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<document id="AC0DE8FA2705ED8EFAEF9C5D889CA9F8" ID-DOI="10.1016/j.phytochem.2015.11.005" ID-ISSN="1873-3700" ID-Zenodo-Dep="10485543" IM.bibliography_approvedBy="julia" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="julia" IM.taxonomicNames_approvedBy="julia" IM.treatments_approvedBy="julia" checkinTime="1704945321634" checkinUser="felipe" docAuthor="Ivanov, Dimitre A., Georgakopoulos, Jorge R. C. &amp; Bernards, Mark A." docDate="2016" docId="702F87A57126FFC1FCC24E00FC02FADD" docLanguage="en" docName="Phytochemistry.122.56-64.pdf" docOrigin="Phytochemistry 122" docSource="http://dx.doi.org/10.1016/j.phytochem.2015.11.005" docStyle="DocumentStyle:9E596C34F4E94307D29315B03ACE1007.6:Phytochemistry.2014-2019.journal_article" docStyleId="9E596C34F4E94307D29315B03ACE1007" docStyleName="Phytochemistry.2014-2019.journal_article" docStyleVersion="6" docTitle="Pythium irregulare subsp. cultures Buisman" docType="treatment" docVersion="1" lastPageNumber="61" masterDocId="8C16FFDD7122FFC4FFE74C04FFFDFFD4" masterDocTitle="The chemoattractant potential of ginsenosides in the ginseng - Pythium irregulare pathosystem" masterLastPageNumber="64" masterPageNumber="56" pageNumber="60" updateTime="1705344678540" updateUser="julia">
<mods:mods id="E0BDC78BA562A8BA2AA6BA7EDBDB52A1" xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:title id="A9C013F89993D9B8AEDD5345367570CD">The chemoattractant potential of ginsenosides in the ginseng - Pythium irregulare pathosystem</mods:title>
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<mods:namePart id="67EA89D3A133BCC4F63327A00E862B18">Ivanov, Dimitre A.</mods:namePart>
<mods:affiliation id="2AA7D39CC50C3664FE0435568E86857A">Department of Biology and the Biotron, The University of Western Ontario, London, ON N 6 A 5 B 7, Canada</mods:affiliation>
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<mods:namePart id="BD052AB9F406B5B6998964D0DF8D5564">Georgakopoulos, Jorge R. C.</mods:namePart>
<mods:affiliation id="DE05E9874CA20CAEC0A0A9657A0A030D">Department of Biology and the Biotron, The University of Western Ontario, London, ON N 6 A 5 B 7, Canada</mods:affiliation>
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<mods:namePart id="C263D4E4DD65B2D07EA9E4C7515037C8">Bernards, Mark A.</mods:namePart>
<mods:affiliation id="4FB4FD8035895CD56DA1A0A40976577B">Department of Biology and the Biotron, The University of Western Ontario, London, ON N 6 A 5 B 7, Canada</mods:affiliation>
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<mods:title id="C2AD9248388831C752D302934F7E1A2D">Phytochemistry</mods:title>
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<subSubSection id="B09C65387126FFC0FCC24E00FC07FDE0" pageId="4" pageNumber="60" type="nomenclature">
<paragraph id="F83936B37126FFC0FCC24E00FC07FDE0" blockId="4.[805,1459,516,564]" pageId="4" pageNumber="60">
<heading id="A37181DF7126FFC0FCC24E00FC07FDE0" fontSize="8" level="3" pageId="4" pageNumber="60" reason="8">
<emphasis id="CAF2EAA17126FFC0FCC24E00FC07FDE0" italics="true" pageId="4" pageNumber="60">
2.3. In vitro exposure of
<taxonomicName id="3F864D307126FFC0FBF54E00FB75FDCC" authority="Buisman" authorityName="Buisman" baseAuthorityName="BR" box="[1042,1160,516,536]" class="Magnoliopsida" family="Araliaceae" genus="Pythium" kingdom="Plantae" order="Apiales" pageId="4" pageNumber="60" phylum="Tracheophyta" rank="species" species="irregulare">P. irregulare</taxonomicName>
to purified ginsenoside extract and pure ginsenosides
</emphasis>
</heading>
</paragraph>
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<subSubSection id="B09C65387126FFC1FCA34E5DFC02FADD" lastPageId="5" lastPageNumber="61" pageId="4" pageNumber="60" type="description">
<paragraph id="F83936B37126FFC0FCA34E5DFB3DFA18" blockId="4.[805,1475,601,2015]" pageId="4" pageNumber="60">
The effects of a purified total ginsenoside extract and selected pure protopanaxadiol (Rb1 (
<emphasis id="CAF2EAA17126FFC0FBAE4E70FBABFD53" bold="true" box="[1097,1110,628,647]" pageId="4" pageNumber="60">1</emphasis>
), F2 (
<emphasis id="CAF2EAA17126FFC0FB734E70FB5CFD53" bold="true" box="[1172,1185,628,647]" pageId="4" pageNumber="60">4</emphasis>
)) and protopanaxatriol ginsenosides (Re,
<emphasis id="CAF2EAA17126FFC0FC5D4E94FC3AFD77" bold="true" box="[954,967,656,675]" pageId="4" pageNumber="60">6</emphasis>
) on the growth of
<emphasis id="CAF2EAA17126FFC0FB6E4E8BFAB0FD77" box="[1161,1357,655,675]" italics="true" pageId="4" pageNumber="60">
<taxonomicName id="3F864D307126FFC0FB6E4E8BFB02FD77" baseAuthorityName="BR" baseAuthorityYear="1068" box="[1161,1279,655,675]" class="Magnoliopsida" family="Araliaceae" genus="Pythium" kingdom="Plantae" order="Apiales" pageId="4" pageNumber="60" phylum="Tracheophyta" rank="species" species="irregulare">P. irregulare</taxonomicName>
in vitro
</emphasis>
allowed us to gain a more complete understanding of the effect a point source of ginsenosides had on the growth pattern of the pathogen. Differential growth of mycelia from
<taxonomicName id="3F864D307126FFC0FBB34EE7FB35FD23" baseAuthorityName="BR" baseAuthorityYear="1068" box="[1108,1224,739,759]" class="Magnoliopsida" family="Araliaceae" genus="Pythium" kingdom="Plantae" order="Apiales" pageId="4" pageNumber="60" phylum="Tracheophyta" rank="species" species="irregulare">
<emphasis id="CAF2EAA17126FFC0FBB34EE7FB35FD23" box="[1108,1224,739,759]" italics="true" pageId="4" pageNumber="60">P. irregulare</emphasis>
</taxonomicName>
was monitored as it progressed toward ginsenoside-treated and non-treated disks on minimal Czapek-Dox Mineral Agar (
