ph.D. students, <span class='nlm-sup'>7</span>Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; <span class='nlm-sup'>3</span>Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and <span class='nlm-sup'>4</span>Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada.</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Joana%2BCoulon%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Coulon%20J&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link last"><a href="/search/author1%3AJoana%2BCoulon%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li></ul></div><div class="author-tooltip-1"><div class="author-tooltip-name">John I. Husnik </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span>Ph.D. student, <span class='nlm-sup'>5</span>Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; <span class='nlm-sup'>2, 6</span>Ph.D. students, <span class='nlm-sup'>7</span>Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; <span class='nlm-sup'>3</span>Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and <span class='nlm-sup'>4</span>Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada.</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=John%2BI.%2BHusnik%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Husnik%20JI&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link last"><a href="/search/author1%3AJohn%2BI.%2BHusnik%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li></ul></div><div class="author-tooltip-2"><div class="author-tooltip-name">Debra L. Inglis </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span>Ph.D. student, <span class='nlm-sup'>5</span>Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; <span class='nlm-sup'>2, 6</span>Ph.D. students, <span class='nlm-sup'>7</span>Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; <span class='nlm-sup'>3</span>Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and <span class='nlm-sup'>4</span>Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada.</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Debra%2BL.%2BInglis%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Inglis%20DL&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link last"><a href="/search/author1%3ADebra%2BL.%2BInglis%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li></ul></div><div class="author-tooltip-3"><div class="author-tooltip-name">George K. van der Merwe </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span>Ph.D. student, <span class='nlm-sup'>5</span>Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; <span class='nlm-sup'>2, 6</span>Ph.D. students, <span class='nlm-sup'>7</span>Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; <span class='nlm-sup'>3</span>Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and <span class='nlm-sup'>4</span>Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada.</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=George%2BK.%2Bvan%2Bder%2BMerwe%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=van%20der%20Merwe%20GK&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link last"><a href="/search/author1%3AGeorge%2BK.%2Bvan%2Bder%2BMerwe%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li></ul></div><div class="author-tooltip-4"><div class="author-tooltip-name">Aline Lonvaud </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span>Ph.D. student, <span class='nlm-sup'>5</span>Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; <span class='nlm-sup'>2, 6</span>Ph.D. students, <span class='nlm-sup'>7</span>Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; <span class='nlm-sup'>3</span>Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and <span class='nlm-sup'>4</span>Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada.</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Aline%2BLonvaud%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Lonvaud%20A&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link last"><a href="/search/author1%3AAline%2BLonvaud%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li></ul></div><div class="author-tooltip-5"><div class="author-tooltip-name">Daniel J. Erasmus </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span>Ph.D. student, <span class='nlm-sup'>5</span>Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; <span class='nlm-sup'>2, 6</span>Ph.D. students, <span class='nlm-sup'>7</span>Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; <span class='nlm-sup'>3</span>Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and <span class='nlm-sup'>4</span>Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada.</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Daniel%2BJ.%2BErasmus%2B" 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class='nlm-sup'>7</span>Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; <span class='nlm-sup'>3</span>Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and <span class='nlm-sup'>4</span>Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada.</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Hennie%2BJ.J.%2Bvan%2BVuuren%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=van%20Vuuren%20HJ&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li 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class="highwire-article-citation highwire-citation-type-highwire-article node5278 tooltip-enable" data-node-nid="5278" id="node-5278--2625608864" data-pisa="ajev;57/2/113" data-pisa-master="ajev;57/2/113" data-apath="/ajev/57/2/113.atom" data-hw-abstract-tooltip-instance="highwire_abstract_tooltip" title="Metabolic Engineering of Saccharomyces cerevisiae to Minimize the Production of Ethyl Carbamate in Wine" data-url="/highwire/article_citation_preview/5278" data-hw-author-tooltip-instance="highwire_author_tooltip"><div class="highwire-cite highwire-cite-highwire-article highwire-citation-jnl-asev-article-list clearfix has-author-tooltip"> <div class="highwire-cite-access"><span class="highwire-citation-access highwire-citation-access-check" data-pisa-id="ajev;57/2/113" data-atom-uri="/ajev/57/2/113.atom" data-request-view="full"><i class="highwire-access-icon highwire-access-icon-user-access user-access icon-ok-sign" title="You have access" aria-hidden="true" style="display:none;"></i><span class="element-invisible highwire-access-icon highwire-access-icon-user-access" style="display:none;">You have access</span><i class="highwire-access-icon highwire-access-icon-no-access no-access icon-lock" title="Restricted access" aria-hidden="true" style="display:none;"></i><span class="element-invisible highwire-access-icon highwire-access-icon-no-access" style="display:none;">Restricted access</span></span></div> <a href="/content/57/2/113" class="highwire-cite-linked-title" data-icon-position="" data-hide-link-title="0"><span class="highwire-cite-title">Metabolic Engineering of <em>Saccharomyces cerevisiae</em> to Minimize the Production of Ethyl Carbamate in Wine</span></a> <div class="highwire-cite-authors"><span class="highwire-citation-authors"><span class="highwire-citation-author first" data-delta="0"><span class="nlm-given-names">Joana</span> <span class="nlm-surname">Coulon</span></span>, <span class="highwire-citation-author" data-delta="1"><span class="nlm-given-names">John I.</span> <span class="nlm-surname">Husnik</span></span>, <span class="highwire-citation-author" data-delta="2"><span class="nlm-given-names">Debra L.