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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Temporal and spatial control of gene expression in horticultural crops</title><author><name sortKey="Dutt, Manjul" sort="Dutt, Manjul" uniqKey="Dutt M" first="Manjul" last="Dutt">Manjul Dutt</name><affiliation><nlm:aff id="aff1"><institution>Citrus Research and Education Center, University of Florida</institution>, 700 Experiment Station Road, Lake Alfred, FL 33850,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Dhekney, Sadanand A" sort="Dhekney, Sadanand A" uniqKey="Dhekney S" first="Sadanand A" last="Dhekney">Sadanand A. Dhekney</name><affiliation><nlm:aff id="aff2"><institution>Department of Plant Sciences, Sheridan Research and Extension Center, University of Wyoming</institution>, Sheridan, WY 82801,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Soriano, Leonardo" sort="Soriano, Leonardo" uniqKey="Soriano L" first="Leonardo" last="Soriano">Leonardo Soriano</name><affiliation><nlm:aff id="aff1"><institution>Citrus Research and Education Center, University of Florida</institution>, 700 Experiment Station Road, Lake Alfred, FL 33850,<country>USA</country></nlm:aff></affiliation><affiliation><nlm:aff id="aff3"><institution>Universidade de Sao Paulo, Centro de Energia Nuclear na Agricultura</institution>, Piracicaba,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Kandel, Raju" sort="Kandel, Raju" uniqKey="Kandel R" first="Raju" last="Kandel">Raju Kandel</name><affiliation><nlm:aff id="aff2"><institution>Department of Plant Sciences, Sheridan Research and Extension Center, University of Wyoming</institution>, Sheridan, WY 82801,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Grosser, Jude W" sort="Grosser, Jude W" uniqKey="Grosser J" first="Jude W" last="Grosser">Jude W. Grosser</name><affiliation><nlm:aff id="aff1"><institution>Citrus Research and Education Center, University of Florida</institution>, 700 Experiment Station Road, Lake Alfred, FL 33850,<country>USA</country></nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">26504550</idno><idno type="pmc">4596326</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4596326</idno><idno type="RBID">PMC:4596326</idno><idno type="doi">10.1038/hortres.2014.47</idno><date when="2014">2014</date><idno type="wicri:Area/Pmc/Corpus">000485</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Temporal and spatial control of gene expression in horticultural crops</title><author><name sortKey="Dutt, Manjul" sort="Dutt, Manjul" uniqKey="Dutt M" first="Manjul" last="Dutt">Manjul Dutt</name><affiliation><nlm:aff id="aff1"><institution>Citrus Research and Education Center, University of Florida</institution>, 700 Experiment Station Road, Lake Alfred, FL 33850,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Dhekney, Sadanand A" sort="Dhekney, Sadanand A" uniqKey="Dhekney S" first="Sadanand A" last="Dhekney">Sadanand A. Dhekney</name><affiliation><nlm:aff id="aff2"><institution>Department of Plant Sciences, Sheridan Research and Extension Center, University of Wyoming</institution>, Sheridan, WY 82801,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Soriano, Leonardo" sort="Soriano, Leonardo" uniqKey="Soriano L" first="Leonardo" last="Soriano">Leonardo Soriano</name><affiliation><nlm:aff id="aff1"><institution>Citrus Research and Education Center, University of Florida</institution>, 700 Experiment Station Road, Lake Alfred, FL 33850,<country>USA</country></nlm:aff></affiliation><affiliation><nlm:aff id="aff3"><institution>Universidade de Sao Paulo, Centro de Energia Nuclear na Agricultura</institution>, Piracicaba,<country>Brazil</country></nlm:aff></affiliation></author><author><name sortKey="Kandel, Raju" sort="Kandel, Raju" uniqKey="Kandel R" first="Raju" last="Kandel">Raju Kandel</name><affiliation><nlm:aff id="aff2"><institution>Department of Plant Sciences, Sheridan Research and Extension Center, University of Wyoming</institution>, Sheridan, WY 82801,<country>USA</country></nlm:aff></affiliation></author><author><name sortKey="Grosser, Jude W" sort="Grosser, Jude W" uniqKey="Grosser J" first="Jude W" last="Grosser">Jude W. Grosser</name><affiliation><nlm:aff id="aff1"><institution>Citrus Research and Education Center, University of Florida</institution>, 700 Experiment Station Road, Lake Alfred, FL 33850,<country>USA</country></nlm:aff></affiliation></author></analytic><series><title level="j">Horticulture Research</title><idno type="eISSN">2052-7276</idno><imprint><date when="2014">2014</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>Biotechnology provides plant breeders an additional tool to improve various traits desired by growers and consumers of horticultural crops. It also provides genetic solutions to major problems affecting horticultural crops and can be a means for rapid improvement of a cultivar. With the availability of a number of horticultural genome sequences, it has become relatively easier to utilize these resources to identify DNA sequences for both basic and applied research. Promoters play a key role in plant gene expression and the regulation of gene expression. In recent years, rapid progress has been made on the isolation and evaluation of plant-derived promoters and their use in horticultural crops, as more and more species become amenable to genetic transformation. Our understanding of the tools and techniques of horticultural plant biotechnology has now evolved from a discovery phase to an implementation phase. The availability of a large number of promoters derived from horticultural plants opens up the field for utilization of native sequences and improving crops using precision breeding. In this review, we look at the temporal and spatial control of gene expression in horticultural crops and the usage of a variety of promoters either isolated from horticultural crops or used in horticultural crop improvement.</p></div></front><back><div1 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article-type="review-article"><pmc-dir>properties open_access</pmc-dir>
<front><journal-meta><journal-id journal-id-type="nlm-ta">Hortic Res</journal-id><journal-id journal-id-type="iso-abbrev">Hortic Res</journal-id><journal-title-group><journal-title>Horticulture Research</journal-title></journal-title-group><issn pub-type="epub">2052-7276</issn><publisher><publisher-name>Nature Publishing Group</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">26504550</article-id><article-id pub-id-type="pmc">4596326</article-id><article-id pub-id-type="pii">hortres201447</article-id><article-id pub-id-type="doi">10.1038/hortres.2014.47</article-id><article-categories><subj-group subj-group-type="heading"><subject>Review Article</subject></subj-group></article-categories><title-group><article-title>Temporal and spatial control of gene expression in horticultural crops</article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Dutt</surname><given-names>Manjul</given-names></name><xref ref-type="aff" rid="aff1">1</xref><xref ref-type="corresp" rid="caf1">*</xref></contrib><contrib contrib-type="author"><name><surname>Dhekney</surname><given-names>Sadanand A</given-names></name><xref ref-type="aff" rid="aff2">2</xref></contrib><contrib contrib-type="author"><name><surname>Soriano</surname><given-names>Leonardo</given-names></name><xref ref-type="aff" rid="aff1">1</xref><xref ref-type="aff" rid="aff3">3</xref></contrib><contrib contrib-type="author"><name><surname>Kandel</surname><given-names>Raju</given-names></name><xref ref-type="aff" rid="aff2">2</xref></contrib><contrib contrib-type="author"><name><surname>Grosser</surname><given-names>Jude W</given-names></name><xref ref-type="aff" rid="aff1">1</xref></contrib><aff id="aff1"><label>1</label><institution>Citrus Research and Education Center, University of Florida</institution>, 700 Experiment Station Road, Lake Alfred, FL 33850,<country>USA</country></aff><aff id="aff2"><label>2</label><institution>Department of Plant Sciences, Sheridan Research and Extension Center, University of Wyoming</institution>, Sheridan, WY 82801,<country>USA</country></aff><aff id="aff3"><label>3</label><institution>Universidade de Sao Paulo, Centro de Energia Nuclear na Agricultura</institution>, Piracicaba,<country>Brazil</country></aff></contrib-group><author-notes><corresp id="caf1"><label>*</label><email>manjul@ufl.edu</email></corresp></author-notes><pub-date pub-type="epub"><day>24</day><month>09</month><year>2014</year></pub-date><pub-date pub-type="collection"><year>2014</year></pub-date><volume>1</volume><fpage>14047</fpage><lpage></lpage><history><date date-type="received"><day>25</day><month>06</month><year>2014</year></date><date date-type="rev-recd"><day>19</day><month>07</month><year>2014</year></date><date date-type="accepted"><day>06</day><month>08</month><year>2014</year></date></history><permissions><copyright-statement>Copyright © 2014 Nanjing Agricultural University</copyright-statement><copyright-year>2014</copyright-year><copyright-holder>Nanjing Agricultural University</copyright-holder><license license-type="open-access" xlink:href="http://creativecommons.org/licenses/by-nc-nd/3.0/"><pmc-comment>author-paid</pmc-comment>
<license-p>This work is licensed under a Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. The images or other third party material in this article are included in the article's Creative Commons license, unless indicated otherwise in the credit line; if the material is not included under the Creative Commons license, users will need to obtain permission from the license holder to reproduce the material. To view a copy of this license, visit <ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by-nc-nd/3.0/">http://creativecommons.org/licenses/by-nc-nd/3.0/</ext-link></license-p></license></permissions><abstract><p>Biotechnology provides plant breeders an additional tool to improve various traits desired by growers and consumers of horticultural crops. It also provides genetic solutions to major problems affecting horticultural crops and can be a means for rapid improvement of a cultivar. With the availability of a number of horticultural genome sequences, it has become relatively easier to utilize these resources to identify DNA sequences for both basic and applied research. Promoters play a key role in plant gene expression and the regulation of gene expression. In recent years, rapid progress has been made on the isolation and evaluation of plant-derived promoters and their use in horticultural crops, as more and more species become amenable to genetic transformation. Our understanding of the tools and techniques of horticultural plant biotechnology has now evolved from a discovery phase to an implementation phase. The availability of a large number of promoters derived from horticultural plants opens up the field for utilization of native sequences and improving crops using precision breeding. In this review, we look at the temporal and spatial control of gene expression in horticultural crops and the usage of a variety of promoters either isolated from horticultural crops or used in horticultural crop improvement.</p></abstract></article-meta></front><body><sec sec-type="intro"><title>Introduction</title><p>Gene expression in prokaryotes as well as in eukaryotes is regulated quantitatively and qualitatively by specific upstream DNA sequences.<sup><xref ref-type="bibr" rid="bib1">1</xref></sup> These DNA sequences are commonly known as gene promoters. Initiation of transcription is in turn mediated by proteins that recognize specific DNA sequences in the promoter, thereby inducing RNA polymerase activity.<sup><xref ref-type="bibr" rid="bib2">2</xref>,<xref ref-type="bibr" rid="bib3">3</xref></sup> Promoters regulate gene expression through DNA recognition sequences, which interact with basic transcription initiation complexes and numerous transcription factors.<sup><xref ref-type="bibr" rid="bib4">4</xref></sup> Such DNA recognition sequences usually include a core promoter with upstream enhancer sequences located close to the structural portion of the gene.<sup><xref ref-type="bibr" rid="bib2">2</xref></sup> Transcription can be activated by these enhancer sequences independent of their location, distance or orientation with respect to the genes promoters.<sup><xref ref-type="bibr" rid="bib5">5</xref></sup></p><p>Promoters in general are divided into two regions: a core promoter region and upstream regulatory regions.<sup><xref ref-type="bibr" rid="bib6">6</xref></sup> The core promoter consists of a 50–100 bp sequence adjacent to the transcription initiation site and flanking sequences.<sup><xref ref-type="bibr" rid="bib7">7</xref></sup> This region interacts with the general transcription machinery <sup><xref ref-type="bibr" rid="bib8">8</xref></sup> and ensures the accurate initiation of transcription by RNA polymerase II.<sup><xref ref-type="bibr" rid="bib9">9</xref></sup> The core promoter consists of two key genetic elements: the TATA box (present in most promoters) and/or an initiator (Inr) element overlapping the transcription start site.<sup><xref ref-type="bibr" rid="bib10">10</xref>,<xref ref-type="bibr" rid="bib11">11</xref></sup> The initiator element binds trans-acting factors for the placement of the start site<sup><xref ref-type="bibr" rid="bib12 bib13 bib14">12–14</xref></sup> and can also mediate transcription initiation in some TATA-less promoters.<sup><xref ref-type="bibr" rid="bib15">15</xref>,<xref ref-type="bibr" rid="bib16">16</xref></sup> The upstream promoter regions of 1–2 kb or more contains several <italic>cis-</italic>regulatory elements that serve as the binding sites for gene-specific regulators.<sup><xref ref-type="bibr" rid="bib7">7</xref></sup> The regulatory sequences that play a role in the qualitative specificity of gene expression have been intensely studied.<sup><xref ref-type="bibr" rid="bib17 bib18 bib19">17–19</xref></sup> Several regulatory sequences present upstream of the 5′ region of plant genes include multiple <italic>cis-</italic>regulatory elements whose distribution and presence contribute to the expression pattern of that particular gene. This interaction between the <italic>cis-</italic>acting elements and the transcription factors is key in the regulation of gene expression.<sup><xref ref-type="bibr" rid="bib20">20</xref></sup> The presence of several <italic>cis-</italic>acting elements can contribute to the complex expression profile of a particular gene<sup><xref ref-type="bibr" rid="bib2">2</xref></sup> and their differential combinatorial interactions with the transcription factors result in expression of the adjacent gene to be either constitutive, induced by external factors, tissue-specific or some combination of these.<sup><xref ref-type="bibr" rid="bib21">21</xref>,<xref ref-type="bibr" rid="bib22">22</xref></sup></p><p>The first biotech crop commercialized in the United States was a horticultural commodity: the Flavr Savr tomato, which was submitted for approval in 1992 and released for consumption in 1994.