<figureCitation id="60BD2A367126FFC0FB504F18FB0AFCFB" box="[1207,1271,796,815]" captionStart="Fig" captionStartId="5.[113,139,827,841]" captionTargetBox="[160,1444,182,798]" captionTargetId="figure-608@5.[159,1445,181,798]" captionTargetPageId="5" captionText="Fig. 5. Effect of ginsenosides on Pythium irregulare BR 1068 growth in vitro. Representative images of 90 mm petri dishes filled with Czapek Dox Mineral Agar containing 6 mm antibiotic-assay disks placed an equal distance from an 8 mm diameter plug of P. irregulare BR 1068 are portrayed. Disks on the left in each column are infused with 100 µL, (A) 10 mg mL–1 purified ginsenoside extract [GSF (G)], (B) 5 mg mL–1 ginsenoside Re (6), (C) 5 mg mL–1 ginsenoside Rb1 (1) or (D) 5 mg mL–1 ginsenoside F2 (4). All disks on the right in each column are controls infused with 100 µL MeOH. Experiment was performed in triplicate (N = 3). Images shown are representative of a single plate, for each treatment, monitored every 24 h for 120 h. Earlier (24 h) images are not included because mycelia had not reached filters and no effects were seen in any treatments. All ginsenosides were dissolved in MeOH. The apparent discolouration observed in the medium around the F2 (4) infused disks is attributed to ginsenosides that had leached into the agar medium from the disk after evaporation of the MeOH solvent. This effect is present in all other ginsenoside infused filters to varying degrees, and arises from manipulation of the lighting used to provide contrast between the mycelia and background medium." figureDoi="http://doi.org/10.5281/zenodo.10485553" httpUri="https://zenodo.org/record/10485553/files/figure.png" pageId="4" pageNumber="60">Fig. 5</figureCitation>
). Mycelial growth appeared to be unaffected by the presence of the control disks (evaporated MeOH) as well as disks infused with
<quantity id="3F7E9B567126FFC0FAD14F50FA87FCB3" box="[1334,1402,852,871]" metricMagnitude="-7" metricUnit="kg" metricValue="5.0" pageId="4" pageNumber="60" unit="mg" value="0.5">0.5 mg</quantity>
of the 20(
<emphasis id="CAF2EAA17126FFC0FCA14F6BFCACFC57" box="[838,849,879,899]" italics="true" pageId="4" pageNumber="60">S</emphasis>
)-protopanaxatriol ginsenoside Re (
<emphasis id="CAF2EAA17126FFC0FB584F74FB31FC57" bold="true" box="[1215,1228,880,899]" pageId="4" pageNumber="60">6</emphasis>
,
<figureCitation id="60BD2A367126FFC0FB3D4F74FADEFC57" box="[1242,1315,880,899]" captionStart="Fig" captionStartId="5.[113,139,827,841]" captionTargetBox="[160,1444,182,798]" captionTargetId="figure-608@5.[159,1445,181,798]" captionTargetPageId="5" captionText="Fig. 5. Effect of ginsenosides on Pythium irregulare BR 1068 growth in vitro. Representative images of 90 mm petri dishes filled with Czapek Dox Mineral Agar containing 6 mm antibiotic-assay disks placed an equal distance from an 8 mm diameter plug of P. irregulare BR 1068 are portrayed. Disks on the left in each column are infused with 100 µL, (A) 10 mg mL–1 purified ginsenoside extract [GSF (G)], (B) 5 mg mL–1 ginsenoside Re (6), (C) 5 mg mL–1 ginsenoside Rb1 (1) or (D) 5 mg mL–1 ginsenoside F2 (4). All disks on the right in each column are controls infused with 100 µL MeOH. Experiment was performed in triplicate (N = 3). Images shown are representative of a single plate, for each treatment, monitored every 24 h for 120 h. Earlier (24 h) images are not included because mycelia had not reached filters and no effects were seen in any treatments. All ginsenosides were dissolved in MeOH. The apparent discolouration observed in the medium around the F2 (4) infused disks is attributed to ginsenosides that had leached into the agar medium from the disk after evaporation of the MeOH solvent. This effect is present in all other ginsenoside infused filters to varying degrees, and arises from manipulation of the lighting used to provide contrast between the mycelia and background medium." figureDoi="http://doi.org/10.5281/zenodo.10485553" httpUri="https://zenodo.org/record/10485553/files/figure.png" pageId="4" pageNumber="60">Fig. 5B</figureCitation>
). Mycelia grew directly through these disks, showing no visible alteration in growth habit up to five days after initiation of growth. Exposure of
<taxonomicName id="3F864D307126FFC0FCA54FC6FC46FC02" baseAuthorityName="BR" baseAuthorityYear="1068" box="[834,955,962,982]" class="Magnoliopsida" family="Araliaceae" genus="Pythium" kingdom="Plantae" order="Apiales" pageId="4" pageNumber="60" phylum="Tracheophyta" rank="species" species="irregulare">
<emphasis id="CAF2EAA17126FFC0FCA54FC6FC46FC02" box="[834,955,962,982]" italics="true" pageId="4" pageNumber="60">P. irregulare</emphasis>
</taxonomicName>
to purified total ginsenoside extract (GSF;
<quantity id="3F7E9B567126FFC0FA6F4FC7FA46FC02" box="[1416,1467,963,982]" metricMagnitude="-6" metricUnit="kg" metricValue="1.0" pageId="4" pageNumber="60" unit="mg" value="1.0">1 mg</quantity>
) resulted in enhanced aerial mycelial growth and a distinct accumulation of mycelia around treated disks, albeit a short distance away from the filter disk (
<figureCitation id="60BD2A367126FFC0FC114813FBC2FBFE" box="[1014,1087,1047,1066]" captionStart="Fig" captionStartId="5.[113,139,827,841]" captionTargetBox="[160,1444,182,798]" captionTargetId="figure-608@5.[159,1445,181,798]" captionTargetPageId="5" captionText="Fig. 5. Effect of ginsenosides on Pythium irregulare BR 1068 growth in vitro. Representative images of 90 mm petri dishes filled with Czapek Dox Mineral Agar containing 6 mm antibiotic-assay disks placed an equal distance from an 8 mm diameter plug of P. irregulare BR 1068 are portrayed. Disks on the left in each column are infused with 100 µL, (A) 10 mg mL–1 purified ginsenoside extract [GSF (G)], (B) 5 mg mL–1 ginsenoside Re (6), (C) 5 mg mL–1 ginsenoside Rb1 (1) or (D) 5 mg mL–1 ginsenoside F2 (4). All disks on the right in each column are controls infused with 100 µL MeOH. Experiment was performed in triplicate (N = 3). Images shown are representative of a single plate, for each treatment, monitored every 24 h for 120 h. Earlier (24 h) images are not included because mycelia had not reached filters and no effects were seen in any treatments. All ginsenosides were dissolved in MeOH. The apparent discolouration observed in the medium around the F2 (4) infused disks is attributed to ginsenosides that had leached into the agar medium from the disk after evaporation of the MeOH solvent. This effect is present in all other ginsenoside infused filters to varying degrees, and arises from manipulation of the lighting used to provide contrast between the mycelia and background medium." figureDoi="http://doi.org/10.5281/zenodo.10485553" httpUri="https://zenodo.org/record/10485553/files/figure.png" pageId="4" pageNumber="60">Fig. 5A</figureCitation>