</span> <span class="nlm-surname">Inglis</span></span>, <span class="highwire-citation-author" data-delta="3"><span class="nlm-given-names">George K.</span> <span class="nlm-surname">van der Merwe</span></span>, <span class="highwire-citation-author" data-delta="4"><span class="nlm-given-names">Aline</span> <span class="nlm-surname">Lonvaud</span></span>, <span class="highwire-citation-author" data-delta="5"><span class="nlm-given-names">Daniel J.</span> <span class="nlm-surname">Erasmus</span></span>, <span class="highwire-citation-author" data-delta="6"><span class="nlm-given-names">Hennie J.J.</span> <span class="nlm-surname">van Vuuren</span></span></span></div> <div class="highwire-cite-metadata"><span class="highwire-cite-metadata-journal highwire-cite-metadata">Am J Enol Vitic.  </span><span class="highwire-cite-metadata-date highwire-cite-metadata">2006  </span><span class="highwire-cite-metadata-volume highwire-cite-metadata">57: </span><span class="highwire-cite-metadata-pages highwire-cite-metadata">113-124  </span><span class="highwire-cite-metadata-doi highwire-cite-metadata">;<span class="label"> DOI:</span> 10.5344/ajev.2006.57.2.113 </span></div> </div> <div id="hw-article-author-popups-node-5278--2625608864" style="display: none;"><div class="author-tooltip-0"><div class="author-tooltip-name">Joana Coulon </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span>Ph.D. student, <span class='nlm-sup'>5</span>Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; <span class='nlm-sup'>2, 6</span>Ph.D. students, <span class='nlm-sup'>7</span>Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; <span class='nlm-sup'>3</span>Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and <span class='nlm-sup'>4</span>Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada.</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Joana%2BCoulon%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Coulon%20J&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link last"><a href="/search/author1%3AJoana%2BCoulon%2B" rel="nofollow" class="" data-icon-position="" 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Husnik </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span>Ph.D. student, <span class='nlm-sup'>5</span>Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; <span class='nlm-sup'>2, 6</span>Ph.D. students, <span class='nlm-sup'>7</span>Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; <span class='nlm-sup'>3</span>Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and <span class='nlm-sup'>4</span>Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada.</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=John%2BI.%2BHusnik%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Husnik%20JI&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link last"><a href="/search/author1%3AJohn%2BI.%2BHusnik%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li></ul></div><div class="author-tooltip-2"><div class="author-tooltip-name">Debra L. Inglis </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span>Ph.D. student, <span class='nlm-sup'>5</span>Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; <span class='nlm-sup'>2, 6</span>Ph.D. students, <span class='nlm-sup'>7</span>Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; <span class='nlm-sup'>3</span>Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and <span class='nlm-sup'>4</span>Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada.</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Debra%2BL.%2BInglis%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Inglis%20DL&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link last"><a href="/search/author1%3ADebra%2BL.%2BInglis%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li></ul></div><div class="author-tooltip-3"><div class="author-tooltip-name">George K. van der Merwe </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span>Ph.D. student, <span class='nlm-sup'>5</span>Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; <span class='nlm-sup'>2, 6</span>Ph.D. students, <span class='nlm-sup'>7</span>Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; <span class='nlm-sup'>3</span>Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and <span class='nlm-sup'>4</span>Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada.</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=George%2BK.%2Bvan%2Bder%2BMerwe%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=van%20der%20Merwe%20GK&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link last"><a href="/search/author1%3AGeorge%2BK.%2Bvan%2Bder%2BMerwe%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li></ul></div><div class="author-tooltip-4"><div class="author-tooltip-name">Aline Lonvaud </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span>Ph.D. student, <span class='nlm-sup'>5</span>Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; <span class='nlm-sup'>2, 6</span>Ph.D. students, <span class='nlm-sup'>7</span>Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; <span class='nlm-sup'>3</span>Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and <span class='nlm-sup'>4</span>Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada.</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Aline%2BLonvaud%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=Lonvaud%20A&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li class="author-site-search-link last"><a href="/search/author1%3AAline%2BLonvaud%2B" rel="nofollow" class="" data-icon-position="" data-hide-link-title="0">Search for this author on this site</a></li></ul></div><div class="author-tooltip-5"><div class="author-tooltip-name">Daniel J. Erasmus </div><div class="author-tooltip-affiliation"><span class="author-tooltip-text"><div class='author-affiliation'><span class='nlm-sup'>1</span>Ph.D. student, <span class='nlm-sup'>5</span>Professor, Faculté d’Oenologie, Unité Associée INRA, Université Victor Segalen Bordeaux 2, Talence, France; <span class='nlm-sup'>2, 6</span>Ph.D. students, <span class='nlm-sup'>7</span>Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; <span class='nlm-sup'>3</span>Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and <span class='nlm-sup'>4</span>Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada.</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Daniel%2BJ.%2BErasmus%2B" 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class='nlm-sup'>7</span>Professor and Eagles Chair, Wine Research Centre, The University of British Columbia, Vancouver, BC, Canada; <span class='nlm-sup'>3</span>Associate professor, Cool Climate Oenology and Viticulture Institute, Brock University, St. Catharines, ON, Canada; and <span class='nlm-sup'>4</span>Assistant professor, Department of Molecular and Cellular Biology, University of Guelph, Guelph, ON, Canada.</div></span></div><ul class="author-tooltip-find-more"><li class="author-tooltip-gs-link first"><a href="/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Hennie%2BJ.J.%2Bvan%2BVuuren%2B" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on Google Scholar</a></li><li class="author-tooltip-pubmed-link"><a href="/lookup/external-ref?access_num=van%20Vuuren%20HJ&link_type=AUTHORSEARCH" target="_blank" class="" data-icon-position="" data-hide-link-title="0">Find this author on PubMed</a></li><li 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