<sup><xref ref-type="bibr" rid="bib23 bib24 bib25">23–25</xref></sup> Numerous horticultural crops in the last 20 years have since been transformed with a wide range of genes and promoter elements. In most studies the introduced genes are controlled by constitutive promoters—the most popular being the 35S promoter obtained from the <italic>Cauliflower mosaic virus</italic> (CaMV).<sup><xref ref-type="bibr" rid="bib26">26</xref>,<xref ref-type="bibr" rid="bib27">27</xref></sup> In many cases, constitutive gene expression may not be required, especially when this does not serve a beneficial purpose.<sup><xref ref-type="bibr" rid="bib28">28</xref>,<xref ref-type="bibr" rid="bib29">29</xref></sup> In such cases, targeted gene expression using tissue-specific or inducible promoters can often provide advantages not seen using constitutive promoters.<sup><xref ref-type="bibr" rid="bib30">30</xref></sup> In recent years, there has been a boom in the availability of promoter information in many promoter databases.<sup><xref ref-type="bibr" rid="bib31 bib32 bib33">31–33</xref></sup> This wealth of information enables the researcher to better understand the role of promoters and their control on plant growth and development. It also allows for the development of improved cultivars containing desirable traits.<sup><xref ref-type="bibr" rid="bib34">34</xref>,<xref ref-type="bibr" rid="bib35">35</xref></sup> In this review, we look at the different promoter elements either isolated from horticultural crops or used to genetically modify a horticultural crop for improved traits (<xref rid="tbl1" ref-type="table">Table 1</xref>).</p></sec><sec><title>Promoters used for constitutive gene expression</title><p>Constitutive promoters direct gene expression uniformly in most tissues and cells at all stages of plant growth and development. Constitutive promoters confer high levels of transgene expression when transferred to plant cells. They generally consist of a core DNA sequence (core promoter) along with other regulatory elements such as enhancers, silencers and other DNA sequences, which interact with DNA binding proteins (transcription factors) to drive transgene expression in various plant cells.<sup><xref ref-type="bibr" rid="bib27">27</xref></sup> Constitutive promoters may provide ectopic gene expression in transgenic plants, not otherwise observed under normal conditions. Significantly variable results may be observed from the use of a constitutive promoter in a monocotyledonous and dicotyledonous species, which makes it essential to identify candidate promoters for specific groups to ensure high transgene expression levels.<sup><xref ref-type="bibr" rid="bib36">36</xref></sup> Most constitutive promoters used for production of transgenic plants derive their origin from viral sequences. Advances in plant genome sequencing initiatives and availability of public genomic databases have led to the identification of numerous plant-derived constitutive promoters, which are increasingly being used in plant transformation.</p><p>The <italic>Cauliflower mosaic virus</italic> 35S (CaMV 35S or simply 35S) promoter is by far the most widely used promoter in plant transformation.<sup><xref ref-type="bibr" rid="bib37">37</xref></sup> The promoter is capable of conferring high gene expression levels in most cells when transferred to plants.<sup><xref ref-type="bibr" rid="bib38">38</xref></sup> The 35S promoter has been extensively studied to identify key regulatory sequences that function to provide high gene expression levels.<sup><xref ref-type="bibr" rid="bib27">27</xref></sup> Sequences analyses of the 35S promoter reveal the presence of several regulatory elements that are dispersed among the entire promoter length. The promoter consists of two domains A and B, which are further subdivided into several subdomains.<sup><xref ref-type="bibr" rid="bib27">27</xref></sup> Deletion analyses studies identified specific <italic>cis-</italic>elements in these subdomains that confer expression in specific tissues of above and below-ground plant parts. Various combinations of <italic>cis</italic>-elements of the 35S promoter can produce gene expression patterns that are not observed with the sole use of such elements, which suggests an interaction between <italic>cis</italic>-elements for expression at various plant growth and developmental stages.<sup><xref ref-type="bibr" rid="bib27">27</xref></sup> Although the 35S promoter is considered to direct constitutive expression, varied expression effects result from its interaction with environmental factors<sup><xref ref-type="bibr" rid="bib39">39</xref></sup> and physiological state of plant development.<sup><xref ref-type="bibr" rid="bib40">40</xref></sup> Gene expression by the 35S promoter also appears to be species-dependent. For instance, high GUS expression levels were observed in pollen of transgenic strawberry plants when a 35S promoter was used, but no expression could be detected in transgenic tomato or petunia plants with similar construct configurations.<sup><xref ref-type="bibr" rid="bib41 bib42 bib43">41–43</xref></sup> In other cases, transgenic chrysanthemum expressing a GUS gene under the control of the 35S promoter exhibited weak transgene expression levels.<sup><xref ref-type="bibr" rid="bib44">44</xref></sup></p><p>In-depth functional analyses of regulatory elements present in the 35S promoter has increased our understanding of the role of individual <italic>cis</italic> and enhancer elements in driving gene transcription.<sup><xref ref-type="bibr" rid="bib27">27</xref>,<xref ref-type="bibr" rid="bib45">45</xref></sup> Such information has been exploited to produce chimeric versions of the 35S promoter that contain duplicated <italic>cis</italic> or enhancer elements.<sup><xref ref-type="bibr" rid="bib46">46</xref></sup> Inclusion of additional viral- and plant-derived sequences in various combinations can provide additional synergy to the 35S promoter. Duplication of 35S enhancer elements in unique orientation along with the core promoter can greatly assist in driving high levels of several genes in a single transformation cassette.<sup><xref ref-type="bibr" rid="bib47">47</xref></sup></p><p>Genetic constructs containing a 35S-derived core promoter and either single or duplicated enhancer elements that controlled fusion gene expression, were arranged in a unidirectional (tandem) or bidirectional (divergent) orientation. Significantly high levels of GFP and GUS expression was observed in grapevine somatic embryos and plants transformed with constructs containing a bidirectional duplex promoter complex, where core promoters and duplicated enhancer elements were arranged in a divergent orientation. This phenomenon was attributed to synergistic activity of core promoters and enhancers arranged in a unique orientation.<sup><xref ref-type="bibr" rid="bib48">48</xref></sup> Similar results were obtained when a grapevine <italic>MybA1</italic> transcription factor encoding anthocyanin expression was fused to viral promoters in various arrangements.<sup><xref ref-type="bibr" rid="bib49">49</xref></sup></p><p>The 35S promoter has been extensively used in horticultural crops for improving abiotic and biotic stress tolerance and quality traits, and for modification of plant architecture. Transgenic papaya that expressed a viral coat protein gene driven by a 35S promoter exhibited enhanced resistance against papaya ring spot virus resistance.<sup><xref ref-type="bibr" rid="bib50">50</xref></sup> Following extensive field trials to confirm stability of resistance, the transgenic lines were used in breeding programs to produce virus resistant cultivars, which were deregulated and released for commercial production.<sup><xref ref-type="bibr" rid="bib51">51</xref></sup> Transgenic ‘Honey Sweet’ plums expressing a plum pox virus coat protein under the control of the 35S promoter exhibited enhanced resistance to plum pox virus, the causal agent of Sharka disease of plum.<sup><xref ref-type="bibr" rid="bib52">52</xref>,<xref ref-type="bibr" rid="bib53">53</xref></sup> ‘Honey Sweet’ was cleared for commercial production in the United States in 2010 following extensive studies by appropriate regulatory agencies.<sup><xref ref-type="bibr" rid="bib54">54</xref></sup> Similar strategies have been used to incorporate virus resistance in other fruit and vegetable crops.<sup><xref ref-type="bibr" rid="bib55">55</xref>,<xref ref-type="bibr" rid="bib56">56</xref></sup></p><p>The 35S promoter has also been fused to a number of genes coding for antimicrobial proteins to improve fungal and bacterial resistance. Improved scab resistance was demonstrated in transgenic apple that expressed a <italic>mbr4</italic> gene driven by the 35S promoter.<sup><xref ref-type="bibr" rid="bib57">57</xref></sup> Genetically, engineered cacao plants constitutively expressing a chitinase gene showed decreased growth of <italic>Colletotrichum gleosporiodes</italic> and reduced symptoms of necrosis compared to the controls.<sup><xref ref-type="bibr" rid="bib58">58</xref></sup></p><p>Transgenic <italic>Citrus</italic> plants expressing an antimicrobial peptide under control of a double-enhanced 35S promoter exhibited reduced symptoms of <italic>Citrus</italic> scab in greenhouse trials.<sup><xref ref-type="bibr" rid="bib59">59</xref></sup> Similar results were observed in transgenic strawberries expressing an antimicrobial protein.<sup><xref ref-type="bibr" rid="bib60">60</xref></sup> Transgenic grapevines expressing either antifungal or antibacterial genes under control of the 35S promoter exhibited enhanced disease resistance and are currently in advanced stages of field testing.<sup><xref ref-type="bibr" rid="bib61">61</xref></sup> A number of PR proteins under the control of the 35S promoter have also been employed to engineer disease resistance in ornamentals. Delayed symptoms of fungal diseases was observed in transgenic lines compared to the controls.<sup><xref ref-type="bibr" rid="bib62">62</xref></sup> Transgenic roses constitutively expressing an antimicrobial peptide exhibited resistance to powdery mildew in greenhouse trials.<sup><xref ref-type="bibr" rid="bib63">63</xref></sup> In other studies insect resistant transgenic fruits and vegetables have been produced by expressing a wide array of genes driven by the 35S promoter.<sup><xref ref-type="bibr" rid="bib64 bib65 bib66 bib67">64–67</xref></sup></p><p>Transgenic horticultural crops with abiotic stress tolerance have been developed by constitutively expressing drought, cold and salinity-related genes. The <italic>Arabidopsis</italic> CBF transcription factors and its homologues from several species have been transferred to a number of fruit and vegetable crops for improving cold/chilling and drought tolerance.<sup><xref ref-type="bibr" rid="bib68 bib69 bib70 bib71 bib72">68–72</xref></sup> A number of antiporter and vacuolar genes have been utilized for enhancing salinity tolerance in several plant species.<sup><xref ref-type="bibr" rid="bib73 bib74 bib75">73–75</xref></sup></p><p>The 35S promoter has been frequently used to downregulate genes involved in ethylene biosynthesis or fruit ripening and subsequently enhance shelf life and fruit quality.<sup><xref ref-type="bibr" rid="bib76 bib77 bib78">76–78</xref></sup> Transgenic tomatoes expressing antisense versions of genes responsible for fruit softening under control of a 35S promoter exhibited enhanced shelf life due to their ability to inhibit or reduce fruit-specific enzymes responsible for softening of the fruit during the ripening process.<sup><xref ref-type="bibr" rid="bib79 bib80 bib81">79–81</xref></sup> Suppression of ripening-specific <italic>N</italic>-glycoprotein modifying enzymes in tomato resulted in an increase in fruit shelf life without adversely affecting other qualitative characteristics.<sup><xref ref-type="bibr" rid="bib82">82</xref></sup></p><p>The 35S promoter has also been used in a number of ornamental crops to modify plant structure, flower color and engineer floral scent in flowers that normally do not produce any fragrance. Enhanced anthocyanin production was observed in transgenic tobacco and petunia plants when a maize leaf color transcription factor was constitutively expressed by a 35S promoter.<sup><xref ref-type="bibr" rid="bib83">83</xref>,<xref ref-type="bibr" rid="bib84">84</xref></sup> Transgenic flower crops with unique colorations not generally observed in wild populations have been created by isolating genes from the pigment biosynthesis pathway and placing them under control of the 35S promoter.<sup><xref ref-type="bibr" rid="bib85">85</xref>,<xref ref-type="bibr" rid="bib86">86</xref></sup> Transgenic roses and carnations expressing unique colorations were also produced and released for commercial production. Transgenic petunia with reduced height and enhanced lateral branching were produced by constitutively expressing a zinc finger transcription factor.<sup><xref ref-type="bibr" rid="bib87">87</xref></sup> The enhanced branching patterns were attributed to alterations in cytokinin metabolism and increase in specific forms of cytokinins. Flowers with improved shelf life have also been produced by expressing various genes under the control of the 35S promoter.<sup><xref ref-type="bibr" rid="bib88">88</xref></sup> Other efforts to improve traits in ornamental plants include the production of dwarf and compact plants and enhanced leaf color.<sup><xref ref-type="bibr" rid="bib89">89</xref></sup> Several attempts to introduce floral scent have been made using genetic engineering; such efforts have achieved partial success, mainly in part due to the absence of key enzymes or precursors that are required for the biosynthesis of the final biochemical compound.<sup><xref ref-type="bibr" rid="bib90">90</xref></sup></p><p>Chimeric promoters that drive constitutive gene expression are created by combining elements from viral-derived sequences other than the 35S promoter.<sup><xref ref-type="bibr" rid="bib91">91</xref>,<xref ref-type="bibr" rid="bib92">92</xref></sup> The Cassava vein mosaic virus (<italic>CVMV</italic>), figwort mosaic virus and Cestrum Yellow Leaf Curling Virus (<italic>CMPS</italic>) have been used to identify regulatory elements that would drive high levels of constitutive gene expression in plants.<sup><xref ref-type="bibr" rid="bib48">48</xref>,<xref ref-type="bibr" rid="bib93 bib94 bib95">93–95</xref></sup> Such chimeric promoters created through shuffling of regulatory elements and inclusion of plant-derived or other viral-derived sequences have shown high levels of transgene expression in several plant species. In some cases, the activity of viral-derived constitutive promoters has been less effective in monocotyledonous species compared to dicotyledonous plant species.<sup><xref ref-type="bibr" rid="bib94">94</xref></sup> In other instances, viral-derived promoter sequences are known to direct high levels of gene expression in a wide array of dicot and monocot species.