). This accumulation persisted for several days, forming a visible ring around the filter disk. Mycelia continued to project out from this ring and accumulate directly around the assay filter disk until the end of the monitoring period of five days. Similarly, exposure of
<taxonomicName id="3F864D307126FFC0FBA24882FB41FB4E" baseAuthorityName="BR" baseAuthorityYear="1068" box="[1093,1212,1158,1178]" class="Magnoliopsida" family="Araliaceae" genus="Pythium" kingdom="Plantae" order="Apiales" pageId="4" pageNumber="60" phylum="Tracheophyta" rank="species" species="irregulare">
<emphasis id="CAF2EAA17126FFC0FBA24882FB41FB4E" box="[1093,1212,1158,1178]" italics="true" pageId="4" pageNumber="60">P. irregulare</emphasis>
</taxonomicName>
to 20(
<emphasis id="CAF2EAA17126FFC0FAE54882FAF0FB4E" box="[1282,1293,1158,1178]" italics="true" pageId="4" pageNumber="60">S</emphasis>
)-protopanaxadiol ginsenoside Rb1 (
<emphasis id="CAF2EAA17126FFC0FC3248A6FC1FFB61" bold="true" box="[981,994,1186,1205]" pageId="4" pageNumber="60">1</emphasis>
) (
<quantity id="3F7E9B567126FFC0FC1048A6FBBDFB62" box="[1015,1088,1186,1206]" metricMagnitude="-7" metricUnit="kg" metricValue="5.0" pageId="4" pageNumber="60" unit="mg" value="0.5">0.5 mg</quantity>
; Rb1 (
<emphasis id="CAF2EAA17126FFC0FB9948A6FB76FB61" bold="true" box="[1150,1163,1186,1205]" pageId="4" pageNumber="60">1</emphasis>
) represents ca. half of the total GSF) (
<figureCitation id="60BD2A367126FFC0FC8448BAFC54FB06" box="[867,937,1214,1234]" captionStart="Fig" captionStartId="5.[113,139,827,841]" captionTargetBox="[160,1444,182,798]" captionTargetId="figure-608@5.[159,1445,181,798]" captionTargetPageId="5" captionText="Fig. 5. Effect of ginsenosides on Pythium irregulare BR 1068 growth in vitro. Representative images of 90 mm petri dishes filled with Czapek Dox Mineral Agar containing 6 mm antibiotic-assay disks placed an equal distance from an 8 mm diameter plug of P. irregulare BR 1068 are portrayed. Disks on the left in each column are infused with 100 µL, (A) 10 mg mL–1 purified ginsenoside extract [GSF (G)], (B) 5 mg mL–1 ginsenoside Re (6), (C) 5 mg mL–1 ginsenoside Rb1 (1) or (D) 5 mg mL–1 ginsenoside F2 (4). All disks on the right in each column are controls infused with 100 µL MeOH. Experiment was performed in triplicate (N = 3). Images shown are representative of a single plate, for each treatment, monitored every 24 h for 120 h. Earlier (24 h) images are not included because mycelia had not reached filters and no effects were seen in any treatments. All ginsenosides were dissolved in MeOH. The apparent discolouration observed in the medium around the F2 (4) infused disks is attributed to ginsenosides that had leached into the agar medium from the disk after evaporation of the MeOH solvent. This effect is present in all other ginsenoside infused filters to varying degrees, and arises from manipulation of the lighting used to provide contrast between the mycelia and background medium." figureDoi="http://doi.org/10.5281/zenodo.10485553" httpUri="https://zenodo.org/record/10485553/files/figure.png" pageId="4" pageNumber="60">Fig. 5C</figureCitation>
) resulted in an accumulation of mycelia around the treated disk. After five days, the ring of mycelia around the filter disk was complete and mycelial growth projected beyond it. That ginsenoside Re (
<emphasis id="CAF2EAA17126FFC0FC324916FC1FFAF1" bold="true" box="[981,994,1298,1317]" pageId="4" pageNumber="60">6</emphasis>
) had no effect, positive or negative, on the growth of
<emphasis id="CAF2EAA17126FFC0FC764929FBA5FA95" box="[913,1112,1325,1345]" italics="true" pageId="4" pageNumber="60">
<taxonomicName id="3F864D307126FFC0FC764929FBF5FA95" baseAuthorityName="BR" baseAuthorityYear="1068" box="[913,1032,1325,1345]" class="Magnoliopsida" family="Araliaceae" genus="Pythium" kingdom="Plantae" order="Apiales" pageId="4" pageNumber="60" phylum="Tracheophyta" rank="species" species="irregulare">P. irregulare</taxonomicName>
in vitro
</emphasis>
, while both GSF and Rb1 (
<emphasis id="CAF2EAA17126FFC0FA88492AFA81FA95" bold="true" box="[1391,1404,1326,1345]" pageId="4" pageNumber="60">1</emphasis>
) alone did, suggests that the protopanaxadiol ginsenosides are also the more bioactive towards this pathogen. These are also the only ginsenosides shown to be enzymatically altered by
<taxonomicName id="3F864D307126FFC0FAC04985FA62FA41" baseAuthorityName="BR" baseAuthorityYear="1068" box="[1319,1439,1409,1429]" class="Magnoliopsida" family="Araliaceae" genus="Pythium" kingdom="Plantae" order="Apiales" pageId="4" pageNumber="60" phylum="Tracheophyta" rank="species" species="irregulare">
<emphasis id="CAF2EAA17126FFC0FAC04985FA62FA41" box="[1319,1439,1409,1429]" italics="true" pageId="4" pageNumber="60">P. irregulare</emphasis>
</taxonomicName>
by extracellular glycosidases (
<bibRefCitation id="9C174B427126FFC0FBA5499AFA9BFA65" author="Yousef, L. F. &amp; Bernards, M. A." box="[1090,1382,1438,1457]" pageId="4" pageNumber="60" pagination="1740 - 1749" refId="ref10407" refString="Yousef, L. F., Bernards, M. A., 2006. In vitro metabolism of ginsenosides by the ginseng root pathogen Pythium irregulare. Phytochemistry 67, 1740 - 1749." type="journal article" year="2006">Yousef and Bernards, 2006</bibRefCitation>
;
<bibRefCitation id="9C174B427126FFC0FA91499AFC65FA18" author="Neculai, M. A. &amp; Ivanov, D. &amp; Bernards, M. A." pageId="4" pageNumber="60" pagination="1948 - 1957" refId="ref9457" refString="Neculai, M. A., Ivanov, D., Bernards, M. A., 2009. Partial purification and characterization of three ginsenoside-metabolizing [beta] - glucosidases from Pythium irregulare. Phytochemistry 70, 1948 - 1957." type="journal article" year="2009">Neculai et al., 2009</bibRefCitation>
;
<bibRefCitation id="9C174B427126FFC0FC4449BEFB4EFA18" author="Ivanov, D. A. &amp; Bernards, M. A." box="[931,1203,1465,1485]" pageId="4" pageNumber="60" pagination="44 - 53" refId="ref8841" refString="Ivanov, D. A., Bernards, M. A., 2012. Ginsenosidases and the pathogenicity of Pythium irregulare. Phytochemistry 78, 44 - 53." type="journal article" year="2012">Ivanov and Bernards, 2012</bibRefCitation>
).
