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<document id="79105A87969FF609FE189CF8A0A0EEB5" ID-DOI="10.1016/j.phytochem.2020.112532" ID-ISSN="1873-3700" ID-Zenodo-Dep="8292694" IM.bibliography_approvedBy="felipe" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="julia" IM.treatments_approvedBy="julia" checkinTime="1693250015429" checkinUser="felipe" docAuthor="Wang, Fumin, Park, Yong-Lak &amp; Gutensohn, Michael" docDate="2020" docId="039B87E9B058FF98FCD4BF0950BD55B5" docLanguage="en" docName="Phytochemistry.180.112532.pdf" docOrigin="Phytochemistry (112532) 180" docSource="http://dx.doi.org/10.1016/j.phytochem.2020.112532" docStyle="DocumentStyle:F36D69FC8B198FBE91029DF9C24697D3.5:Phytochemistry.2020-.journal_article" docStyleId="F36D69FC8B198FBE91029DF9C24697D3" docStyleName="Phytochemistry.2020-.journal_article" docStyleVersion="5" docTitle="Solanum habrochaites subsp. chemotypes S. Knapp &amp; D. M. Spooner" docType="treatment" docVersion="3" lastPageNumber="2" masterDocId="FFA2FF91B059FF99FFE6BE16556D5215" masterDocTitle="Glandular trichome-derived sesquiterpenes of wild tomato accessions (Solanum habrochaites) affect aphid performance and feeding behavior" masterLastPageNumber="11" masterPageNumber="1" pageNumber="2" updateTime="1693406035079" updateUser="julia">
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<mods:title id="A457405F7D727429DBBB7ED66D7C0E55">Glandular trichome-derived sesquiterpenes of wild tomato accessions (Solanum habrochaites) affect aphid performance and feeding behavior</mods:title>
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<paragraph id="8B8D36FFB058FF98FCD4BF095127535B" blockId="1.[818,1471,287,306]" lastBlockId="1.[818,1098,315,334]" pageId="1" pageNumber="2">
<emphasis id="B946EAEDB058FF98FCD4BF095127535B" bold="true" italics="true" pageId="1" pageNumber="2">
<heading id="D0C58193B058FF98FCD4BF0950D25327" bold="true" box="[818,1471,287,306]" centered="true" fontSize="36" level="1" pageId="1" pageNumber="2" reason="1">
2.1.
<taxonomicName id="4C324D7CB058FF98FC84BF09514A5327" ID-CoL="4Y2MW" ID-ENA="62890" authority="S. Knapp &amp; D. M. Spooner" authorityName="S. Knapp &amp; D. M. Spooner" box="[866,1063,287,306]" class="Magnoliopsida" family="Solanaceae" genus="Solanum" kingdom="Plantae" order="Solanales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="species" species="habrochaites">Solanum habrochaites</taxonomicName>
chemotypes produce distinct sets of glandular
</heading>
<heading id="D0C58193B058FF98FCD4BF2D5127535B" box="[818,1098,315,334]" fontSize="8" level="3" pageId="1" pageNumber="2" reason="8">trichome derived sesquiterpenes</heading>
</emphasis>
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<paragraph id="8B8D36FFB058FF98FCB7BF6556C45478" blockId="1.[818,1488,371,1952]" pageId="1" pageNumber="2">
To identify glandular trichome derived terpene traits in tomato that have the potential to improve the plant defense against piercing-sucking pests such as the potato aphid (
<taxonomicName id="4C324D7CB058FF98FBB9BFBD518D53A8" box="[1119,1248,426,446]" class="Insecta" family="Aphididae" genus="Macrosiphum" kingdom="Animalia" order="Hemiptera" pageId="1" pageNumber="2" phylum="Arthropoda" rank="species" species="euphorbiae">
<emphasis id="B946EAEDB058FF98FBB9BFBD518D53A8" bold="true" box="[1119,1248,426,446]" italics="true" pageId="1" pageNumber="2">M. euphorbiae</emphasis>
</taxonomicName>
) we have assembled and screened a collection of two
<taxonomicName id="4C324D7CB058FF98FB90BFD1506753CC" box="[1142,1290,454,474]" class="Magnoliopsida" family="Solanaceae" genus="Solanum" kingdom="Plantae" order="Solanales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="species" species="lycopersicum">
<emphasis id="B946EAEDB058FF98FB90BFD1506753CC" bold="true" box="[1142,1290,454,474]" italics="true" pageId="1" pageNumber="2">S. lycopersicum</emphasis>
</taxonomicName>
cultivars and ten
<taxonomicName id="4C324D7CB058FF98FCD4BFF456D253E0" box="[818,959,482,501]" class="Magnoliopsida" family="Solanaceae" genus="Solanum" kingdom="Plantae" order="Solanales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="species" species="habrochaites">
<emphasis id="B946EAEDB058FF98FCD4BFF456D253E0" bold="true" box="[818,959,482,501]" italics="true" pageId="1" pageNumber="2">S. habrochaites</emphasis>
</taxonomicName>
accessions that based on previous analyses (