<sup><xref ref-type="bibr" rid="bib96">96</xref></sup></p><p>Advances in genome sequencing of major crops of commercial importance and availability of high throughput sequence analyses have led to the isolation of several constitutive promoters from plant species. Promoters of constitutively expressed genes such as ubiquitin are ideal candidates due to their ability to drive high gene expression levels in transformed cells. Several grapevine promoters have been isolated from the sequenced genome and analyzed for their ability to direct gene expression in various plant tissues.<sup><xref ref-type="bibr" rid="bib97">97</xref></sup> Among the various candidates tested, ubiquitin promoters exhibited the highest activity levels when tested in grape somatic embryos and tobacco callus cultures, leaves and floral tissues. Two promoters Ubi-6-1 and Ubi7-2 exhibited gene expression levels comparable to a doubly enhanced 35S promoter when fused to the <italic>gus</italic> and anthocyanin reporter genes. Higher levels of gene expression could be correlated with an increased number of <italic>cis-</italic>elements in these promoters, which underlines the significance of identifying specific sequences in promoter regions for predicting expression levels. An ubiquitin extension promoter (<italic>uep1</italic>) identified in oil palm exhibited constitutive expression in the native species as well as in tobacco, thereby indicating its utility in monocot and dicot groups of plants.<sup><xref ref-type="bibr" rid="bib98">98</xref></sup> A comparison of the activity of plant- and viral-derived promoter sequences in transgenic <italic>Gladioulus</italic> found no differences in expression levels of GUS during the culture stage.<sup><xref ref-type="bibr" rid="bib99">99</xref></sup> However greenhouse-grown transgenic lines exhibited higher gene expression levels when the <italic>gus</italic> gene was driven by an <italic>Arabidopsis</italic>-derived <italic>rolD</italic> promoter. Transgenic chrysanthemums exhibited higher GUS expression levels when fused to a potato <italic>Lhca3.St.1</italic> promoter than the 35S promoter.<sup><xref ref-type="bibr" rid="bib100">100</xref></sup> Such effects were attributed to potential post-transcriptional modifications leading to greater stability of the mRNA and higher expression levels.</p></sec><sec><title>Promoters involved in fruit-specific gene expression</title><p>The ability of constitutive promoters to direct high levels of transgene expression can be a limiting factor when temporal and spatial gene expression patterns are required to achieve manipulation of specific plant organs or developmental stages. Constitutive expression of transcription factors by the 35S promoter may interfere with normal plant development resulting in abnormal phenotypes.<sup><xref ref-type="bibr" rid="bib70">70</xref>,<xref ref-type="bibr" rid="bib101">101</xref></sup> In other cases, the 35S promoter may not be active in specific plant tissues thereby rendering it ineffective for directing high levels of spatial transgene expression.<sup><xref ref-type="bibr" rid="bib43">43</xref></sup> Tissue-specific promoters may be useful for directing transgenic expression in specific plant tissues without interfering with normal plant growth and development processes. A number of promoters involved in various stages of fruit growth, maturity and ripening have been identified and can be used as genetic engineering tools to improve fruit yield, quality and post-harvest shelf life. Fruit-specific promoters with unique positive and negative regulatory elements may function efficiently in restricting tissue-specific expression of genes and avoiding the possibility of abnormal plant growth often observed with constitutive promoters. Fruit-specific promoters from both plant species that exhibit climacteric and non-climacteric ripening patterns have been studied.</p><p>A number of fruit-specific promoters are regulated by ethylene, which is involved in a number of developmental processes including fruit maturity, ripening and senescence. Promoters of ethylene responsive genes such as the <italic>E4</italic> and <italic>E8</italic> genes have been well studied to identify activator and suppressor elements that ensure spatial and temporal gene expression.<sup><xref ref-type="bibr" rid="bib102">102</xref>,<xref ref-type="bibr" rid="bib103">103</xref></sup> Promoters from genes such as the <italic>ACC</italic> ox<italic>idase</italic> and <italic>ACO synthase</italic> isoforms that catalyze the key steps in ethylene biosynthesis have also been analyzed in a number of plant species to identify specific <italic>cis-</italic>elements involved in the regulatory process.<sup><xref ref-type="bibr" rid="bib104 bib105 bib106">104–106</xref></sup> Deletion analysis of a peach <italic>ACC oxidase</italic> promoter fused to the GUS gene revealed the presence of regulatory regions that controlled gene expression at specific stages of fruit ripening.<sup><xref ref-type="bibr" rid="bib107">107</xref></sup> Longer sequences of the promoter enhanced GUS expression in transgenic tomato, which was attributed to the presence of an enhancer element. Genes involved in tomato fruit development from the immature-green to mature-green stages have been identified using large-scale microarray analysis to identify fruit-specific promoters that direct gene expression from ovary development to ripening.<sup><xref ref-type="bibr" rid="bib108">108</xref></sup> Analysis of a sour cherry expansin gene and its promoter region revealed the presence of a TATA box and several CAAT boxes that are conserved among promoter sequences.<sup><xref ref-type="bibr" rid="bib109">109</xref></sup> Additionally, sequences that were responsive to hormones (ethylene and gibberellins), an anaerobic responsive element, GATA boxes, pyrimidine box and other <italic>cis-</italic>elements that conferred tissue specificity were identified in the 5′ upstream region. Such sequences were highly conserved with previously identified <italic>cis-</italic>elements in other plant species. Promoter deletion analysis studies confirmed specific <italic>cis-</italic>elements that acted as positive regulators of gene expression in fruits at various stages of development. Similar results were obtained with the analysis of a cucumber fruit-specific expansion gene, <italic>CsExp</italic>.<sup><xref ref-type="bibr" rid="bib110">110</xref></sup> In addition to the TATA and CAAT boxes, light and hormone-responsive <italic>cis-</italic>elements with a high degree of homology with other similar elements in other species were identified. Genes responsible for sex expression in cucumber and expressed during fruit development were studied along with their promoter regions.<sup><xref ref-type="bibr" rid="bib111">111</xref></sup> Sequence analysis for two female-specific genes revealed gene duplication except for differences in the promoter regions. No differences were observed in the proximal promoter region of the <italic>CsACS1G</italic> and <italic>CsACS1</italic> genes, which has <italic>cis-</italic>elements that acted as repressors of gibberellins. <italic>In silico</italic> analysis of the distal regions indicated the presence of auxin-responsive elements in the CsACS1G promoter, which could potentially confer responsiveness of this gene to specific hormonal factors and control female sex expression.<sup><xref ref-type="bibr" rid="bib111">111</xref></sup></p><p>The strawberry gene <italic>Faxy1</italic> coding for a fruit-specific β-xylosidase and potentially involved in hemicellulose degradation during fruit ripening was isolated along with its 5′ flanking region.<sup><xref ref-type="bibr" rid="bib112">112</xref></sup> Analysis of the promoter region revealed the presence of several hormone, light and abiotic stress-related regulatory regions in addition to the TATA box and several CAAT boxes. While abscisic acid (ABA) treatment of peduncles enhanced gene expression and protein levels, a reduction was observed with NAA, GA<sub>3</sub> and ethylene treatment thereby indicating the presence of <italic>cis-</italic>elements that were positively and negatively regulated by specific hormones. Light responsive <italic>cis-</italic>elements such as ACE, SP1 G-box and MRE sequences were identified. The promoter region also included a number of cold, drought and heat-responsive elements.</p><p>A number of promoter sequences that are involved in the expression of genes involved in biochemical changes of fruit composition during development and ripening have been studied.<sup><xref ref-type="bibr" rid="bib113 bib114 bib115">113–115</xref></sup> A banana sucrose phosphate synthase (<italic>SPS</italic>) promoter that is involved in sucrose accumulation during ripening was analyzed to identify regulatory elements and their interaction with transcription factors.<sup><xref ref-type="bibr" rid="bib116">116</xref></sup> The presence of <italic>cis-</italic>elements regulated by light and hormonal interactions in addition to the TATA box and CAAT box indicated an interaction of plant hormones and environmental factors during the process of fruit ripening. In watermelon, the ADP-glucose pryophosphorylase gene, which is involved in carbohydrate metabolism during fruit ripening, was negatively regulated in the vegetative tissues.<sup><xref ref-type="bibr" rid="bib115">115</xref></sup> Removal of the <italic>cis-</italic>elements involved in negative regulation by fine promoter deletion analysis led to constitutive expression of the gene in leaf epidermal cells. Novel fruit-specific elements were identified in a cucumisin gene that is expressed in ripe melon fruits.<sup><xref ref-type="bibr" rid="bib114">114</xref></sup> Deletion analysis identified a fruit-specific enhancer element, and an I-box-like sequence, which negatively regulated cucumin biosynthesis in tissues other than the fruit. Similar elements with positive and negative regulatory functions were identified in a <italic>Citrus</italic> C11 promoter that was specifically expressed in juice sacs of ripening lemon fruit.<sup><xref ref-type="bibr" rid="bib117">117</xref></sup> Heterologous expression of the promoter: <italic>gus</italic> chimeric fusion in tomato revealed GUS expression specifically in the anthers and ovaries but not in vegetative tissues.</p><p>Promoters coding for metallothionin expression have been isolated from oil palm and <italic>Citrus</italic>.<sup><xref ref-type="bibr" rid="bib118">118</xref>,<xref ref-type="bibr" rid="bib119">119</xref></sup> The oil palm promoter exhibited higher activity in the mesocarp tissue compared to leaf tissues. A core sequence that specified mesocarp expression while negatively regulating constitutive expression was identified in addition to specific enhancer elements that promoted expression in fruit tissues. Thus, tissue-specific expression appeared to be controlled by the combination of the positive and negative regulatory elements in the promoter region.<sup><xref ref-type="bibr" rid="bib119">119</xref></sup> Analysis of the <italic>Citrus methallothionin</italic> gene indicated the promoter to be in the TATA-less group of plant promoters such as those involved in photosynthesis. A number of fruit-specific <italic>cis</italic>-elements were identified in the promoter region and their function was confirmed by heterologous expression in <italic>Arabidopsis</italic>.</p><p>A number of genes for pigment production in fruits during the ripening phase have been well characterized.<sup><xref ref-type="bibr" rid="bib113">113</xref>,<xref ref-type="bibr" rid="bib120 bib121 bib122">120–122</xref></sup> The <italic>VvMybA1</italic> transcription factor is known to bind to specific regulatory elements of genes involved in the phenylalanine pathway, thereby promoting anthocyanin biosynthesis in grape berries post-veraison. A difference in the production of red and white colored berries in various grape cultivars is attributed to the insertion of a grape retrotransposon element GRET 1, which causes lack of pigment production resulting in white colored berries.<sup><xref ref-type="bibr" rid="bib123">123</xref></sup> Analysis of the grape dihydroflavonal reductase (<italic>dfr</italic>) gene promoter region revealed the presence of regulatory elements that conferred expression in fruits during ripening.<sup><xref ref-type="bibr" rid="bib124">124</xref></sup> A transcription factor <italic>LcMybA1</italic> that accumulated anthocyanin in litchi pericarp was found to be upregulated by light and ABA.<sup><xref ref-type="bibr" rid="bib125">125</xref></sup> Promoter analysis of the <italic>LcMybA1</italic> gene revealed the presence of light, hormone and abiotic stress-responsive <italic>cis-</italic>elements that were involved in positive and negative regulation of gene expression. A <italic>Gentiana lutea</italic> carotenoid-related zeaxanthin epoxidase (<italic>GIZEP</italic>) gene and promoter region was analyzed for its function in carotenoid biosynthesis.<sup><xref ref-type="bibr" rid="bib126">126</xref></sup> Heterologous expression of a <italic>GIZEP</italic>:<italic>gus</italic> fusion in transgenic tomato specified carotenoid expression in flowers and ripe fruit but minimal levels in vegetative tissues, roots and immature fruit containing chloroplast. <italic>Cis-</italic>elements that are responsive to hormones and abiotic stress factors were identified in the promoter region and may be involved in carotenoid biosynthesis at specific developmental stages. In other studies, two fruit-specific promoters in tomato, SIACS4 and SIEXP1 contained regulatory elements that conferred gene expression specifically in seed, embryo and endosperm tissues.<sup><xref ref-type="bibr" rid="bib127">127</xref></sup> Candidate promoter sequences have also been identified from other fruits that exhibit seed-specific expression in heterologous species, indicating the presence of conserved <italic>cis-</italic>elements.<sup><xref ref-type="bibr" rid="bib128">128</xref></sup></p><p>Fruit-specific promoters have been used to either express or downregulate transgenic proteins at specific stages of development for enhancing fruit yield and quality.<sup><xref ref-type="bibr" rid="bib129 bib130 bib131">129–131</xref></sup> Transgenic tomatoes expressing miraculin, a taste modifying glycoprotein under control of an E8 promoter accumulated uniformly high levels of the transgenic protein in ripening fruits compared to fruits expressing the protein under a 35S promoter, where protein accumulation occurred predominantly in the exocarp.<sup><xref ref-type="bibr" rid="bib132">132</xref></sup> Targeted expression of a yeast <italic>S</italic>-adenosylmethionine decarboxylase gene under the control of a fruit-specific E8 promoter significantly increased spermine and spermidine levels in transgenic tomato fruit, resulting in enhanced shelf life and higher lycopene content.