</paragraph>
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<emphasis id="CAF2EAA17126FFC0FFB04931FF74FA97" bold="true" box="[87,137,1333,1347]" pageId="4" pageNumber="60">Fig. 4.</emphasis>
Time course of ginseng infection by
<taxonomicName id="3F864D307126FFC0FE514930FDBBFA97" authorityName="BR" authorityYear="1068" box="[438,582,1332,1347]" class="Magnoliopsida" family="Araliaceae" genus="Pythium" kingdom="Plantae" order="Apiales" pageId="4" pageNumber="60" phylum="Tracheophyta" rank="species" species="irregulare">
<emphasis id="CAF2EAA17126FFC0FE514930FDBBFA97" box="[438,582,1332,1347]" italics="true" pageId="4" pageNumber="60">Pythium irregulare</emphasis>
</taxonomicName>
. Ginseng plants were soaked in either ddH O or 10 mg mL
<superScript id="0FF39BFB7126FFC0FE6F494CFE64FA86" attach="left" box="[392,409,1352,1362]" fontSize="4" pageId="4" pageNumber="60">–1</superScript>
(ddH O) purified ginsenoside extract and
<subScript id="640234F67126FFC0FEE14957FE2DFA89" attach="both" box="[262,464,1363,1373]" fontSize="4" pageId="4" pageNumber="60">2 2</subScript>
planted 8 cm apart. Uninoculated plants: ΦNO values for (A) one- and (B) two year old plants treated with 10 mg mL
<superScript id="0FF39BFB7126FFC0FE944971FE79FA54" attach="left" box="[371,388,1397,1408]" fontSize="4" pageId="4" pageNumber="60">–1</superScript>
(dissolved in ddH O) purified ginsenosides
<subScript id="640234F67126FFC0FDC54984FDD4FA5E" attach="both" box="[546,553,1408,1418]" fontSize="4" pageId="4" pageNumber="60">2</subScript>
extract (filled circles) were collected every day for 15 days (one-year old) and for 20 days (two-year old). Inoculated plants: A 13 mm column of sand was removed 4 cm away from the base of each plant, in the middle of the pot, and filled with a
<taxonomicName id="3F864D307126FFC0FFB049D0FEC2FA37" authority="(BR 1068)" baseAuthorityName="BR" baseAuthorityYear="1068" box="[87,319,1492,1507]" class="Magnoliopsida" family="Araliaceae" genus="Pythium" kingdom="Plantae" order="Apiales" pageId="4" pageNumber="60" phylum="Tracheophyta" rank="species" species="irregulare">
<emphasis id="CAF2EAA17126FFC0FFB049D0FF14FA37" box="[87,233,1492,1507]" italics="true" pageId="4" pageNumber="60">Pythium irregulare</emphasis>
(BR 1068)
</taxonomicName>
-sand mix inoculum. ΦNO values for (A) one- and (B) two year old plants treated with ddH O (open circles) or 10 mg mL
<superScript id="0FF39BFB7126FFC0FD9049ECFD75FA26" attach="left" box="[631,648,1512,1522]" fontSize="4" pageId="4" pageNumber="60">–1</superScript>
(dissolved in
<subScript id="640234F67126FFC0FE6149F7FE70FA29" attach="both" box="[390,397,1523,1533]" fontSize="4" pageId="4" pageNumber="60">2</subScript>
ddH
<subScript id="640234F67126FFC0FF9E4A0DFF7DF9C7" attach="both" box="[121,128,1545,1555]" fontSize="4" pageId="4" pageNumber="60">2</subScript>
O) purified ginsenosides extract (open triangles) were collected every day for 15 days (one-year old) and for 20 days (two-year old). All ΦNO values are expressed as Mean ± SE.
<emphasis id="CAF2EAA17126FFC0FF354A34FF23F9EB" box="[210,222,1584,1599]" italics="true" pageId="4" pageNumber="60">N</emphasis>
= 6 for one-year old plant treatments;
<emphasis id="CAF2EAA17126FFC0FDD04A34FDBEF9EB" box="[567,579,1584,1599]" italics="true" pageId="4" pageNumber="60">N</emphasis>
= 4 for one-year old control;
<emphasis id="CAF2EAA17126FFC0FF7B4A43FF55F982" box="[156,168,1607,1622]" italics="true" pageId="4" pageNumber="60">N</emphasis>
= 8 for two-year old plant treatments and control.
</paragraph>
</caption>
<paragraph id="F83936B37126FFC0FCA349D1FCA3F92B" blockId="4.[805,1475,601,2015]" pageId="4" pageNumber="60">
The ring of enhanced mycelial growth around assay disks infused with GSF or ginsenoside Rb1
<emphasis id="CAF2EAA17126FFC0FB5A49F5FB37F9D0" bold="true" box="[1213,1226,1521,1540]" pageId="4" pageNumber="60">1</emphasis>
presumably represents the point of primary contact between the pathogen and the ginsenosides because otherwise
<taxonomicName id="3F864D307126FFC0FB864A2CFB20F9E8" baseAuthorityName="BR" baseAuthorityYear="1068" box="[1121,1245,1576,1596]" class="Magnoliopsida" family="Araliaceae" genus="Pythium" kingdom="Plantae" order="Apiales" pageId="4" pageNumber="60" phylum="Tracheophyta" rank="species" species="irregulare">
<emphasis id="CAF2EAA17126FFC0FB864A2CFB20F9E8" box="[1121,1245,1576,1596]" italics="true" pageId="4" pageNumber="60">P. irregulare</emphasis>
</taxonomicName>
mycelia grew thinly and in all directions until within close proximity to the treated disks. While there was extensive mycelial growth and hyphal branching observed around the GSF and Rb1 (
<emphasis id="CAF2EAA17126FFC0FB0C4A79FB05F944" bold="true" box="[1259,1272,1661,1680]" pageId="4" pageNumber="60">1</emphasis>
) treated disks, upon close inspection, it was evident that hyphal growth was not inhibited, and spread directly through the medium around and under the treated filter disks the same way it did through the control disks.
</paragraph>
<paragraph id="F83936B37126FFC1FCA34B0DFE68FBB6" blockId="4.[805,1475,601,2015]" lastBlockId="5.[113,784,1075,2015]" lastPageId="5" lastPageNumber="61" pageId="4" pageNumber="60">
In contrast, the growth of mycelia was clearly inhibited around the assay disk infused with ginsenoside F2 (
<emphasis id="CAF2EAA17126FFC0FB144B20FAFDF8E3" bold="true" box="[1267,1280,1828,1847]" pageId="4" pageNumber="60">4</emphasis>
) (
<figureCitation id="60BD2A367126FFC0FAFF4B20FA99F8E3" box="[1304,1380,1828,1847]" captionStart="Fig" captionStartId="5.[113,139,827,841]" captionTargetBox="[160,1444,182,798]" captionTargetId="figure-608@5.[159,1445,181,798]" captionTargetPageId="5" captionText="Fig. 5. Effect of ginsenosides on Pythium irregulare BR 1068 growth in vitro. Representative images of 90 mm petri dishes filled with Czapek Dox Mineral Agar containing 6 mm antibiotic-assay disks placed an equal distance from an 8 mm diameter plug of P. irregulare BR 1068 are portrayed. Disks on the left in each column are infused with 100 µL, (A) 10 mg mL–1 purified ginsenoside extract [GSF (G)], (B) 5 mg mL–1 ginsenoside Re (6), (C) 5 mg mL–1 ginsenoside Rb1 (1) or (D) 5 mg mL–1 ginsenoside F2 (4). All disks on the right in each column are controls infused with 100 µL MeOH. Experiment was performed in triplicate (N = 3). Images shown are representative of a single plate, for each treatment, monitored every 24 h for 120 h. Earlier (24 h) images are not included because mycelia had not reached filters and no effects were seen in any treatments. All ginsenosides were dissolved in MeOH. The apparent discolouration observed in the medium around the F2 (4) infused disks is attributed to ginsenosides that had leached into the agar medium from the disk after evaporation of the MeOH solvent. This effect is present in all other ginsenoside infused filters to varying degrees, and arises from manipulation of the lighting used to provide contrast between the mycelia and background medium." figureDoi="http://doi.org/10.5281/zenodo.10485553" httpUri="https://zenodo.org/record/10485553/files/figure.png" pageId="4" pageNumber="60">Fig. 5D</figureCitation>
;
<quantity id="3F7E9B567126FFC0FA964B20FA45F8E3" box="[1393,1464,1828,1847]" metricMagnitude="-7" metricUnit="kg" metricValue="5.0" pageId="4" pageNumber="60" unit="mg" value="0.5">0.5 mg</quantity>
). Within two days of growth, inhibition of mycelial growth towards the F2 (
<emphasis id="CAF2EAA17126FFC0FC944B58FC7DF8BB" bold="true" box="[883,896,1884,1903]" pageId="4" pageNumber="60">4</emphasis>
) infused assay disk was visually evident. While the inhibitory effect of F2 (
<emphasis id="CAF2EAA17126FFC0FC194B7CFBF6F85F" bold="true" box="[1022,1035,1912,1931]" pageId="4" pageNumber="60">4</emphasis>
) persisted throughout the five day time course, it did appear to diminish, as mycelia began to grow around the filter disk, almost completely surrounding it by the end of the experiment. However, mycelia were never seen to grow through the filter; neither was any enhanced aerial growth observed at any time during the time course.