<bibRefCitation id="EFA34B0EB058FF98FA6BBFF556985004" author="Gonzales-Vigil, E. &amp; Hufnagel, D. E. &amp; Kim, J. &amp; Last, R. L. &amp; Barry, C. S." pageId="1" pageNumber="2" pagination="921 - 935" refId="ref12830" refString="Gonzales-Vigil, E., Hufnagel, D. E., Kim, J., Last, R. L., Barry, C. S., 2012. Evolution of TPS 20 - related terpene synthases influences chemical diversity in the glandular trichomes of the wild tomato relative Solanum habrochaites. Plant J. 71, 921 - 935." type="journal article" year="2012">Gonzales-Vigil et al., 2012</bibRefCitation>
) represent five distinct chemotypes with specific terpene profiles. To test if the results of this previous characterization of glandular trichome derived terpenes could be verified for tomato plants grown under our conditions, we performed leaf dip extractions for these tomato accessions and analyzed the resulting extracts by combined gas chromatography-mass spectrometry (GC-MS). Indeed, our analysis confirmed that the blend of terpenes produced in glandular trichomes differed significantly between
<taxonomicName id="4C324D7CB058FF98FBADBCD751BE50C1" box="[1099,1235,705,724]" class="Magnoliopsida" family="Solanaceae" genus="Solanum" kingdom="Plantae" order="Solanales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="species" species="lycopersicum">
<emphasis id="B946EAEDB058FF98FBADBCD751BE50C1" bold="true" box="[1099,1235,705,724]" italics="true" pageId="1" pageNumber="2">S. lycopersicum</emphasis>
</taxonomicName>
and
<taxonomicName id="4C324D7CB058FF98FB19BCD750E850C1" box="[1279,1413,705,724]" class="Magnoliopsida" family="Solanaceae" genus="Solanum" kingdom="Plantae" order="Solanales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="species" species="habrochaites">
<emphasis id="B946EAEDB058FF98FB19BCD750E850C1" bold="true" box="[1279,1413,705,724]" italics="true" pageId="1" pageNumber="2">S. habrochaites</emphasis>
</taxonomicName>
, as well as among different
<taxonomicName id="4C324D7CB058FF98FC1DBCCB51E550E5" box="[1019,1160,733,752]" class="Magnoliopsida" family="Solanaceae" genus="Solanum" kingdom="Plantae" order="Solanales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="species" species="habrochaites">
<emphasis id="B946EAEDB058FF98FC1DBCCB51E550E5" bold="true" box="[1019,1160,733,752]" italics="true" pageId="1" pageNumber="2">S. habrochaites</emphasis>
</taxonomicName>
accessions (Table S1). The two
<taxonomicName id="4C324D7CB058FF98FCD4BCEF56D05119" box="[818,957,761,780]" class="Magnoliopsida" family="Solanaceae" genus="Solanum" kingdom="Plantae" order="Solanales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="species" species="lycopersicum">
<emphasis id="B946EAEDB058FF98FCD4BCEF56D05119" bold="true" box="[818,957,761,780]" italics="true" pageId="1" pageNumber="2">S. lycopersicum</emphasis>
</taxonomicName>
cultivars (c.v. M82 and c.v. Moneymaker) produced a mixture of monoterpenes with β- phellandrene being the most prominent compound besides smaller amounts of α- pinene, δ-2-carene, and α- phellandrene. In addition, both cultivars produced minor amounts of the sesquiterpenes β- caryophyllene and α- humulene, while δ- elemene was only found in extracts of c.v. M82. In contrast to the
<taxonomicName id="4C324D7CB058FF98FAA0BD9350A25182" box="[1350,1487,900,920]" class="Magnoliopsida" family="Solanaceae" genus="Solanum" kingdom="Plantae" order="Solanales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="species" species="lycopersicum">
<emphasis id="B946EAEDB058FF98FAA0BD9350A25182" bold="true" box="[1350,1487,900,920]" italics="true" pageId="1" pageNumber="2">S. lycopersicum</emphasis>
</taxonomicName>
cultivars the
<taxonomicName id="4C324D7CB058FF98FC56BDB7515751A6" box="[944,1082,928,948]" class="Magnoliopsida" family="Solanaceae" genus="Solanum" kingdom="Plantae" order="Solanales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="species" species="habrochaites">
<emphasis id="B946EAEDB058FF98FC56BDB7515751A6" bold="true" box="[944,1082,928,948]" italics="true" pageId="1" pageNumber="2">S. habrochaites</emphasis>
</taxonomicName>
accessions lack monoterpenes and can be sorted into five different chemotypes based on the profile of sesquiterpenes produced (Table S1). Chemotype 1 includes the
<taxonomicName id="4C324D7CB058FF98FAA0BDCF50A251FE" box="[1350,1487,984,1004]" class="Magnoliopsida" family="Solanaceae" genus="Solanum" kingdom="Plantae" order="Solanales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="species" species="habrochaites">
<emphasis id="B946EAEDB058FF98FAA0BDCF50A251FE" bold="true" box="[1350,1487,984,1004]" italics="true" pageId="1" pageNumber="2">S. habrochaites</emphasis>
</taxonomicName>
accessions LA1691 and LA2650, and primarily produces 7-epizingiberene and smaller quantities of