<sup><xref ref-type="bibr" rid="bib133">133</xref></sup> Transgenic tomatoes expressing an <italic>Agrobacterium rolB</italic> gene under control of an ovary-specific promoter produced parthenocarpic fruit.<sup><xref ref-type="bibr" rid="bib134">134</xref></sup> No differences in fruit morphology were observed compared to the non-transformed fruit. In other studies, tomato fruits with enhanced rot resistance and shelf life were obtained by expressing a tomato anionic peroxidase under control of a fruit-specific E8 promoter.<sup><xref ref-type="bibr" rid="bib135">135</xref></sup> Transgenic avocado plants harboring a <italic>S</italic>-adenosine <italic>L</italic>-methionine hydrolyase gene under control of a fruit-specific cellulose promoter have been produced to study the potential for improving fruit shelf life.<sup><xref ref-type="bibr" rid="bib136">136</xref></sup> Targeted expression of a bacterial-derived auxin biosynthesis gene under control of a ovule-specific promoter significantly enhanced fecundity of transgenic ‘Silcora’ and ‘Thompson Seedless’ grapevines by improving berry and cluster size without compromising qualitative characteristics.<sup><xref ref-type="bibr" rid="bib137">137</xref></sup> Similar results of improved yield along with the production of parthenocarpic fruit were obtained in transgenic strawberry and raspberry plants.<sup><xref ref-type="bibr" rid="bib138">138</xref></sup></p></sec><sec><title>Promoters active in the seeds</title><p>The expression of genes that produce seed storage proteins is highly regulated. Deletion analysis of seed-specific promoters has led to identification of proximal regions that confer seed specificity and distal regions that are responsible for modulating gene expression.<sup><xref ref-type="bibr" rid="bib139 bib140 bib141">139–141</xref></sup> Many seed storage protein genes have been cloned from diverse plant species, and their promoters have been analyzed in detail to identify several <italic>cis</italic>- and <italic>trans</italic>-acting elements involved in gene regulation.<sup><xref ref-type="bibr" rid="bib139">139</xref>,<xref ref-type="bibr" rid="bib140">140</xref></sup> Although such proteins exhibit wide structural variations, their promoters have a number of common properties.<sup><xref ref-type="bibr" rid="bib142">142</xref></sup> They allow the synthesis of proteins at high levels in specific tissues of the seed and at certain stages of plant development.<sup><xref ref-type="bibr" rid="bib143">143</xref></sup> The tightly regulated promoters make them ideal candidates for improving seed-specific traits such as nutritional value without potentially altering existing desirable characteristics.<sup><xref ref-type="bibr" rid="bib144">144</xref></sup></p><p>The 2S albumin gene promoter from a number of horticultural species has been used to direct seed-specific gene expression.<sup><xref ref-type="bibr" rid="bib145 bib146 bib147">145–147</xref></sup> DNA sequence analysis of a seed-specific 2S albumin promoter region derived from grape (<italic>Vitis vinifera</italic> L.) indicated that several conserved seed-specific regulatory motifs were clustered within a 0.6 kb region upstream of the transcription start site. A high level of GFP expression was observed in the cotyledonary but not hypocotyl and vegetative tissues of grape and tobacco indicating the ability of the promoter to direct seed-specific gene expression.<sup><xref ref-type="bibr" rid="bib145">145</xref></sup> This promoter region contained DNA motifs with core sequences identical to that of cotyledon box (CATGCA), F1 (ACGT) motif, F2 (CACCTC) motif, F3 (CACGTC) and AGGA box that have been previously characterized in 2S albumin and related seed-specific promoters of other species<sup><xref ref-type="bibr" rid="bib147 bib148 bib149">147–149</xref></sup> Substitution mutation analysis of the napin promoter using promoter–reporter gene fusions in stable transgenic tobacco showed synergistic interactions between the B-box and RY/G <italic>cis</italic>-elements within these complexes. It was further determined that elements in the B-box constitute an ABA-responsive complex and the seed-specific activity of the <italic>napA</italic> gene promoter relies on the combinatorial interaction between the RY/G complex and the B-box ABA-responsive complex during ABA response in seed development.<sup><xref ref-type="bibr" rid="bib150">150</xref></sup> The B-box is highly conserved in all 2S promoters and displays similarity to abscisic acid response elements.<sup><xref ref-type="bibr" rid="bib151">151</xref></sup></p><p>Legumin gene promoters have also been well studied in a number of plant species. In <italic>Pisum sativum</italic>, they are coded for by a multigene family.<sup><xref ref-type="bibr" rid="bib152">152</xref></sup> The promoter regions of <italic>legA</italic>, <italic>legB</italic> and <italic>legC</italic> were analyzed and were found to be identical including the TATA box and CAAT box.<sup><xref ref-type="bibr" rid="bib153">153</xref></sup> Deletion analysis of the pea <italic>legA</italic> major seed storage protein gene identified a minimal 549 bp upstream flanking sequence that was required for seed-specific expression.<sup><xref ref-type="bibr" rid="bib154">154</xref></sup> This fragment contained the <italic>leg</italic> box element, a 28 bp conserved sequence found in the legumin-type genes of <italic>Vicia, Pisum</italic>, <italic>Glycine</italic> and <italic>Helianthus</italic>. Larger promoter fragments significantly increased levels of seed-specific gene expression.<sup><xref ref-type="bibr" rid="bib143">143</xref></sup> DNA binding assays, however, indicated that the <italic>leg</italic> box element is not the sole promoter determinant in legumin gene expression since the −124 bp fragment which included the <italic>leg</italic> box did not bind to nuclear proteins.<sup><xref ref-type="bibr" rid="bib155">155</xref></sup> In addition, deletion of the <italic>leg</italic> box with its seed protein gene-specific CATGCATG motif has no obvious effects on expression levels. A 2.4 kb fragment containing the 5′-flanking region and the 5′-noncoding sequence of the <italic>Vicia faba</italic> legumin gene LeB4 was observed to mediate high level of seed-specific expression in transgenic tobacco plants. Deletion analysis revealed that a 1 kb of 5′-flanking sequence was sufficient for high-levels of expression.<sup><xref ref-type="bibr" rid="bib156">156</xref></sup> Similar to that observed with the pea legA promoter, positive regulatory, enhancer-like <italic>cis</italic>-elements are present within 566 bp of the upstream sequence. However, these elements are only fully functional in conjunction with the core motif CATGCATG of the legumin box present around position −95.<sup><xref ref-type="bibr" rid="bib157">157</xref></sup></p><p>Seed specificity within the 5′-upstream region of a <italic>Vicia faba</italic> non-storage seed protein gene, called <italic>usp</italic> was mainly determined by the −68/+51 region. Deletion analysis of the promoter revealed the 0.4 kb of usp upstream sequence contain at least six distinct interspersed <italic>cis-</italic>elements including an AT-rich sequence, a G-box element and a CATGCATG motif.<sup><xref ref-type="bibr" rid="bib158">158</xref></sup> The beta-phaseolin gene (<italic>phas</italic>), encoding the major seed storage protein of bean (<italic>Phaseolus vulgaris</italic>), is confined to the cotyledons of developing embryos. Promoter analysis revealed that although <italic>cis</italic>-elements extending 1470 bp upstream of the transcription start site can modulate gene expression, the proximal 295 bp is sufficient to drive high levels of seed-specific GUS activity.<sup><xref ref-type="bibr" rid="bib141">141</xref></sup> The <italic>cis-</italic>regulatory CACGTG motif (G box) was identified as a major <italic>cis-</italic>acting regulatory element conferring spatial and temporal control of beta-phaseolin<sup><xref ref-type="bibr" rid="bib159">159</xref></sup> as substitution mutation of this motif reduced promoter activity by 75%.<sup><xref ref-type="bibr" rid="bib160">160</xref></sup> In addition, there are three CANNTG motifs and two AG-1-binding sites in the beta-phaseolin promoter that play a critical role in gene transcription.<sup><xref ref-type="bibr" rid="bib160">160</xref></sup> Combinational interactions between multiple sequence motifs such as two upstream activating sequences UAS1 (−295 to −109) and UAS2 (−468 to −391) affected the spatial and temporal regulation of the promoter. While UAS1 was sufficient for seed-specific expression, UAS2 extended gene activity to the hypocotyl. Deletion of either of the two negative regulatory sequences, NRS1 (−391 to −295) and NRS2 (−518 to −418), resulted in premature onset of GUS expression, indicating their role in the temporal control of gene expression.<sup><xref ref-type="bibr" rid="bib161">161</xref></sup></p><p><italic>Dc3</italic> is a carrot <italic>lea</italic> class gene expressed during embryogenesis in developing seeds and in vegetative tissues subject to drought and treatment with exogenous ABA.<sup><xref ref-type="bibr" rid="bib162">162</xref></sup> The proximal promoter region (−117 to +26) is responsible for mediating the embryo-specific expression.<sup><xref ref-type="bibr" rid="bib163">163</xref></sup> The <italic>Dc3</italic> promoter directed ABA and mannitol-inducible GUS expression in <italic>Arabidopsis</italic> guard cells and the two treatments were additive.<sup><xref ref-type="bibr" rid="bib164">164</xref></sup> A small family of bZIP transcription factors are involved in the seed-specific and ABA-responsive expression of the <italic>Dc3</italic> gene. <italic>Dc3</italic> binds to three DNA binding proteins, DPBF-1, 2 and 3. These DPBFs are bZIP factors that have been postulated to be global regulators of seed-specific and ABA-inducible genes.<sup><xref ref-type="bibr" rid="bib165">165</xref></sup> Deletion analysis of the promoter region led to the delineation of a proximal promoter region and a distal promoter region. The proximal promoter region contains <italic>cis-</italic>acting elements responsible for the developmental regulation of <italic>Dc3</italic> expression in seeds. Both distal promoter region and proximal promoter region interact with common nuclear proteins that are present in embryos and inducible by ABA in vegetative tissues.<sup><xref ref-type="bibr" rid="bib162">162</xref></sup> Following a 3-day water stress cycle, leaf GUS expression increased about 200-fold while there was a 16-fold increase in free ABA. These effects were reversed by re-watering indicating the drought inducibility of this promoter. In addition, 10 µM ABA resulted in more than 10-fold induction within 8 h.<sup><xref ref-type="bibr" rid="bib166">166</xref></sup></p><p>Other <italic>cis</italic>-elements involved in seed-specific promoter expression such as a number of A/T-rich sequences and a CATGCAT/A sequence are present in the 5′-upstream regions of genes encoding concanavalin A (<italic>ConA</italic>) and canavalin, two major seed storage proteins of <italic>Canavalia gladiata</italic>, the sword bean. Deletion analysis of the promoter regions of both genes revealed positive regulatory elements located in the −894/−602 and −602/−74 regions of the <italic>ConA</italic> gene, and in the −428/−376, −281/−155 and −155/−50 regions of the canavalin gene.<sup><xref ref-type="bibr" rid="bib167">167</xref></sup></p><p>Progressive 5′ deletions of the pea lectin (<italic>Psl</italic>) gene promoter identified a 22 bp element (W1), important for seed-specific expression when coupled as a trimer to a heterologous TATA box.<sup><xref ref-type="bibr" rid="bib168">168</xref></sup> Within the 469 bp upstream region of the seed-specific pea lectin gene, a trimer of the 22 bp fragment conferred high gene expression in seeds. This 22 bp fragment contains the binding site for the cloned basic domain/leucine zipper (bZIP) proteins TGA1a and Opaque-2 (O2), which in turn binds to the C-box <italic>cis</italic>-element (ATGAGTCAT).<sup><xref ref-type="bibr" rid="bib169">169</xref></sup> In a majority of the promoters, most of the <italic>cis</italic>-elements are located within 1 kb upstream of the ATG sequence. However, in the HaG3-A sunflower promoter that directs helianthinin gene expression, <italic>cis-</italic>regulatory elements located in a 2.4 kb upstream region were responsible for expression in a heterologous system.<sup><xref ref-type="bibr" rid="bib170">170</xref></sup> Similarly, the 2.4 kb in the 5′ upstream region of the <italic>CuMFT1</italic> (citrus FT/TFL1 homolog from Satsuma mandarin (<italic>Citrus unshiu</italic> Marc.)) contained RY (CATGCAT), E-box (CANNTG) and distant B-box (GCCACTTGTC) <italic>cis-</italic>elements, all of which have been reported to promote seed-specific gene expression in plants. Seed-specific expression was confirmed by expressing the <italic>gus</italic> gene in <italic>Arabidopsis</italic>.<sup><xref ref-type="bibr" rid="bib171">171</xref></sup> A 0.8 kb fragment from the 5′-flanking region of a French bean <italic>beta-phaseolin</italic> gene yielded strong, temporally regulated and embryo-specific GUS expression in transgenic tobacco plants.<sup><xref ref-type="bibr" rid="bib140">140</xref></sup> Expression levels were observed to be similar as that obtained using the phaseolin seed protein promoter.<sup><xref ref-type="bibr" rid="bib172">172</xref></sup></p><p>Several promoters expressed in the seeds can also be expressed in other plant organs. The <italic>strictosidine synthase</italic> (<italic>Str</italic>) gene promoter from <italic>Catharanthus roseus</italic> contains a G-box sequence which helps to direct seed-specific expression independently of other regulatory sequences. G-box-directed expression in leaves, however, required the presence of an enhancer region from the 35S promoter.<sup><xref ref-type="bibr" rid="bib173">173</xref></sup> The fruit and seed-specific expression of two tomato fruit-specific promoters <italic>SIACS4</italic> and <italic>SIEXP1</italic> was analyzed in transgenic tomato lines expressing the promoter: <italic>gus</italic> fusion constructs. The <italic>SIACS4</italic> promoter (−1 to −373) showed GUS activity restricted specifically to flower buds and seeds in fruits. On the contrary, the <italic>SIEXP1</italic> promoter (−1 to −769) showed high level of expression in seeds as compared to fruit tissues at different stages of fruit ripening.<sup><xref ref-type="bibr" rid="bib127">127</xref></sup> The seed-specific expression shown by these promoters might be due to the presence of Prolamin box and E-boxes, which are conserved sequences found in the promoters of many seed storage proteins.<sup><xref ref-type="bibr" rid="bib150">150</xref></sup></p></sec><sec><title>Promoters active in the floral tissues</title><p>In contrast to other plant organs, flowers are composite structures composed of several organs that form an ordered pattern.