</paragraph>
<paragraph id="F83936B37126FFC0FF914B90FA3EFE34" blockId="4.[87,757,1689,2015]" lastBlockId="4.[805,1475,182,481]" pageId="4" pageNumber="60">
Alternatively, the presence of relatively large doses of ginsenosides in the rhizosphere may have simply altered the growth pattern of the pathogen in the soil, as seen when
<taxonomicName id="3F864D307126FFC0FD834BCEFD21F80A" baseAuthorityName="BR" baseAuthorityYear="1068" box="[612,732,1994,2014]" class="Magnoliopsida" family="Araliaceae" genus="Pythium" kingdom="Plantae" order="Apiales" pageId="4" pageNumber="60" phylum="Tracheophyta" rank="species" species="irregulare">
<emphasis id="CAF2EAA17126FFC0FD834BCEFD21F80A" box="[612,732,1994,2014]" italics="true" pageId="4" pageNumber="60">P. irregulare</emphasis>
</taxonomicName>
is grown in medium supplemented with ginsenosides (
<bibRefCitation id="9C174B427126FFC0FAAB4CB2FCA4FF31" author="Nicol, R. W. &amp; Traquair, J. A. &amp; Bernards, M. A." pageId="4" pageNumber="60" pagination="557 - 562" refId="ref9534" refString="Nicol, R. W., Traquair, J. A., Bernards, M. A., 2002. Ginsenosides as host resistance factors in American ginseng (Panax quinquefolius). Can. J. Bot. 80, 557 - 562." type="journal article" year="2002">Nicol et al., 2002</bibRefCitation>
,
<bibRefCitation id="9C174B427126FFC0FC8F4CD6FC67FF31" author="Nicol, R. W. &amp; Yousef, L. &amp; Traquair, J. A. &amp; Bernards, M. A." box="[872,922,210,229]" pageId="4" pageNumber="60" pagination="257 - 264" refId="ref9582" refString="Nicol, R. W., Yousef, L., Traquair, J. A., Bernards, M. A., 2003. Ginsenosides stimulate the growth of soilborne pathogens of American ginseng. Phytochemistry 64, 257 - 264." type="journal article" year="2003">2003</bibRefCitation>
; see also below). Moreover,
<taxonomicName id="3F864D307126FFC0FB3C4CD5FAABFF31" authorityName="BR" authorityYear="1068" box="[1243,1366,209,229]" class="Magnoliopsida" family="Araliaceae" genus="Pythium" kingdom="Plantae" order="Apiales" pageId="4" pageNumber="60" phylum="Tracheophyta" rank="species" species="irregulare">
<emphasis id="CAF2EAA17126FFC0FB3C4CD5FAABFF31" box="[1243,1366,209,229]" italics="true" pageId="4" pageNumber="60">P. irregulare</emphasis>
</taxonomicName>
has been shown to directly interact with ginsenosides
<emphasis id="CAF2EAA17126FFC0FB004CE9FAD0FED5" box="[1255,1325,237,257]" italics="true" pageId="4" pageNumber="60">in vitro</emphasis>
, by deglycosylating them into ginsenoside F2 (
<emphasis id="CAF2EAA17126FFC0FB674D0EFB70FEC9" bold="true" box="[1152,1165,266,285]" pageId="4" pageNumber="60">4</emphasis>
) (
<bibRefCitation id="9C174B427126FFC0FB424D0EFA43FEC9" author="Yousef, L. F. &amp; Bernards, M. A." box="[1189,1470,266,285]" pageId="4" pageNumber="60" pagination="1740 - 1749" refId="ref10407" refString="Yousef, L. F., Bernards, M. A., 2006. In vitro metabolism of ginsenosides by the ginseng root pathogen Pythium irregulare. Phytochemistry 67, 1740 - 1749." type="journal article" year="2006">Yousef and Bernards, 2006</bibRefCitation>
;
<bibRefCitation id="9C174B427126FFC0FCC24D22FC0BFEED" author="Neculai, M. A. &amp; Ivanov, D. &amp; Bernards, M. A." box="[805,1014,294,313]" pageId="4" pageNumber="60" pagination="1948 - 1957" refId="ref9457" refString="Neculai, M. A., Ivanov, D., Bernards, M. A., 2009. Partial purification and characterization of three ginsenoside-metabolizing [beta] - glucosidases from Pythium irregulare. Phytochemistry 70, 1948 - 1957." type="journal article" year="2009">Neculai et al., 2009</bibRefCitation>
;
<bibRefCitation id="9C174B427126FFC0FBE24D22FAE2FEED" author="Ivanov, D. A. &amp; Bernards, M. A." box="[1029,1311,294,313]" pageId="4" pageNumber="60" pagination="44 - 53" refId="ref8841" refString="Ivanov, D. A., Bernards, M. A., 2012. Ginsenosidases and the pathogenicity of Pythium irregulare. Phytochemistry 78, 44 - 53." type="journal article" year="2012">Ivanov and Bernards, 2012</bibRefCitation>
). It is possible, therefore, that this interaction results in altered patterns of growth in the soil, which may have the secondary effect of delaying infection. Unfortunately, our attempts to directly assess the extent of
<taxonomicName id="3F864D307126FFC0FCC24D90FC64FE7C" authorityName="BR" authorityYear="1068" box="[805,921,404,424]" class="Magnoliopsida" family="Araliaceae" genus="Pythium" kingdom="Plantae" order="Apiales" pageId="4" pageNumber="60" phylum="Tracheophyta" rank="species" species="irregulare">
<emphasis id="CAF2EAA17126FFC0FCC24D90FC64FE7C" box="[805,921,404,424]" italics="true" pageId="4" pageNumber="60">P. irregulare</emphasis>
</taxonomicName>
growth in the pots, using ITS region nuclear ribosomal DNA markers (
<bibRefCitation id="9C174B427126FFC0FC2C4DB5FB4EFE10" author="Okubara, P. A. &amp; Schroeder, K. L. &amp; Paulitz, T. C." box="[971,1203,433,453]" pageId="4" pageNumber="60" pagination="300 - 313" refId="ref9628" refString="Okubara, P. A., Schroeder, K. L., Paulitz, T. C., 2005. Real-time polymerase chain reaction: applications to studies on soilborne pathogens. Can. J. Plant Pathol. 27, 300 - 313." type="journal article" year="2005">Okubara et al., 2005</bibRefCitation>
;