<emphasis id="B946EAEDB058FF98FBACBA0751355631" bold="true" box="[1098,1112,1041,1060]" italics="true" pageId="1" pageNumber="2">R</emphasis>
-curcumene. Leaf dip extracts from chemotype 2, represented by the accessions LA 1721 and LA1927, contained large amounts of γ- elemene in addition to smaller amounts of δ- elemene. A similar qualitative composition was found for extracts from chemotype 3, including accessions LA1978 and LA2155; however, these contained large amounts of δ- elemene and only small quantities of γ- elemene. Extracts from chemotype 4, represented by accessions LA1775 and LA1779, also contained large amounts of γ- elemene and small amounts of δ- elemene like those from chemotype 2. However, accessions of chemotype 4 are characterized by the production of substantial amounts of α- santalene, α- bergamotene and β- bergamotene as well as a few minor compounds including β- elemene and (
<emphasis id="B946EAEDB058FF98FCDCBB49562B5767" bold="true" box="[826,838,1375,1394]" italics="true" pageId="1" pageNumber="2">Z</emphasis>
)-α- farnesene. Extracts from chemotype 5, comprised of accessions LA 1624 and LA2860, contained the two sesquiterpenes β- caryophyllene and α- humulene that were also found in
<taxonomicName id="4C324D7CB058FF98FB48BB81505A57BF" box="[1198,1335,1431,1450]" class="Magnoliopsida" family="Solanaceae" genus="Solanum" kingdom="Plantae" order="Solanales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="species" species="lycopersicum">
<emphasis id="B946EAEDB058FF98FB48BB81505A57BF" bold="true" box="[1198,1335,1431,1450]" italics="true" pageId="1" pageNumber="2">S. lycopersicum</emphasis>
</taxonomicName>
, however, at up to 26- and 51-fold higher quantities, respectively. While δ- elemene, γ- elemene, and β- elemene were found in the extracts of many accessions, including those of chemotypes 2, 3, 4, and c.v. M82, these are likely the result of Cope rearrangements occurring upon sample injection into the hot (220
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C) GC injector port (
<bibRefCitation id="EFA34B0EB058FF98FBB3B83451935420" author="Colby, S. M. &amp; Crock, J. &amp; Dowdle-Rizzo, B. &amp; Lemaux, P. G. &amp; Croteau, R." box="[1109,1278,1570,1590]" pageId="1" pageNumber="2" pagination="2216 - 2221" refId="ref12248" refString="Colby, S. M., Crock, J., Dowdle-Rizzo, B., Lemaux, P. G., Croteau, R., 1998. Germacrene C synthase from Lycopersicon esculentum cv. VFNT Cherry tomato: cDNA isolation, characterization, and bacterial expression of the multiple product sesquiterpene cyclase. Proc. Natl. Acad. Sci. U. S. A. 95, 2216 - 2221." type="journal article" year="1998">Colby et al., 1998</bibRefCitation>
), thus suggesting that glandular trichomes of these accessions produce germacrene C, B, and A, respectively.
</paragraph>
<paragraph id="8B8D36FFB058FF98FCB7B86050BD55B5" blockId="1.[818,1488,371,1952]" pageId="1" pageNumber="2">
The analysis of tomato leaf dip extracts revealed the accumulation of terpenes in glandular trichomes, which can affect aphids once they are on the leaf surface. However, the behavior of aphids could already be modified prior to landing on the host plant through terpenes emitted from leaves into the atmosphere. Thus, we also performed headspace collections from leaves followed by GC-MS analysis to characterize the profile of emitted terpenes for all the tomato accessions qualitatively and quantitatively. In summary the results of this analysis of emitted terpenes (Table S2) are in line with the previous analysis of leaf dip extracts and further confirm the observed differences in the terpene profiles between
<taxonomicName id="4C324D7CB058FF98FC36B99B513555B5" box="[976,1112,1933,1952]" class="Magnoliopsida" family="Solanaceae" genus="Solanum" kingdom="Plantae" order="Solanales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="species" species="lycopersicum">
<emphasis id="B946EAEDB058FF98FC36B99B513555B5" bold="true" box="[976,1112,1933,1952]" italics="true" pageId="1" pageNumber="2">S. lycopersicum</emphasis>
</taxonomicName>
and the five
<taxonomicName id="4C324D7CB058FF98FB29B99B50A055B5" box="[1231,1485,1933,1952]" class="Magnoliopsida" family="Solanaceae" genus="Solanum" kingdom="Plantae" order="Solanales" pageId="1" pageNumber="3" phylum="Tracheophyta" rank="subSpecies" species="habrochaites" subSpecies="chemotypes">
<emphasis id="B946EAEDB058FF98FB29B99B503855B5" bold="true" box="[1231,1365,1933,1952]" italics="true" pageId="1" pageNumber="2">S. habrochaites</emphasis>
chemotypes
</taxonomicName>
.
</paragraph>
</subSubSection>
</treatment>
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