<sup><xref ref-type="bibr" rid="bib174">174</xref></sup> The typical flower consists of four organs arranged in whorls. The sepals consist of the outermost whorl followed by the petals in the next whorl and stamens (male reproductive organs) in the third whorl and carpels (female reproductive organs) in the innermost whorl.<sup><xref ref-type="bibr" rid="bib175">175</xref></sup> Each of these whorls consist of unique genes targeted to the specific organ and several homeotic genes that affect the fate of organ primordia.<sup><xref ref-type="bibr" rid="bib176">176</xref></sup> Targeted genetic engineering, by utilizing promoters obtained from genes specifically expressed in a specific whorl is highly desirable for targeted gene expression and can be exploited by using specific promoters.<sup><xref ref-type="bibr" rid="bib177">177</xref></sup> Some of the traits that can be engineered in the floral tissues include increased vase life,<sup><xref ref-type="bibr" rid="bib178 bib179 bib180 bib181 bib182">178–182</xref></sup> flower color modification,<sup><xref ref-type="bibr" rid="bib181">181</xref>,<xref ref-type="bibr" rid="bib183 bib184 bib185">183–185</xref></sup> fragrance<sup><xref ref-type="bibr" rid="bib185 bib186 bib187">185–187</xref></sup> and male and female sterility<sup><xref ref-type="bibr" rid="bib188 bib189 bib190 bib191 bib192">188–192</xref></sup> among others.</p><p>Chalcone synthase (<italic>CHS</italic>) is synthesized in the flower corolla, tube and anthers<sup><xref ref-type="bibr" rid="bib193">193</xref></sup> and is important for the biosynthesis of flavonoid antimicrobial phytoalexins and anthocyanin pigments in plants.<sup><xref ref-type="bibr" rid="bib194">194</xref></sup> Various CHS promoters has been studied extensively in many plants, especially in <italic>Phaseolus vulgaris</italic>, antirrhinum, petunia and parsley.<sup><xref ref-type="bibr" rid="bib195 bib196 bib197">195–197</xref></sup> A 1.4 kb promoter fragment of the bean <italic>CHS8</italic> gene was highly active in the root apical meristem and in petals and weakly expressed in other floral organs, mature leaves, and stems.<sup><xref ref-type="bibr" rid="bib198">198</xref></sup> Gene expression strongly depended on the G-box and H-box,<sup><xref ref-type="bibr" rid="bib199">199</xref></sup> as a synthetic 39 bp DNA fragment containing the two elements and linked to the minimal cauliflower mosaic virus 35S promoter conferred a high level of tissue-specific expression. Mutations in either the G-box or H-box motifs abolished tissue-specific gene expression.<sup><xref ref-type="bibr" rid="bib195">195</xref></sup> A mutation in the G-box did not exhibit impaired promoter response to wounding, but demonstrated a 19% reduction in the response to HgCl<sub>2</sub> and TMV. A mutation at the H-box resulted in a 30% increase in promoter response to wounding and reductions of 36% and 54% in the response to HgCl<sub>2</sub> and TMV, respectively, demonstrating the differential utilization of regulatory <italic>cis-</italic>elements.<sup><xref ref-type="bibr" rid="bib200">200</xref></sup> A silencer element present between positions −140 and −326 contained three binding sites for a bean nuclear factor (SBF-1).<sup><xref ref-type="bibr" rid="bib201">201</xref></sup> The region between −326 and −130 contained both activator and silencer elements.<sup><xref ref-type="bibr" rid="bib202">202</xref></sup> The petunia genome contains eight chalcone synthase genes, of which four are differentially expressed in floral tissues and UV light-induced seedlings.<sup><xref ref-type="bibr" rid="bib197">197</xref></sup> The <italic>chs</italic>A promoter contains a 220 bp <italic>cis-</italic>acting element region conferring flower-specific and UV-inducible expression <sup><xref ref-type="bibr" rid="bib203">203</xref></sup> and its expression was enhanced when plant tissues were exposed to high carbohydrate levels.<sup><xref ref-type="bibr" rid="bib204">204</xref></sup> A promoter fragment from −67 to +1, was able to direct low level flower-specific gene expression, but could not drive UV-inducible expression in transgenic <italic>Petunia</italic> seedlings.<sup><xref ref-type="bibr" rid="bib205">205</xref></sup> Histochemical analyses of GUS expression demonstrated that <italic>CHS</italic> promoters are not only active in pigmented cell types (epidermal cells of the flower corolla and tube and subepidermal cells of the flower stem), but also in a number of non-pigmented cell types (mesophylic cells of the corolla, several cell types in the ovary and the seed coat).<sup><xref ref-type="bibr" rid="bib197">197</xref></sup> The highest level of expression directed by the 1.1 kb snapdragon chalcone synthase promoter was observed in immature seeds. Deletions analysis identified regions of the promoter required for expression in roots, stems, leaves, seeds and flower petals of transgenic plants. A promoter fragment truncated to −39 activates transcription in roots of 4-week-old seedlings, whereas a fragment extending to −197 bp directed expression in petals and seeds.<sup><xref ref-type="bibr" rid="bib206">206</xref>,<xref ref-type="bibr" rid="bib207">207</xref></sup> The positive regulatory element in the promoter consists of a 47 bp direct repeat between positions −564 and −670.<sup><xref ref-type="bibr" rid="bib208">208</xref></sup> 150 bp of the 5′ flanking region contained <italic>cis-</italic>acting signals for UV light-induced expression.<sup><xref ref-type="bibr" rid="bib209">209</xref></sup> The <italic>GTCHS1</italic> promoter from <italic>Gentiana triflora</italic> contains a sequence of the MYB protein-binding site, five consensus sequences of the MYC protein-binding site, one core sequence of a G-box and three P-box-like sequences. Gene expression is strongly directed flower limbs and the inner epidermis<sup><xref ref-type="bibr" rid="bib210">210</xref></sup> and is dependent on the G-box.<sup><xref ref-type="bibr" rid="bib211">211</xref></sup></p><p>In efforts to produce high transgene expression in petal tissue of ray florets of chrysanthemum, expression levels were compared with four petal-specific promoters: ubiquitin extension protein (<italic>UEP</italic>1) promoter from chrysanthemum chalcone synthase (<italic>chs-A</italic>) a zinc finger transcription factor (<italic>EPF</italic>2-5) from petunia, eceriferum (<italic>CER</italic>6) from <italic>Arabidopsis</italic> and multicystatin (<italic>PMC</italic>) from potato. The highest expression in petal tissue of ray and disc florets was conferred by the <italic>UEP</italic>1 promoter, followed by <italic>CER</italic>6 and <italic>EPF</italic>2-5. The <italic>UEP</italic>1 promoter in ray florets was reported to confer over 50-fold enhancement in expression as compared to CaMV 35S-based promoters.<sup><xref ref-type="bibr" rid="bib212">212</xref></sup></p><p>Promoters targeting other parts of the flower have also been evaluated. When a 2.4 kb fragment of the pistil-specific thaumatin/PR5-like protein (<italic>PsTL1</italic>) promoter from Japanese pear (<italic>Pyrus serotina</italic>) was evaluated,<sup><xref ref-type="bibr" rid="bib213">213</xref></sup> it was observed that <italic>PsTL1</italic> accumulated in pistils but not in other floral and vegetative organs which constitute a novel pistil-specific class of thaumatin/PR5-like protein.<sup><xref ref-type="bibr" rid="bib214">214</xref>,<xref ref-type="bibr" rid="bib215">215</xref></sup> Other parts of the flower targeted include the flower receptacle. Promoters targeting other parts of the flower have been evaluated. When a 2.4 kb fragment of the pistil-specific thaumatin/PR5-like protein (<italic>PsTL1</italic>) promoter from Japanese pear (<italic>Pyrus serotina</italic>) was evaluated,<sup><xref ref-type="bibr" rid="bib213">213</xref></sup> it was observed that PsTL1 accumulated in pistils, but not in other floral and vegetative organs which constitute a novel pistil-specific class of thaumatin/PR5-like protein.<sup><xref ref-type="bibr" rid="bib214">214</xref>,<xref ref-type="bibr" rid="bib215">215</xref></sup> Several reports exist on the isolation, characterization and use of promoters targeted to the flower receptacles,<sup><xref ref-type="bibr" rid="bib216">216</xref>,<xref ref-type="bibr" rid="bib217">217</xref></sup> stamen,<sup><xref ref-type="bibr" rid="bib218 bib219 bib220">218–220</xref></sup> anthers<sup><xref ref-type="bibr" rid="bib221 bib222 bib223">221–223</xref></sup> and ovaries.<sup><xref ref-type="bibr" rid="bib134">134</xref></sup> The potato <italic>SK2</italic> gene promoter directed pistil-specific gene expression. It was observed that the regulatory elements responsible for pistil-specific expression were located within a 230 bp fragment.<sup><xref ref-type="bibr" rid="bib224">224</xref></sup></p><p>Numerous genes and their promoters that are expressed at the various stages during male gametogenesis have been cloned.<sup><xref ref-type="bibr" rid="bib225">225</xref></sup> Most of these have been isolated from agronomic crops such as maize,<sup><xref ref-type="bibr" rid="bib226">226</xref>,<xref ref-type="bibr" rid="bib227">227</xref></sup> rice,<sup><xref ref-type="bibr" rid="bib228">228</xref>,<xref ref-type="bibr" rid="bib229">229</xref></sup> tobacco<sup><xref ref-type="bibr" rid="bib230">230</xref>,<xref ref-type="bibr" rid="bib231">231</xref></sup> and wheat<sup><xref ref-type="bibr" rid="bib232">232</xref></sup> as well as the model plant <italic>Arabidopsis</italic>.<sup><xref ref-type="bibr" rid="bib233 bib234 bib235 bib236">233–236</xref></sup> A few have also been isolated from horticultural crops.<sup><xref ref-type="bibr" rid="bib237">237</xref></sup> These promoters fused to a cytotoxic gene have been used to induce male sterility.<sup><xref ref-type="bibr" rid="bib226">226</xref>,<xref ref-type="bibr" rid="bib238">238</xref></sup> The <italic>LAT52</italic> and <italic>LAT59</italic> anther-specific gene promoters from tomato have been evaluated in various crops for their anther-specific activity.<sup><xref ref-type="bibr" rid="bib237">237</xref>,<xref ref-type="bibr" rid="bib239">239</xref>,<xref ref-type="bibr" rid="bib240">240</xref></sup> These genes are very critical during tomato pollen development. In their absence, pollen germinates abnormally and is sterile.<sup><xref ref-type="bibr" rid="bib241">241</xref></sup> All major <italic>cis-</italic>regulatory elements required for pollen-specific transcription in the LAT52 promoter were located within −492 to −52.<sup><xref ref-type="bibr" rid="bib242">242</xref></sup> Both promoters became active with the onset of microspore mitosis and increased progressively until anthesis,<sup><xref ref-type="bibr" rid="bib223">223</xref></sup> although the LAT52 promoter demonstrated a minor temporal difference in activity when tested in different plant species.<sup><xref ref-type="bibr" rid="bib243 bib244 bib245">243–245</xref></sup> The LAT52 promoter was highly active in electroporated pollen protoplasts isolated from <italic>Lilium longiflorum</italic>.<sup><xref ref-type="bibr" rid="bib246">246</xref></sup> The antisense <italic>Bcp1</italic> gene under the control of the LAT52 promoter induced sterility in cauliflower pollen.<sup><xref ref-type="bibr" rid="bib247">247</xref></sup> Similarly, a 0.44 kb chiA PA2 promoter fragment from petunia drove pollen-specific gene expression and a 1.75 kb chiB PB promoter fragment conferred anther-specific (pollen and tapetum cells) expression to the <italic>gus</italic> gene.<sup><xref ref-type="bibr" rid="bib222">222</xref></sup> The <italic>TomA108</italic> gene promoter from tomato was also highly active from early-meiosis to free microspores production in the tapetum.<sup><xref ref-type="bibr" rid="bib248">248</xref></sup> Deletion analysis of the <italic>BAN215</italic>-6 gene promoter isolated from the Chinese cabbage identified a 383 bp (−274–+109) region that was observed to be sufficient for the anther-specific expression of the <italic>gus</italic> gene. GUS expression was first detected in uninucleate microspores, increased during anther development and reached its highest level in mature pollens.<sup><xref ref-type="bibr" rid="bib249">249</xref></sup> Similar observation were made with the 2.7 kb promoter fragment of a pea <italic>END1</italic> gene. This promoter was evaluated in several species and observed to be fully functional in the anthers.<sup><xref ref-type="bibr" rid="bib250">250</xref></sup></p></sec><sec><title>Promoters active in the root system</title><p>Plant roots have been essential for the evolution of vascular plants enabling them to meet the requirements for anchorage and the acquisition of water and nutrients.<sup><xref ref-type="bibr" rid="bib251">251</xref></sup> Roots are multifunctional and involved in the acquisition of water and nutrients, anchorage of the plant and storage functions.<sup><xref ref-type="bibr" rid="bib252">252</xref>,<xref ref-type="bibr" rid="bib253">253</xref></sup> In fact, plant productivity is dependent on a heathy root system<sup><xref ref-type="bibr" rid="bib254">254</xref></sup> as problem with root health directly affects the above ground part.<sup><xref ref-type="bibr" rid="bib255">255</xref>,<xref ref-type="bibr" rid="bib256">256</xref></sup> Roots interact with its surrounding environment <sup><xref ref-type="bibr" rid="bib257">257</xref></sup> and can be susceptible to a multitude of problems stemming from the environment in which it lives.<sup><xref ref-type="bibr" rid="bib252">252</xref>,<xref ref-type="bibr" rid="bib258 bib259 bib260">258–260</xref></sup> Targeted gene expression by using root-specific promoters can allow for the development of horticultural plants better suited for growth in a range of soil types, soil pH and under microbial stress.<sup><xref ref-type="bibr" rid="bib261">261</xref>,<xref ref-type="bibr" rid="bib262">262</xref></sup> Several root-specific promoters have been evaluated in horticultural plants. The <italic>SLREO</italic> gene isolated from tomato is highly expressed in roots, but had a very low level of expression in aerial plant organs.<sup><xref ref-type="bibr" rid="bib263">263</xref></sup> The RB7 protein from tobacco,<sup><xref ref-type="bibr" rid="bib262">262</xref></sup> is a membrane channel aquaporin, allowing the diffusion of amino acids and/or peptides from the vacuolar compartment to the cytoplasm.<sup><xref ref-type="bibr" rid="bib264">264</xref>,<xref ref-type="bibr" rid="bib265">265</xref></sup> This promoter is root-specific and has been used to drive the <italic>Arabidopsis</italic> thionin (<italic>Thi2.1</italic>) gene in tomato.<sup><xref ref-type="bibr" rid="bib266">266</xref></sup> A strawberry homolog (<italic>FaRB7</italic>) behaves in the same way as the tobacco RB7 promoter.<sup><xref ref-type="bibr" rid="bib267">267</xref></sup> Other promoters identified include a 2 kb promoter fragment of the <italic>MipB</italic> gene from <italic>Mesembryanthemum crystallinum</italic> that was observed to be expressed strongly in the tobacco root. However, gene expression was also observed in other rapidly expanding cells and cells with high water flux capacity.