<bibRefCitation id="9C174B427126FFC0FB214DB5FA43FE10" author="Schroeder, K. L. &amp; Okubara, P. A. &amp; Tambong, J. T. &amp; Levesque, C. A. &amp; Paulitz, T. C." box="[1222,1470,433,453]" pageId="4" pageNumber="60" pagination="637 - 647" refId="ref9982" refString="Schroeder, K. L., Okubara, P. A., Tambong, J. T., Levesque, C. A., Paulitz, T. C., 2006. Identification and quantification of pathogenic Pythium spp. from soils in Eastern Washington using real-time polymerase chain reaction. Ecol. Epidemiol. 96, 637 - 647." type="journal article" year="2006">Schroeder et al., 2006</bibRefCitation>
;
<bibRefCitation id="9C174B427126FFC0FCC24DC9FBF7FE34" author="Kernaghan, G. &amp; Reeleder, R. D. &amp; Hoke, S. M. T." box="[805,1034,461,481]" pageId="4" pageNumber="60" pagination="508 - 516" refId="ref8930" refString="Kernaghan, G., Reeleder, R. D., Hoke, S. M. T., 2007. Quantification of Cylindrocarpon destructans f. sp. panacis in soils by real-time PCR. Plant Pathol. 56, 508 - 516." type="journal article" year="2007">Kernaghan et al., 2007</bibRefCitation>
;
<bibRefCitation id="9C174B427126FFC0FBF34DC9FB08FE34" author="Kernaghan, G. &amp; Reeleder, R. D. &amp; Hoke, S. M. T." box="[1044,1269,461,481]" pageId="4" pageNumber="60" pagination="447 - 455" refId="ref8977" refString="Kernaghan, G., Reeleder, R. D., Hoke, S. M. T., 2008. Quantification of Pythium populations in ginseng soils by culture dependent and real-time PCR methods. Appl. Soil Ecol. 40, 447 - 455." type="journal article" year="2008">Kernaghan et al., 2008</bibRefCitation>
) were unsuccessful.
</paragraph>
<caption id="ACF9663B7127FFC1FF964F3FFC57FC3D" ID-DOI="http://doi.org/10.5281/zenodo.10485553" ID-Zenodo-Dep="10485553" httpUri="https://zenodo.org/record/10485553/files/figure.png" pageId="5" pageNumber="61" startId="5.[113,139,827,841]" targetBox="[160,1444,182,798]" targetPageId="5" targetType="figure">
<paragraph id="F83936B37127FFC1FF964F3FFC57FC3D" blockId="5.[113,1502,826,1002]" pageId="5" pageNumber="61">
<emphasis id="CAF2EAA17127FFC1FF964F3FFF59FC9D" bold="true" box="[113,164,827,841]" pageId="5" pageNumber="61">Fig. 5.</emphasis>
Effect of ginsenosides on
<taxonomicName id="3F864D307127FFC1FE9B4F3EFDA5FC9D" authority="BR 1068" authorityName="BR" authorityYear="1068" box="[380,600,826,842]" class="Magnoliopsida" family="Araliaceae" genus="Pythium" kingdom="Plantae" order="Apiales" pageId="5" pageNumber="61" phylum="Tracheophyta" rank="species" species="irregulare">
<emphasis id="CAF2EAA17127FFC1FE9B4F3EFDF3FC9D" box="[380,526,826,841]" italics="true" pageId="5" pageNumber="61">Pythium irregulare</emphasis>
BR 1068
</taxonomicName>
growth
<emphasis id="CAF2EAA17127FFC1FD794F3EFD2AFC9D" box="[670,727,826,841]" italics="true" pageId="5" pageNumber="61">in vitro</emphasis>
. Representative images of 90 mm petri dishes filled with Czapek Dox Mineral Agar containing 6 mm antibiotic-assay disks placed an equal distance from an 8 mm diameter plug of
<taxonomicName id="3F864D307127FFC1FCCD4F55FC2CFCB4" authority="BR 1068" authorityName="BR" authorityYear="1068" box="[810,977,849,864]" class="Magnoliopsida" family="Araliaceae" genus="Pythium" kingdom="Plantae" order="Apiales" pageId="5" pageNumber="61" phylum="Tracheophyta" rank="species" species="irregulare">
<emphasis id="CAF2EAA17127FFC1FCCD4F55FC75FCB4" box="[810,904,849,864]" italics="true" pageId="5" pageNumber="61">P. irregulare</emphasis>
BR 1068
</taxonomicName>
are portrayed. Disks on the left in each column are infused with 100 µL, (A) 10 mg mL
<superScript id="0FF39BFB7127FFC1FEFA4F61FED3FCBB" attach="left" box="[285,302,869,879]" fontSize="4" pageId="5" pageNumber="61">–1</superScript>
purified ginsenoside extract [GSF (G)], (B) 5 mg mL
<superScript id="0FF39BFB7127FFC1FD2A4F61FD23FCBB" attach="left" box="[717,734,869,879]" fontSize="4" pageId="5" pageNumber="61">–1</superScript>
ginsenoside Re (
<emphasis id="CAF2EAA17127FFC1FC814F6DFC8DFCA3" bold="true" box="[870,880,873,887]" pageId="5" pageNumber="61">6</emphasis>
), (C) 5 mg mL
<superScript id="0FF39BFB7127FFC1FC064F61FC0FFCBB" attach="left" box="[993,1010,869,879]" fontSize="4" pageId="5" pageNumber="61">–1</superScript>
ginsenoside Rb1 (
<emphasis id="CAF2EAA17127FFC1FB624F6DFB72FCA3" bold="true" box="[1157,1167,873,887]" pageId="5" pageNumber="61">1</emphasis>
) or (D) 5 mg mL
<superScript id="0FF39BFB7127FFC1FAF34F61FAD8FCBB" attach="left" box="[1300,1317,869,879]" fontSize="4" pageId="5" pageNumber="61">–1</superScript>
ginsenoside F2 (
<emphasis id="CAF2EAA17127FFC1FA4A4F6DFA4AFCA3" bold="true" box="[1453,1463,873,887]" pageId="5" pageNumber="61">4</emphasis>
). All disks on the right in each column are controls infused with 100 µL MeOH. Experiment was performed in triplicate (
<emphasis id="CAF2EAA17127FFC1FBF54F7BFBE3FC5A" box="[1042,1054,895,910]" italics="true" pageId="5" pageNumber="61">N</emphasis>
= 3). Images shown are representative of a single plate, for each treatment, monitored every 24 h for 120 h. Earlier (24 h) images are not included because mycelia had not reached filters and no effects were seen in any treatments. All ginsenosides were dissolved in MeOH. The apparent discolouration observed in the medium around the F2 (
<emphasis id="CAF2EAA17127FFC1FC0A4FA9FC0AFC6F" bold="true" box="[1005,1015,941,955]" pageId="5" pageNumber="61">4</emphasis>
) infused disks is attributed to ginsenosides that had leached into the agar medium from the disk after evaporation of the MeOH solvent. This effect is present in all other ginsenoside infused filters to varying degrees, and arises from manipulation of the lighting used to provide contrast between the mycelia and background medium.