<sup><xref ref-type="bibr" rid="bib268">268</xref></sup></p><p>Several root nodule-specific promoters have been identified from leguminous plants.<sup><xref ref-type="bibr" rid="bib269">269</xref></sup> A 1.3 kb fragment of the French bean <italic>gln-gamma</italic> gene promoter is strongly induced during nodule development.<sup><xref ref-type="bibr" rid="bib270">270</xref></sup> The <italic>Vicia faba VfLb29</italic> gene promoter was found to be specifically active not only in the infected cells of the nitrogen-fixing zone of root nodules but also in arbuscule-containing cells of transgenic <italic>V. faba</italic> roots colonized by the endomycorrhizal fungus <italic>Glomus intraradices</italic>.<sup><xref ref-type="bibr" rid="bib271">271</xref></sup> A promoter fragment (−692/41) encoding the <italic>Vicia faba</italic> early nodulin <italic>VfEnod12</italic> and containing a putative binding site for the transcription factor ENBP1, mediated reporter gene expression in root cortical cells, nodule primordia and the prefixing zone II of transgenic <italic>Vicia hirsute</italic> root nodules.<sup><xref ref-type="bibr" rid="bib272">272</xref></sup> A 1.9 kb fragment of the <italic>Sesbania rostrata</italic> leghemoglobin <italic>glb3</italic> 5′-upstream region was found to direct a high level of nodule-specific GUS activity in lotus. Replacement of the −161 to −48 region, containing the <italic>glb3</italic> CAAT and TATA boxes, with the heterologous truncated promoters delta-p35S and delta-pnos, resulted in a loss of nodule specificity and reduction of GUS activity restricted to the <italic>Rhizobium</italic>-infected cells of the nodules.<sup><xref ref-type="bibr" rid="bib273">273</xref></sup> Promoter analyses of pea <italic>PsENOD12A</italic> and <italic>PsENOD12B</italic>, nodulin gene promoters showed that the 200 bp immediately upstream of the transcription start are sufficient to direct nodule-specific and Nod factor-induced gene expression.<sup><xref ref-type="bibr" rid="bib274">274</xref></sup> GUS activity was only detected in the infected cells of the nodules of lotus transgenic plants when a Npv30 promoter isolated from <italic>Phaseolus vulgaris</italic> fused to the <italic>gus</italic> reporter gene was used.<sup><xref ref-type="bibr" rid="bib275">275</xref></sup></p><p>Several genes are highly upregulated in tubers.<sup><xref ref-type="bibr" rid="bib276 bib277 bib278">276–278</xref></sup> Many of these storage gene promoters have been exploited for horticultural crop improvement. Patatin is a major tuber protein and is very tissue-specific.<sup><xref ref-type="bibr" rid="bib279">279</xref></sup> The 1.5 kb 5′-upstream region of the class I patatin gene <italic>B33</italic> directed strong expression of the GUS reporter gene in potato tubers which was on average 100- to 1000-fold higher in tubers as compared to leaf, stem and roots.<sup><xref ref-type="bibr" rid="bib280">280</xref></sup> Gene expression was also induced by sucrose application.<sup><xref ref-type="bibr" rid="bib278">278</xref></sup> Deletion analysis identified a tuber-specific element located downstream from position −195. Sequences between −40 and −400 bp and between −400 and −957 bp of the transcriptional start site were able to confer tuber-specific expression on a heterologous truncated promoter.<sup><xref ref-type="bibr" rid="bib281">281</xref></sup> Sucrose inducibility was controlled by sequences downstream of position −228. <sup><xref ref-type="bibr" rid="bib282">282</xref>,<xref ref-type="bibr" rid="bib283">283</xref></sup> High levels of mature human serum albumin was expressed in potato tubers using the potato patatin B33 tuber-specific promoter.<sup><xref ref-type="bibr" rid="bib284">284</xref></sup></p><p>Sporamin accounts for more than 80% of the total soluble proteins of tuberous roots of sweet potato<sup><xref ref-type="bibr" rid="bib285">285</xref></sup> and can be induced by wounding and sucrose.<sup><xref ref-type="bibr" rid="bib286">286</xref></sup> Two wound response-like elements, a G box-like element and a GCC core-like sequence, were found in the sporamin gene promoter.<sup><xref ref-type="bibr" rid="bib287">287</xref></sup> When overexpressed in potato, the sporamin promoter was highly active in leaves, stems and different size tubers.<sup><xref ref-type="bibr" rid="bib288">288</xref></sup> Deletion of the sporamin A promoter sequences extending from position −305 (relative to the transcription start site) to −283 and from −146 to −87 resulted in an approximately 40-fold enhancement in GUS reporter expression. It was observed that the sequence between positions −282 and −165 contained to two <italic>cis-</italic>acting elements, termed CMSREs (carbohydrate metabolite signal responsive elements) 1 and 2 are responsible for the sucrose-responsiveness of the promoter.<sup><xref ref-type="bibr" rid="bib289">289</xref></sup></p></sec><sec><title>Promoters active in the vascular tissues</title><p>The plant's vascular system acts as a bridge between the leaves and other parts of the shoot, with the roots.<sup><xref ref-type="bibr" rid="bib290">290</xref></sup> This system, comprised of two kinds of conducting tissue, the xylem and phloem enables efficient long-distance transport between the organs.<sup><xref ref-type="bibr" rid="bib291">291</xref></sup> Xylem is primarily responsible for water transport and movement of soluble mineral nutrients from the roots throughout the plant.<sup><xref ref-type="bibr" rid="bib292">292</xref></sup> Phloem, on the other hand, transports sugars from source tissues such as the photosynthetic leaf cells to sink or storage tissues such as the roots, flowers or fruits.<sup><xref ref-type="bibr" rid="bib293">293</xref>,<xref ref-type="bibr" rid="bib294">294</xref></sup> Targeting a transgene into the vasculature using either a xylem or phloem-specific promoter allows gene expression at the site of infection and can potentially control vascular maladies. It can also provide a rapid response in response to wounding and for the control of aphids and other sap sucking insects.<sup><xref ref-type="bibr" rid="bib295 bib296 bib297">295–297</xref></sup></p><p>A 494 bp promoter fragment of the glycine-rich wall protein GRP 1.8 from the French bean translationally fused to the <italic>gus</italic> gene expressed the gene in vascular tissue of roots, stems, leaves and flowers. Four <italic>cis-</italic>acting regulatory regions, SE1 and SE2 (stem elements), a negative regulatory element and a root-specific element, were found to control the tissue-specific expression.<sup><xref ref-type="bibr" rid="bib298">298</xref></sup> The vs-1 motif in the GRP 1.8 promoter was a <italic>cis-</italic>element that specifically bound to a transcription activation factor VSF-1 protein and allows xylem-specific expression.<sup><xref ref-type="bibr" rid="bib299">299</xref></sup> The gene was developmentally expressed during differentiation of both primary and secondary vascular tissue and was also rapidly induced (within <30 min) after excision-wounding of young stems.<sup><xref ref-type="bibr" rid="bib300">300</xref></sup> The bean phenylalanine ammonia-lyase gene 2 (<italic>PAL2</italic>) is expressed in the early stages of vascular development at the inception of xylem differentiation. Deletion analysis revealed the presence of <italic>cis</italic>-elements located between nucleotides −289 and −74 relative to the transcription start site being essential for xylem expression.<sup><xref ref-type="bibr" rid="bib301">301</xref></sup> Expression of the PAL2 promoter in the vascular system involves positive and negative regulatory <italic>cis</italic>-elements. Among these elements is an AC-rich motif implicated in xylem expression.<sup><xref ref-type="bibr" rid="bib302">302</xref></sup> Similarly, the citrus PAL gene (<italic>CsPP</italic>) promoter fused to the <italic>gus</italic> gene and transformed into tobacco and ‘Valencia’ sweet orange preferentially, but not exclusively, conferred gene expression in xylem tissues of tobacco. Weaker GUS staining was also detected throughout the petiole region in tobacco and citrus CsPP transgenic plants.<sup><xref ref-type="bibr" rid="bib303">303</xref></sup> The <italic>Arabidopsis</italic> PAL promoter when transformed into citrus expressed exclusively in the xylem parenchyma.<sup><xref ref-type="bibr" rid="bib304">304</xref></sup></p><p>The full-length promoter and a series of 5′ deletions of the pea cytosolic glutamine synthetase <italic>GS3A</italic> gene were fused to the <italic>gus</italic> gene and introduced into tobacco and alfalfa. The <italic>GS3A</italic> promoter directed GUS expression in the phloem cells of the vasculature in leaves, stems and roots. Interestingly, the promoter was found to be active even when deleted to −132 relative to the start of transcription.<sup><xref ref-type="bibr" rid="bib305">305</xref></sup> The <italic>Arabidopsis</italic> sucrose-H<sup>+</sup> symporter <italic>AtSUC2</italic><sup><xref ref-type="bibr" rid="bib306">306</xref></sup> has been used to direct phloem-specific gene expression in a number of horticultural crops, such as Mexican lime,<sup><xref ref-type="bibr" rid="bib307">307</xref></sup> sweet orange,<sup><xref ref-type="bibr" rid="bib308">308</xref></sup> pears<sup><xref ref-type="bibr" rid="bib309">309</xref></sup> and strawberries.<sup><xref ref-type="bibr" rid="bib310">310</xref></sup> Two alleles of the <italic>Citrus sinensis</italic> sucrose synthase-1 promoter (<italic>CsSUS1p</italic>) were inserted upstream of the <italic>gus</italic> gene to test their ability to drive expression in the phloem of transgenic <italic>A. thaliana</italic> and <italic>N. tabacum</italic>. Although both promoter variants were capable of conferring localized GUS expression in the phloem, the <italic>CsSUS1p-2</italic> allele also generated a significant level of expression in non-target tissues. Deletion analysis of the CsSUS1p suggested that a fragment comprising nucleotides −410 to −268 relative to the transcriptional start site contained elements required for phloem-specific expression, while nucleotides −268 to −103 contained elements necessary for wound-specific expression.<sup><xref ref-type="bibr" rid="bib311">311</xref></sup> In citrus, the CsSUS promoter appeared leaky with some laminar tissue staining.<sup><xref ref-type="bibr" rid="bib312">312</xref></sup> A citrus phloem protein 2 (CsPP2) promoter was also evaluated in sweet orange and gene was observed to be preferentially expressed in the phloem.<sup><xref ref-type="bibr" rid="bib308">308</xref></sup> Two heterologous promoters, rolC and CoYMVP, were fused with the <italic>gus</italic> reporter gene and evaluated in the vegetative tissues of apple. It was observed that the CoYMV promoter was slightly more active than the rolC promoter, although both expressed GUS at a lower level than the CaMV 35S promoter. This analysis demonstrated that with both the rolC and CoYMV promoters the reporter gene activity was primarily localized to vascular tissues, particularly the phloem.<sup><xref ref-type="bibr" rid="bib313">313</xref></sup></p></sec><sec><title>Inducible promoters</title><p>These promoters are induced by either physical factors such as biotic and abiotic factors or chemical agents and is a powerful tool to regulate the expression of genes at certain stages of plant or tissue development.<sup><xref ref-type="bibr" rid="bib314 bib315 bib316 bib317 bib318">314–318</xref></sup> Examples of physically regulated promoters include heat shock promoters,<sup><xref ref-type="bibr" rid="bib319">319</xref></sup> cold inducible promoters,<sup><xref ref-type="bibr" rid="bib320">320</xref></sup> light inducible promoters,<sup><xref ref-type="bibr" rid="bib321">321</xref></sup> light repressible promoters<sup><xref ref-type="bibr" rid="bib322">322</xref></sup> or wound inducible promoters.<sup><xref ref-type="bibr" rid="bib323">323</xref></sup> Chemically inducible promoters include alcohol regulated promoters,<sup><xref ref-type="bibr" rid="bib324">324</xref></sup> tetracycline regulated promoters,<sup><xref ref-type="bibr" rid="bib325">325</xref></sup> steroid responsive promoters such as glucocorticoid receptor promoters, estrogen and ecdysone receptor promoters,<sup><xref ref-type="bibr" rid="bib316">316</xref>,<xref ref-type="bibr" rid="bib326">326</xref></sup> metal-responsive promoters<sup><xref ref-type="bibr" rid="bib327">327</xref></sup> and pathogenesis related promoters.<sup><xref ref-type="bibr" rid="bib328">328</xref></sup> Some of these promoters have been isolated from horticultural crops or used for horticultural plant improvement.</p><p>The potato proteinase inhibitor II gene (<italic>pinII</italic>) is a chymotrypsin and trypsin inhibitor<sup><xref ref-type="bibr" rid="bib329">329</xref></sup> and is wound and UV irradiation inducible.<sup><xref ref-type="bibr" rid="bib330">330</xref>,<xref ref-type="bibr" rid="bib331">331</xref></sup> The sequence TATAAA is found 26 nucleotides upstream of the transcription initiation site and the sequence CAAAT at position –103 in the promoter.<sup><xref ref-type="bibr" rid="bib332">332</xref></sup> The wound inducibility of this promoter has been evaluated in several plant species to test gene function that involve cell-specific and systemic induction.<sup><xref ref-type="bibr" rid="bib333">333</xref></sup> The PinII promoter has been utilized in the wound-inducible expression of the bacterial isopentenyl transferase (<italic>ipt</italic>) gene into <italic>Nicotiana plumbaginifolia</italic>.<sup><xref ref-type="bibr" rid="bib334">334</xref></sup> In transgenic rice plants, the expression of the pinII-<italic>gus</italic> fusion gene displayed a systemic wound response.<sup><xref ref-type="bibr" rid="bib335">335</xref></sup> In alfalfa, GUS expression was observed in leaf and root vascular tissue, and in some plants, expression was observed in leaf mesophyll cells. Mechanical wounding of leaves increased GUS expression approximately twofold over 24 h.<sup><xref ref-type="bibr" rid="bib336">336</xref></sup> The PinII promoter is active in monocot species also. Localized induced gene expression was obtained in white spruce seedlings (<italic>Picea glauca</italic>) using a similar pinII-<italic>gus</italic> construct.<sup><xref ref-type="bibr" rid="bib337">337</xref></sup> In rice, the wound-inducible expression of the <italic>pinII</italic> gene driven by its own promoter, together with the first intron of the rice actin 1 gene (<italic>act1</italic>), resulted in high-level accumulation of the PINII protein in the transgenic plants.<sup><xref ref-type="bibr" rid="bib338">338</xref></sup> The <italic>wun1</italic> gene is another wound inducible gene from potato.