</paragraph>
</caption>
<paragraph id="F83936B37127FFC1FF76486FFEFEFAF1" blockId="5.[113,784,1075,2015]" pageId="5" pageNumber="61">
The effects of all ginsenoside treatments on mycelial growth, except ginsenoside Re (
<emphasis id="CAF2EAA17127FFC1FE814882FE8EFB4D" bold="true" box="[358,371,1158,1177]" pageId="5" pageNumber="61">6</emphasis>
), were dose dependent (see Supporting Information
<figureCitation id="60BD2A367127FFC1FF1E48A6FE98FB62" box="[249,357,1186,1206]" captionStart="Fig" captionStartId="1.[113,139,1496,1510]" captionTargetBox="[386,1217,187,1452]" captionTargetId="figure-286@1.[378,1227,181,1467]" captionTargetPageId="1" captionText="Fig. 1. Ginsenosides of Panax quinquefolius L.Ginsenosides Rb1 (1), Rd (2), gypenoside XVII (GXVII, 3) and F2 (4) are 20(S)-protopanaxadiols. Ginsenosides Rg1 (5) and Re (6) are 20(S)-protopanaxatriols. The core structure of ginsenoside F2 (4) is highlighted in BOLD in the structures of Ginsenosides Rb1 (1), Rd (2), gypenoside XVII (GXVII,3) and F2 (4), emphasizing that selective ss(1–6) and ss(1–2) glucosidase metabolism of sugar residues from 20(S)-protopanaxadiol ginsenosides can lead to formation of F2 (4)." figureDoi="http://doi.org/10.5281/zenodo.10485545" httpUri="https://zenodo.org/record/10485545/files/figure.png" pageId="5" pageNumber="61">Figs. S1–S</figureCitation>
<figureCitation id="60BD2A367127FFC1FE8248A7FE8FFB62" box="[357,370,1187,1206]" captionStart="Fig" captionStartId="4.[87,113,1333,1347]" captionTargetBox="[96,747,181,1304]" captionTargetId="figure-808@4.[96,747,181,1304]" captionTargetPageId="4" captionText="Fig. 4. Time course of ginseng infection by Pythium irregulare. Ginseng plants were soaked in either ddH O or 10 mg mL–1 (ddH O) purified ginsenoside extract and 2 2 planted 8 cm apart. Uninoculated plants: ΦNO values for (A) one- and (B) two year old plants treated with 10 mg mL –1 (dissolved in ddH O) purified ginsenosides 2 extract (filled circles) were collected every day for 15 days (one-year old) and for 20 days (two-year old). Inoculated plants: A 13 mm column of sand was removed 4 cm away from the base of each plant, in the middle of the pot, and filled with a Pythium irregulare (BR 1068)-sand mix inoculum. ΦNO values for (A) one- and (B) two year old plants treated with ddH O (open circles) or 10 mg mL–1 (dissolved in 2 ddH2O) purified ginsenosides extract (open triangles) were collected every day for 15 days (one-year old) and for 20 days (two-year old). All ΦNO values are expressed as Mean ± SE. N = 6 for one-year old plant treatments; N = 4 for one-year old control; N = 8 for two-year old plant treatments and control." figureDoi="http://doi.org/10.5281/zenodo.10485551" httpUri="https://zenodo.org/record/10485551/files/figure.png" pageId="5" pageNumber="61">4</figureCitation>
). That is, enhanced mycelial growth increased with ginsenoside concentration around filters treated with GSF and ginsenoside Rb1 (
<emphasis id="CAF2EAA17127FFC1FE2348DEFE2CFB39" bold="true" box="[452,465,1242,1261]" pageId="5" pageNumber="61">1</emphasis>
), while the zone of inhibition around disks infused with ginsenoside F2 (
<emphasis id="CAF2EAA17127FFC1FDB748F2FDA0FADD" bold="true" box="[592,605,1270,1289]" pageId="5" pageNumber="61">4</emphasis>
) increased with concentration.
</paragraph>
<paragraph id="F83936B37127FFC1FF76492AFE69F9A0" blockId="5.[113,784,1075,2015]" pageId="5" pageNumber="61">
It is not clear from these
<emphasis id="CAF2EAA17127FFC1FE784929FE15FA95" box="[415,488,1325,1345]" italics="true" pageId="5" pageNumber="61">in vitro</emphasis>
data whether the enhanced growth of
<taxonomicName id="3F864D307127FFC1FF05494DFEA6FA89" authorityName="BR" authorityYear="1068" box="[226,347,1353,1373]" class="Magnoliopsida" family="Araliaceae" genus="Pythium" kingdom="Plantae" order="Apiales" pageId="5" pageNumber="61" phylum="Tracheophyta" rank="species" species="irregulare">
<emphasis id="CAF2EAA17127FFC1FF05494DFEA6FA89" box="[226,347,1353,1373]" italics="true" pageId="5" pageNumber="61">P. irregulare</emphasis>
</taxonomicName>
in the presence of protopanaxadiols was due to ginsenosides acting as chemoattractants or growth factors. Furthermore, while ginsenoside Rb1 (
<emphasis id="CAF2EAA17127FFC1FDE24985FDEFFA40" bold="true" box="[517,530,1409,1428]" pageId="5" pageNumber="61">1</emphasis>
) appeared to stimulate pathogen growth on its own, the effect of the ginsenoside mixture was greater and correlated to the increased concentration of ginsenosides applied. Ginsenoside Rb1 (
<emphasis id="CAF2EAA17127FFC1FE0949D1FE06FA3C" bold="true" box="[494,507,1493,1512]" pageId="5" pageNumber="61">1</emphasis>
) applied in the same concentration as the ginsenoside mixture showed an equivalent level of growth stimulation (Supporting Information
<figureCitation id="60BD2A367127FFC1FD8E4A09FCFCF9F4" box="[617,769,1549,1568]" captionStart="Fig" captionStartId="4.[87,113,1333,1347]" captionTargetBox="[96,747,181,1304]" captionTargetId="figure-808@4.[96,747,181,1304]" captionTargetPageId="4" captionText="Fig. 4. Time course of ginseng infection by Pythium irregulare. Ginseng plants were soaked in either ddH O or 10 mg mL–1 (ddH O) purified ginsenoside extract and 2 2 planted 8 cm apart. Uninoculated plants: ΦNO values for (A) one- and (B) two year old plants treated with 10 mg mL –1 (dissolved in ddH O) purified ginsenosides 2 extract (filled circles) were collected every day for 15 days (one-year old) and for 20 days (two-year old). Inoculated plants: A 13 mm column of sand was removed 4 cm away from the base of each plant, in the middle of the pot, and filled with a Pythium irregulare (BR 1068)-sand mix inoculum. ΦNO values for (A) one- and (B) two year old plants treated with ddH O (open circles) or 10 mg mL–1 (dissolved in 2 ddH2O) purified ginsenosides extract (open triangles) were collected every day for 15 days (one-year old) and for 20 days (two-year old). All ΦNO values are expressed as Mean ± SE. N = 6 for one-year old plant treatments; N = 4 for one-year old control; N = 8 for two-year old plant treatments and control." figureDoi="http://doi.org/10.5281/zenodo.10485551" httpUri="https://zenodo.org/record/10485551/files/figure.png" pageId="5" pageNumber="61">
Figs.