<sup><xref ref-type="bibr" rid="bib339">339</xref></sup> Histochemical analysis of transgenic tobacco plants that expressing the wun1–<italic>gus</italic> fusions demonstrated the wound-inducible and cell-specific wun1 promoter activity in plants containing the −1022 bp fragment.<sup><xref ref-type="bibr" rid="bib340">340</xref></sup></p><p>The tomato <italic>Lehsp23.8</italic> heat shock protein gene's expression is induced by treatment with high or low temperatures, heavy metal or ABA. Using the <italic>gus</italic> reporter gene system, the developmental and tissue-specific expression of the <italic>gus</italic> gene controlled by the <italic>Lehsp23.8</italic> promoter was characterized in transgenic tomato plants. The optimal heat-shock temperatures leading to the maximal GUS activity in the pericarp of green, breaker, pink and red fruits were 42, 36, 39 and 39 °C, respectively.<sup><xref ref-type="bibr" rid="bib341">341</xref></sup> Deletion analysis of the <italic>Lehsp23.8</italic> promoter revealed a proximal region (−565 to −23 bp) to harbor <italic>cis-</italic>regulatory elements that conferred high levels of heat-induced expression in transgenic tobacco. Mutation of the five proximal HSEs (HSE1 to 5) led to an absence of heat inducibility.<sup><xref ref-type="bibr" rid="bib342">342</xref></sup> The tomato chloroplast small heat shock protein (HSP), <italic>HSP21</italic>, is also induced by heat treatment in leaves.<sup><xref ref-type="bibr" rid="bib343">343</xref></sup> Several sunflower genes encode small HSPs.<sup><xref ref-type="bibr" rid="bib344">344</xref>,<xref ref-type="bibr" rid="bib345">345</xref></sup> In vegetative tissues, these mRNAs accumulated in response to either heat shock (42 °C), ABA or mild water stress treatments. The <italic>Hahsp17</italic>.7<italic>G4</italic> mRNA is also active during zygotic embryogenesis at 25 °C. Developmental induction of the G4 promoter was faithfully reproduced during zygotic embryogenesis in transgenic plants containing G4: GUS translational fusions. Distal sequences of this promoter (between −1132 and −395) were needed to confer a preferential spatial expression of GUS activity in the cotyledons while proximal regions confer responses to ABA and heat shock<sup><xref ref-type="bibr" rid="bib346">346</xref></sup> This −83 to +163 fragment was observed to be sufficient to support a promoter activity in tobacco galls induced by the root–knot nematode <italic>Meloidogyne incognita</italic>. GUS activity was largely restricted to giant cells within the galls.<sup><xref ref-type="bibr" rid="bib347">347</xref></sup> However, the <italic>Hahsp17.6G1</italic> (<italic>G1</italic>) promoter which is not induced by heat shock, was observed to be silent in these giant cells, indicating that the high metabolic rate of giant cells produced as a result of nematode infection may somehow mimic heat-shock and/or other stress responses.<sup><xref ref-type="bibr" rid="bib348">348</xref></sup> Other examples include the strong oxidative stress-inducible peroxidase <italic>SWPA2</italic> promoter from sweet potato. This promoter contained several <italic>cis-</italic>element sequences implicated in oxidative stress such as GCN-4, AP-1, HSTF and SP-1 reported in animal cells and a plant-specific G-box. A 1314 bp promoter fragment fused to the <italic>gus</italic> gene and transformed into tobacco exhibited about 30 times higher GUS expression than the CaMV 35S promoter in response to environmental stresses including hydrogen peroxide, wounding and UV treatment.<sup><xref ref-type="bibr" rid="bib349">349</xref></sup> similarly, when potatoes were transformed with a stress inducible <italic>Arabidopsis</italic> rd29A promoter driving the cold tolerance CBF genes, freezing tolerance was increased by 2 °C.<sup><xref ref-type="bibr" rid="bib350">350</xref></sup></p><p>Several promoters are chemically induced. Ethylene treatment or leaves wounding rapidly induced the melon <italic>ACC oxidase</italic> gene, CM-ACO1-<italic>gus</italic> gene in transgenic tobacco plants.<sup><xref ref-type="bibr" rid="bib351">351</xref></sup> Jasmonates and alpha-linolenic acid strongly induced the expression of the wound-induced 4CL promoter in parsley cell cultures and transgenic tobacco plants expressing 4CL1–<italic>GUS</italic> gene fusions. This supported a role for jasmonates in mediating wound-induced gene expression.<sup><xref ref-type="bibr" rid="bib352">352</xref></sup> Two wound response-like elements, a G box-like element and a GCC core-like sequence were found within a 1.25 kb <italic>sporamin</italic> promoter. Transgenic tobacco containing this promoter driving the <italic>gus</italic> gene was wounded and a high level of GUS activity was observed in stems and leaves of, but not in roots. Exogenous application of methyl jasmonate also activated the sporamin promoter in leaves and stems of sweet potato.<sup><xref ref-type="bibr" rid="bib287">287</xref></sup> The chemically inducible PR-1a tobacco promoter was fused to the <italic>Bacillus thuringiensis cry1Ab</italic> gene and transformed into broccoli. Two progeny lines expressed the <italic>cry1Ab</italic> gene and provided insect resistance when treated with the chemical inducers 2,6-dichloroiso-nicotinic acid or 1,2,3-benzothiadiazole-7-carbothioic acid <italic>S</italic>-methyl ester.<sup><xref ref-type="bibr" rid="bib353">353</xref></sup> Other examples include the alfalfa pathogen-inducible PR 10 promoter. This promoter fused to the <italic>Vitis</italic> stilbene synthase 1 (<italic>VvSS1</italic>) gene was introduced into the grape rootstock genome. Transgenic plants accumulated 5- to 100-fold resveratrol in leaves infected with <italic>Botrytis</italic> using an <italic>in vitro</italic> test.<sup><xref ref-type="bibr" rid="bib354">354</xref></sup></p><p>Some promoters can be regulated both physically and chemically. A 2.2 kb promoter region of the tomato prosystemin gene fused to the <italic>gus</italic> gene and transformed back into tomato contains elements conferring its correct temporal and spatial expression in the vascular bundles of transgenic tomato plants by wounding and by treatment of the plants with methyl jasmonate.<sup><xref ref-type="bibr" rid="bib323">323</xref></sup></p></sec><sec sec-type="conclusions"><title>Conclusions</title><p>The global human population is increasing at an unprecedented rate and is projected to cross 11 billion before the end of this century.<sup><xref ref-type="bibr" rid="bib355">355</xref></sup> This doubling of the population and a rapid increase in global food demand creates huge challenges for the sustainability both of food production and the ability to grow more from a shrinking cultivable land mass. Thus far, the combined effects of improved varieties, increased fertilizer use and irrigation coupled with increased pesticide use have been instrumental in allowing world food production to double in the last 35 years<sup><xref ref-type="bibr" rid="bib356">356</xref></sup> A multifaceted and linked global strategy to increase food production from shrinking land and water resources will ensure sustainable and equitable food security.<sup><xref ref-type="bibr" rid="bib357">357</xref>,<xref ref-type="bibr" rid="bib358">358</xref></sup> Fruits and vegetables claimed an increasing share of the world agricultural trade, from 10.6% in 1961 to 17% in 2001.<sup><xref ref-type="bibr" rid="bib359">359</xref></sup> It is expected that demand for horticultural commodities, especially fruits, vegetables and flowers will continue to increase with the increase in the purchasing ability of the expanding middle class and an growing awareness of the many health benefits associated with an increased consumption of fruits and vegetables.<sup><xref ref-type="bibr" rid="bib360">360</xref>,<xref ref-type="bibr" rid="bib361">361</xref></sup></p><p>Acreage under genetically modified crop plants has increased substantially in recent years as more and more acreage is consumed to feed, clothe and sustain a growing world population.<sup><xref ref-type="bibr" rid="bib362">362</xref></sup> However, there has been limited progress in the commercialization of genetically modified horticultural commodities, with the exception of the Hawaiian papaya cultivars resistant to papaya ringspot virus <sup><xref ref-type="bibr" rid="bib363">363</xref>,<xref ref-type="bibr" rid="bib364">364</xref></sup> and color-altered varieties of carnation flowers.<sup><xref ref-type="bibr" rid="bib365">365</xref></sup> Development of genetically modified horticultural cultivars that can alleviate consumer concerns and the related reluctance of food processors and marketers to accept new biotech horticultural commodities can speed up the introduction of horticultural products already developed.<sup><xref ref-type="bibr" rid="bib366">366</xref></sup></p><p>In recent years, molecular advancement in the field of bioinformatics has been rapid.<sup><xref ref-type="bibr" rid="bib37">37</xref></sup> With the genome of a number of horticultural species being sequenced and the availability of numerous online databases for analyzing, identifying and characterizing promoters from different horticultural species,<sup><xref ref-type="bibr" rid="bib367 bib368 bib369 bib370 bib371">367–371</xref></sup> it has become relatively easier to identify and characterize plant derived promoters and other genetic elements. Identification and incorporation of plant promoter and other genetic sequences by exploiting the expanding public databases and bioinformatics services can potentially alleviate some of the public concerns about safety issues with the use of a genetically modified horticultural crop.<sup><xref ref-type="bibr" rid="bib372">372</xref>,<xref ref-type="bibr" rid="bib373">373</xref></sup> Development of precision breeding techniques (previously termed as cisgenic or intragenic genetic improvement)<sup><xref ref-type="bibr" rid="bib374">374</xref></sup> will enable more precise genetic modification of plants.<sup><xref ref-type="bibr" rid="bib375">375</xref></sup> The resulting horticultural plant, devoid of DNA from other gene pool and restricted to a modulation of existing traits from the sexually compatible gene pool, could also result in less comprehensive regulation towards the release of a precision bred plant, thereby decreasing the regulatory approval costs.<sup><xref ref-type="bibr" rid="bib373">373</xref></sup></p></sec></body><back><notes><p>The authors declare no conflict of interest.</p></notes><ref-list><ref id="bib1"><mixed-citation publication-type="journal">Mitsuhara I, Ugaki M, Hirochika H et al. <article-title>Efficient promoter cassettes for enhanced expression of foreign genes in dicotyledonous and monocotyledonous plants.</article-title><source>Plant Cell Physiol</source><year>1996</year>; <volume>37</volume>: <fpage>49</fpage>–59.<pub-id pub-id-type="pmid">8720924</pub-id></mixed-citation></ref><ref id="bib2"><mixed-citation publication-type="book">Komarnytsky S, Borisjuk N. 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align="left" valign="top" charoff="50">Promoter</th><th align="left" valign="top" charoff="50">Origin</th><th align="left" valign="top" charoff="50">Crop use</th><th align="left" valign="top" charoff="50">References</th></tr></thead><tbody valign="top"><tr><td colspan="4" align="left" valign="top" charoff="50"><bold>Constitutive expression</bold><hr></hr></td></tr><tr><td align="left" valign="top" charoff="50">BSV</td><td align="left" valign="top" charoff="50">Banana streak badnavirus</td><td align="left" valign="top" charoff="50">Banana, Sunflower</td><td align="left" valign="top" charoff="50">96</td></tr><tr><td align="left" valign="top" charoff="50">CaMV 35S</td><td align="left" valign="top" charoff="50">Cauliflower mosaic virus</td><td align="left" valign="top" charoff="50">Apple, broccoli, citrus, chrysanthemum, cocoa, collard, grape, Indian Mustard, Lilium, <italic>Nicotiana glutinosa</italic>, papaya, peach, petunia, plum, poplar, rose, strawberry, tomato, torenia</td><td align="left" valign="top" charoff="50">41–44, 47–63, 65–79, 84, 85, 87</td></tr><tr><td align="left" valign="top" charoff="50">CMPS</td><td align="left" valign="top" charoff="50">Cestrum Yellow Leaf curling virus</td><td align="left" valign="top" charoff="50">Grape</td><td align="left" valign="top" charoff="50">95</td></tr><tr><td align="left" valign="top" charoff="50">Lhca3.St.1</td><td align="left" valign="top" charoff="50">Potato</td><td align="left" valign="top" charoff="50">Chrysanthemum</td><td align="left" valign="top" charoff="50">100</td></tr><tr><td align="left" valign="top" charoff="50">Mannopin synthase</td><td align="left" valign="top" charoff="50">Gladiolus</td><td align="left" valign="top" charoff="50">Gladiolus</td><td align="left" valign="top" charoff="50">99</td></tr><tr><td align="left" valign="top" charoff="50">RolD</td><td align="left" valign="top" charoff="50"><italic>A. rhizogenes</italic></td><td align="left" valign="top" charoff="50">Gladiolus</td><td align="left" valign="top" charoff="50">99</td></tr><tr><td align="left" valign="top" charoff="50">Uep1</td><td align="left" valign="top" charoff="50">Oilpalm</td><td align="left" valign="top" charoff="50">Oilpalm, tobacco</td><td align="left" valign="top" charoff="50">98</td></tr><tr><td align="left" valign="top" charoff="50">Ubiquitin</td><td align="left" valign="top" charoff="50">Grape, gladiolus</td><td align="left" valign="top" charoff="50">Grape, gladiolus</td><td align="left" valign="top" charoff="50">49, 99</td></tr><tr><td colspan="4" align="left" valign="top" charoff="50"><bold>Fruit-specific expression</bold><hr></hr></td></tr><tr><td align="left" valign="top" charoff="50">ACC-oxidase</td><td align="left" valign="top" charoff="50">Peach, apple, tomato, banana</td><td align="left" valign="top" charoff="50">Tomato, banana</td><td align="left" valign="top" charoff="50">104–107</td></tr><tr><td align="left" valign="top" charoff="50">ADP-glucose pyrophosphorylase</td><td align="left" valign="top" charoff="50">Watermelon</td><td align="left" valign="top" charoff="50">Tomato</td><td align="left" valign="top" charoff="50">115</td></tr><tr><td align="left" valign="top" charoff="50">Expansin</td><td align="left" valign="top" charoff="50">Cherry, cucumber</td><td align="left" valign="top" charoff="50">Tomato, cucumber</td><td align="left" valign="top" charoff="50">109, 110</td></tr><tr><td align="left" valign="top" charoff="50">Cucumisin</td><td align="left" valign="top" charoff="50">Melon</td><td align="left" valign="top" charoff="50">Melon</td><td align="left" valign="top" charoff="50">114</td></tr><tr><td align="left" valign="top" charoff="50">C11</td><td align="left" valign="top" charoff="50">Citrus</td><td align="left" valign="top" charoff="50">Lemon</td><td align="left" valign="top" charoff="50">117</td></tr><tr><td align="left" valign="top" charoff="50">CsACS1G/CsACS1</td><td align="left" valign="top" charoff="50">Cucumber</td><td align="left" valign="top" charoff="50"><italic>Arabidopsis</italic></td><td align="left" valign="top" charoff="50">111</td></tr><tr><td align="left" valign="top" charoff="50">CsExp</td><td align="left" valign="top" charoff="50">Cucumber</td><td align="left" valign="top" charoff="50">Cucumber</td><td align="left" valign="top" charoff="50">110</td></tr><tr><td align="left" valign="top" charoff="50">DefH9</td><td align="left" valign="top" charoff="50">Grape</td><td align="left" valign="top" charoff="50">Grape</td><td align="left" valign="top" charoff="50">138</td></tr><tr><td align="left" valign="top" charoff="50">DFR</td><td align="left" valign="top" charoff="50">Grape</td><td align="left" valign="top" charoff="50">Grape</td><td align="left" valign="top" charoff="50">124</td></tr><tr><td align="left" valign="top" charoff="50">E8</td><td align="left" valign="top" charoff="50">Tomato</td><td align="left" valign="top" charoff="50">Tomato</td><td align="left" valign="top" charoff="50">133</td></tr><tr><td align="left" valign="top" charoff="50">Faxyl1</td><td align="left" valign="top" charoff="50">Strawberry</td><td align="left" valign="top" charoff="50">Strawberry</td><td align="left" valign="top" charoff="50">112</td></tr><tr><td align="left" valign="top" charoff="50">GIZEP</td><td align="left" valign="top" charoff="50"><italic>Gentiana lutea</italic></td><td align="left" valign="top" charoff="50">Tomato</td><td align="left" valign="top" charoff="50">126</td></tr><tr><td align="left" valign="top" charoff="50">Metallothionin</td><td align="left" valign="top" charoff="50">Citrus, oilpalm</td><td align="left" valign="top" charoff="50"><italic>Arabidopsis</italic></td><td align="left" valign="top" charoff="50">118, 119</td></tr><tr><td align="left" valign="top" charoff="50">Pac1</td><td align="left" valign="top" charoff="50">Yeast</td><td align="left" valign="top" charoff="50">Avocado</td><td align="left" valign="top" charoff="50">136</td></tr><tr><td align="left" valign="top" charoff="50">RolB</td><td align="left" valign="top" charoff="50"><italic>A. rhizogenes</italic></td><td align="left" valign="top" charoff="50">Tomato</td><td align="left" valign="top" charoff="50">134</td></tr><tr><td align="left" valign="top" charoff="50">SPS</td><td align="left" valign="top" charoff="50">Banana</td><td align="left" valign="top" charoff="50">Banana</td><td align="left" valign="top" charoff="50">116</td></tr><tr><td align="left" valign="top" charoff="50">SIACS4/SIEXP1</td><td align="left" valign="top" charoff="50">Tomato</td><td align="left" valign="top" charoff="50">Tomato</td><td align="left" valign="top" charoff="50">127</td></tr><tr><td colspan="4" align="left" valign="top" charoff="50"><bold>Seed-specific expression</bold><hr></hr></td></tr><tr><td align="left" valign="top" charoff="50">2S</td><td align="left" valign="top" charoff="50">Grape</td><td align="left" valign="top" charoff="50">Grape, tobacco</td><td align="left" valign="top" charoff="50">145, 147</td></tr><tr><td align="left" valign="top" charoff="50">CuMFT1</td><td align="left" valign="top" charoff="50">Citrus</td><td align="left" valign="top" charoff="50"><italic>Arabidopsis</italic></td><td align="left" valign="top" charoff="50">171</td></tr><tr><td align="left" valign="top" charoff="50">Dc3</td><td align="left" valign="top" charoff="50">Carrot</td><td align="left" valign="top" charoff="50"><italic>Arabidopsis</italic></td><td align="left" valign="top" charoff="50">163</td></tr><tr><td align="left" valign="top" charoff="50">HaG3-A</td><td align="left" valign="top" charoff="50">Sunflower</td><td align="left" valign="top" charoff="50">Tobacco</td><td align="left" valign="top" charoff="50">170</td></tr><tr><td align="left" valign="top" charoff="50">LeB4</td><td align="left" valign="top" charoff="50"><italic>Vicia faba</italic></td><td align="left" valign="top" charoff="50">Tobacco</td><td align="left" valign="top" charoff="50">156</td></tr><tr><td align="left" valign="top" charoff="50">LegA</td><td align="left" valign="top" charoff="50">Pea</td><td align="left" valign="top" charoff="50"><italic>Helianthus</italic></td><td align="left" valign="top" charoff="50">143</td></tr><tr><td align="left" valign="top" charoff="50">NapA</td><td align="left" valign="top" charoff="50"><italic>Brassica napus</italic></td><td align="left" valign="top" charoff="50">Tobacco</td><td align="left" valign="top" charoff="50">150</td></tr><tr><td align="left" valign="top" charoff="50">Phas</td><td align="left" valign="top" charoff="50">Bean</td><td align="left" valign="top" charoff="50">Tobacco</td><td align="left" valign="top" charoff="50">140, 172</td></tr><tr><td align="left" valign="top" charoff="50">Psl</td><td align="left" valign="top" charoff="50">Pea</td><td align="left" valign="top" charoff="50">Tobacco</td><td align="left" valign="top" charoff="50">168</td></tr><tr><td align="left" valign="top" charoff="50">Str</td><td align="left" valign="top" charoff="50"><italic>Catharanthus roseus</italic></td><td align="left" valign="top" charoff="50">Tobacco</td><td align="left" valign="top" charoff="50">173</td></tr><tr><td align="left" valign="top" charoff="50">USP</td><td align="left" valign="top" charoff="50"><italic>Vicia faba</italic></td><td align="left" valign="top" charoff="50">Tomato</td><td align="left" valign="top" charoff="50">158</td></tr><tr><td colspan="4" align="left" valign="top" charoff="50"><bold>Floral tissue-specific expression</bold><hr></hr></td></tr><tr><td align="left" valign="top" charoff="50">BAN215-6</td><td align="left" valign="top" charoff="50"><italic>Brassica campestris</italic></td><td align="left" valign="top" charoff="50">Tobacco</td><td align="left" valign="top" charoff="50">249</td></tr><tr><td align="left" valign="top" charoff="50">CHS</td><td align="left" valign="top" charoff="50">Bean</td><td align="left" valign="top" charoff="50">Petunia, tobacco</td><td align="left" valign="top" charoff="50">197, 198</td></tr><tr><td align="left" valign="top" charoff="50">END1</td><td align="left" valign="top" charoff="50">Pea</td><td align="left" valign="top" charoff="50">Tobacco</td><td align="left" valign="top" charoff="50">250</td></tr><tr><td align="left" valign="top" charoff="50">GTCHS1</td><td align="left" valign="top" charoff="50"><italic>Gentiana triflora</italic></td><td align="left" valign="top" charoff="50">Petunia</td><td align="left" valign="top" charoff="50">210</td></tr><tr><td align="left" valign="top" charoff="50">LAT52</td><td align="left" valign="top" charoff="50">Tomato</td><td align="left" valign="top" charoff="50"><italic>Lilium longiflorum</italic></td><td align="left" valign="top" charoff="50">246</td></tr><tr><td align="left" valign="top" charoff="50">PsTL1</td><td align="left" valign="top" charoff="50"><italic>Pyrus serotina</italic></td><td align="left" valign="top" charoff="50">Tobacco</td><td align="left" valign="top" charoff="50">214</td></tr><tr><td align="left" valign="top" charoff="50">SK2</td><td align="left" valign="top" charoff="50">Potato</td><td align="left" valign="top" charoff="50">Potato</td><td align="left" valign="top" charoff="50">224</td></tr><tr><td align="left" valign="top" charoff="50">TomA108</td><td align="left" valign="top" charoff="50">Tomato</td><td align="left" valign="top" charoff="50">Tobacco</td><td align="left" valign="top" charoff="50">248</td></tr><tr><td colspan="4" align="left" valign="top" charoff="50"><bold>Root-specific expression</bold><hr></hr></td></tr><tr><td align="left" valign="top" charoff="50">B33</td><td align="left" valign="top" charoff="50">Potato</td><td align="left" valign="top" charoff="50">Potato</td><td align="left" valign="top" charoff="50">284</td></tr><tr><td align="left" valign="top" charoff="50">FaRB7</td><td align="left" valign="top" charoff="50">Strawberry</td><td align="left" valign="top" charoff="50">Tobacco</td><td align="left" valign="top" charoff="50">267</td></tr><tr><td align="left" valign="top" charoff="50">Glb3 5'</td><td align="left" valign="top" charoff="50"><italic>Sesbania rostrata</italic></td><td align="left" valign="top" charoff="50">Lotus</td><td align="left" valign="top" charoff="50">273</td></tr><tr><td align="left" valign="top" charoff="50">MipB</td><td align="left" valign="top" charoff="50"><italic>Mesembryanthemum crystallinum</italic></td><td align="left" valign="top" charoff="50">Tobacco</td><td align="left" valign="top" charoff="50">268</td></tr><tr><td align="left" valign="top" charoff="50">Npv30</td><td align="left" valign="top" charoff="50">Bean</td><td align="left" valign="top" charoff="50">Lotus</td><td align="left" valign="top" charoff="50">275</td></tr><tr><td align="left" valign="top" charoff="50">PsENOD12A/PsENOD12B</td><td align="left" valign="top" charoff="50">Pea</td><td align="left" valign="top" charoff="50"><italic>Vicia hirsuta</italic></td><td align="left" valign="top" charoff="50">274</td></tr><tr><td align="left" valign="top" charoff="50">RB7</td><td align="left" valign="top" charoff="50">Tobacco</td><td align="left" valign="top" charoff="50">Tomato</td><td align="left" valign="top" charoff="50">267</td></tr><tr><td align="left" valign="top" charoff="50">SLREO</td><td align="left" valign="top" charoff="50">Tomato</td><td align="left" valign="top" charoff="50">Tomato</td><td align="left" valign="top" charoff="50">263</td></tr><tr><td align="left" valign="top" charoff="50">VfLb29</td><td align="left" valign="top" charoff="50"><italic>Vicia faba</italic></td><td align="left" valign="top" charoff="50"><italic>Vicia faba</italic></td><td align="left" valign="top" charoff="50">271</td></tr><tr><td align="left" valign="top" charoff="50">Sporamin</td><td align="left" valign="top" charoff="50">Sweet potato</td><td align="left" valign="top" charoff="50">Potato, tobacco</td><td align="left" valign="top" charoff="50">287, 288</td></tr><tr><td colspan="4" align="left" valign="top" charoff="50"><bold>Vascular tissue-specific expression</bold><hr></hr></td></tr><tr><td align="left" valign="top" charoff="50">AtSUC2</td><td align="left" valign="top" charoff="50"><italic>Arabidopsis</italic></td><td align="left" valign="top" charoff="50">Citrus, pear, strawberries</td><td align="left" valign="top" charoff="50">307, 309, 310</td></tr><tr><td align="left" valign="top" charoff="50">CoYMVP</td><td align="left" valign="top" charoff="50">Commelina Yellow Mottle Virus</td><td align="left" valign="top" charoff="50">Apple</td><td align="left" valign="top" charoff="50">313</td></tr><tr><td align="left" valign="top" charoff="50">CsPP2</td><td align="left" valign="top" charoff="50">Citrus</td><td align="left" valign="top" charoff="50">Sweet orange</td><td align="left" valign="top" charoff="50">308</td></tr><tr><td align="left" valign="top" charoff="50">CsSUS1p</td><td align="left" valign="top" charoff="50">Citrus</td><td align="left" valign="top" charoff="50"><italic>Arabidopsis</italic>/tobacco</td><td align="left" valign="top" charoff="50">311</td></tr><tr><td align="left" valign="top" charoff="50">GRP 1.8</td><td align="left" valign="top" charoff="50">Bean</td><td align="left" valign="top" charoff="50">Tobacco</td><td align="left" valign="top" charoff="50">300</td></tr><tr><td align="left" valign="top" charoff="50">GS3A</td><td align="left" valign="top" charoff="50">Pea</td><td align="left" valign="top" charoff="50">Alfalfa</td><td align="left" valign="top" charoff="50">305</td></tr><tr><td align="left" valign="top" charoff="50">PAL2</td><td align="left" valign="top" charoff="50">Bean</td><td align="left" valign="top" charoff="50">Tobacco</td><td align="left" valign="top" charoff="50">302</td></tr><tr><td align="left" valign="top" charoff="50">RolC</td><td align="left" valign="top" charoff="50"><italic>A. rhizogenes</italic></td><td align="left" valign="top" charoff="50">Apple</td><td align="left" valign="top" charoff="50">313</td></tr><tr><td align="left" valign="top" charoff="50">RTBV</td><td align="left" valign="top" charoff="50">Rice Tungro Virus</td><td align="left" valign="top" charoff="50">Citrus</td><td align="left" valign="top" charoff="50">307</td></tr><tr><td align="left" valign="top" charoff="50">Rice sucrose synthase l</td><td align="left" valign="top" charoff="50">Rice</td><td align="left" valign="top" charoff="50">Citrus</td><td align="left" valign="top" charoff="50">307</td></tr><tr><td colspan="4" align="left" valign="top" charoff="50"><bold>Inducible expression</bold><hr></hr></td></tr><tr><td align="left" valign="top" charoff="50">4CL</td><td align="left" valign="top" charoff="50">Parsley/tobacco</td><td align="left" valign="top" charoff="50">Parsley/tobacco</td><td align="left" valign="top" charoff="50">352</td></tr><tr><td align="left" valign="top" charoff="50">CM-ACO1</td><td align="left" valign="top" charoff="50">Melon</td><td align="left" valign="top" charoff="50">Tobacco</td><td align="left" valign="top" charoff="50">351</td></tr><tr><td align="left" valign="top" charoff="50">HSP</td><td align="left" valign="top" charoff="50">Tomato</td><td align="left" valign="top" charoff="50">Sunflower</td><td align="left" valign="top" charoff="50">344</td></tr><tr><td align="left" valign="top" charoff="50">Lehsp23.8</td><td align="left" valign="top" charoff="50">Tomato</td><td align="left" valign="top" charoff="50">Tomato</td><td align="left" valign="top" charoff="50">341</td></tr><tr><td align="left" valign="top" charoff="50">PinII</td><td align="left" valign="top" charoff="50">Potato</td><td align="left" valign="top" charoff="50">Alfalfa, <italic>Nicotiana plumbaginifolia</italic>, rice</td><td align="left" valign="top" charoff="50">334–336</td></tr><tr><td align="left" valign="top" charoff="50">PR-1</td><td align="left" valign="top" charoff="50">Tobacco</td><td align="left" valign="top" charoff="50">Broccoli</td><td align="left" valign="top" charoff="50">353</td></tr><tr><td align="left" valign="top" charoff="50">PR 10</td><td align="left" valign="top" charoff="50">Alfalfa</td><td align="left" valign="top" charoff="50">Grape</td><td align="left" valign="top" charoff="50">354</td></tr><tr><td align="left" valign="top" charoff="50">Prosystemin</td><td align="left" valign="top" charoff="50">Tomato</td><td align="left" valign="top" charoff="50">Tomato</td><td align="left" valign="top" charoff="50">323</td></tr><tr><td align="left" valign="top" charoff="50">Rd29A</td><td align="left" valign="top" charoff="50"><italic>Arabidopsis</italic></td><td align="left" valign="top" charoff="50">Potato</td><td align="left" valign="top" charoff="50">350</td></tr><tr><td align="left" valign="top" charoff="50">SWPA2</td><td align="left" valign="top" charoff="50">Sweet potato</td><td align="left" valign="top" charoff="50">Tobacco</td><td align="left" valign="top" charoff="50">349</td></tr><tr><td align="left" valign="top" charoff="50">Wun1</td><td align="left" valign="top" charoff="50">Potato</td><td align="left" valign="top" charoff="50">Tobacco</td><td align="left" valign="top" charoff="50">340</td></tr></tbody></table></table-wrap></floats-group></pmc></record>Pour manipuler ce document sous Unix (Dilib)
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