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–
<geoCoordinate id="9DB250747127FFC1FD304A09FCFCF9F4" box="[727,769,1549,1568]" degrees="4.4" direction="south" orientation="latitude" pageId="5" pageNumber="61" precision="5555" value="-4.4">S4.4</geoCoordinate>
</figureCitation>
). Therefore, while ginsenoside Rb1 (
<emphasis id="CAF2EAA17127FFC1FE074A2DFE10F9E8" bold="true" box="[480,493,1577,1596]" pageId="5" pageNumber="61">1</emphasis>
) appears to be involved in the growth stimulation of
<taxonomicName id="3F864D307127FFC1FE754A40FDF0F98C" authorityName="BR" authorityYear="1068" box="[402,525,1604,1624]" class="Magnoliopsida" family="Araliaceae" genus="Pythium" kingdom="Plantae" order="Apiales" pageId="5" pageNumber="61" phylum="Tracheophyta" rank="species" species="irregulare">
<emphasis id="CAF2EAA17127FFC1FE754A40FDF0F98C" box="[402,525,1604,1624]" italics="true" pageId="5" pageNumber="61">P. irregulare</emphasis>
</taxonomicName>
, it is not the only ginsenoside that has this effect.
</paragraph>
<paragraph id="F83936B37127FFC1FF764A79FC02FADD" blockId="5.[113,784,1075,2015]" lastBlockId="5.[831,1501,1074,1289]" pageId="5" pageNumber="61">
Based on these data, it remains unclear why
<taxonomicName id="3F864D307127FFC1FDA94A78FD3EF944" authorityName="BR" authorityYear="1068" box="[590,707,1660,1680]" class="Magnoliopsida" family="Araliaceae" genus="Pythium" kingdom="Plantae" order="Apiales" pageId="5" pageNumber="61" phylum="Tracheophyta" rank="species" species="irregulare">
<emphasis id="CAF2EAA17127FFC1FDA94A78FD3EF944" box="[590,707,1660,1680]" italics="true" pageId="5" pageNumber="61">P. irregulare</emphasis>
</taxonomicName>
invests the energy to convert the common ginsenosides (e.g., Rb1 (
<emphasis id="CAF2EAA17127FFC1FD344A9DFD1DF978" bold="true" box="[723,736,1689,1708]" pageId="5" pageNumber="61">1</emphasis>
), Rd (
<emphasis id="CAF2EAA17127FFC1FF9D4AB0FF7AF913" bold="true" box="[122,135,1716,1735]" pageId="5" pageNumber="61">2</emphasis>
), gypenoside XIIV (
<emphasis id="CAF2EAA17127FFC1FEB64AB0FEA3F913" bold="true" box="[337,350,1716,1735]" pageId="5" pageNumber="61">3</emphasis>
)) into ginsenoside F2 (
<emphasis id="CAF2EAA17127FFC1FDAA4AB0FDA7F913" bold="true" box="[589,602,1716,1735]" pageId="5" pageNumber="61">4</emphasis>
) via extracellular glycosidases (
<bibRefCitation id="9C174B427127FFC1FF1B4AD4FDF5F937" author="Yousef, L. F. &amp; Bernards, M. A." box="[252,520,1744,1763]" pageId="5" pageNumber="61" pagination="1740 - 1749" refId="ref10407" refString="Yousef, L. F., Bernards, M. A., 2006. In vitro metabolism of ginsenosides by the ginseng root pathogen Pythium irregulare. Phytochemistry 67, 1740 - 1749." type="journal article" year="2006">Yousef and Bernards, 2006</bibRefCitation>
), when in fact the growth of this organism is visibly inhibited by the pure F2 (
<emphasis id="CAF2EAA17127FFC1FD964AE8FD83F92B" bold="true" box="[625,638,1772,1791]" pageId="5" pageNumber="61">4</emphasis>
) compound. It is possible that the presence of common ginsenosides stimulates a change in
<taxonomicName id="3F864D307127FFC1FF074B27FEA4F8E3" authorityName="BR" authorityYear="1068" box="[224,345,1827,1847]" class="Magnoliopsida" family="Araliaceae" genus="Pythium" kingdom="Plantae" order="Apiales" pageId="5" pageNumber="61" phylum="Tracheophyta" rank="species" species="irregulare">
<emphasis id="CAF2EAA17127FFC1FF074B27FEA4F8E3" box="[224,345,1827,1847]" italics="true" pageId="5" pageNumber="61">P. irregulare</emphasis>
</taxonomicName>
metabolism, leading to the expression of genes encoding extracellular glycosidases, as well as genes associated with the utilization of F2 (
<emphasis id="CAF2EAA17127FFC1FE564B58FE43F8BB" bold="true" box="[433,446,1884,1903]" pageId="5" pageNumber="61">4</emphasis>
) (e.g., as a growth stimulant). By contrast, direct exposure to pure F2 (
<emphasis id="CAF2EAA17127FFC1FE004B7CFE09F85F" bold="true" box="[487,500,1912,1931]" pageId="5" pageNumber="61">4</emphasis>
) may circumvent the metabolic shift needed to utilize this compound and avoid the potential deleterious effects it can have on growth. Regardless, the impact of GSF and Rb1 on
<taxonomicName id="3F864D307127FFC1FEF44BCEFE7AF80A" authorityName="BR" authorityYear="1068" box="[275,391,1994,2014]" class="Magnoliopsida" family="Araliaceae" genus="Pythium" kingdom="Plantae" order="Apiales" pageId="5" pageNumber="61" phylum="Tracheophyta" rank="species" species="irregulare">
<emphasis id="CAF2EAA17127FFC1FEF44BCEFE7AF80A" box="[275,391,1994,2014]" italics="true" pageId="5" pageNumber="61">P. irregulare</emphasis>
</taxonomicName>
growth habit in the
<emphasis id="CAF2EAA17127FFC1FDB44BCEFD65F80A" box="[595,664,1994,2014]" italics="true" pageId="5" pageNumber="61">in vitro</emphasis>
assays may inform our observations in the
<emphasis id="CAF2EAA17127FFC1FB914836FB4BFB92" box="[1142,1206,1074,1094]" italics="true" pageId="5" pageNumber="61">in vivo</emphasis>
assays. That is, when hyphae of
<taxonomicName id="3F864D307127FFC1FCB0484AFC36FBB6" authorityName="BR" authorityYear="1068" box="[855,971,1102,1122]" class="Magnoliopsida" family="Araliaceae" genus="Pythium" kingdom="Plantae" order="Apiales" pageId="5" pageNumber="61" phylum="Tracheophyta" rank="species" species="irregulare">
<emphasis id="CAF2EAA17127FFC1FCB0484AFC36FBB6" box="[855,971,1102,1122]" italics="true" pageId="5" pageNumber="61">P. irregulare</emphasis>
</taxonomicName>
encounter ginsenosides in the soil surrounding roots of ginseng, it is likely that forward progress is temporarily halted while the organism metabolizes these compounds, building up hyphal mass in the process. Such a change in growth habit would explain the apparent delay in infection of ginsenoside-treated roots, which would have a higher concentration of ginsenosides in the rhizosphere.
</paragraph>
</subSubSection>
</treatment>
</document>
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