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<mods:title id="23D08BFDE38CD5C5C801F019CF0D1D20">What pollinators see does not match what they smell: Absence of color-fragrance association in the deceptive orchid Ionopsis utricularioides</mods:title>
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<mods:namePart id="C22C4DAC54CBAA2D212B9374E824C077">Aguiar, João Marcelo Robazzi Bignelli Valente</mods:namePart>
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<mods:namePart id="5768A460DE908673B0951B3B918784EF">Ferreira, Gabriel de Souza</mods:namePart>
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<mods:namePart id="D0E2C888B9341CBFE159F52EAC1076A3">Sanches, Patricia Alessandra</mods:namePart>
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2.1. What is the scent composition of
<taxonomicName id="97D28CE26B50FB1BFB07DEC1D103FA33" ID-CoL="6N8VC" ID-ENA="154687" authority="(Sw.) Lindl." authorityName="Lindl." baseAuthorityName="Sw." box="[1164,1312,1516,1536]" class="Liliopsida" family="Orchidaceae" genus="Ionopsis" kingdom="Plantae" order="Asparagales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="species" species="utricularioides">I. utricularioides</taxonomicName>
?
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Eighteen different volatile components were found in the flowers of
<taxonomicName id="97D28CE26B50FB1BFCB9DD6CD7E0F99F" box="[818,963,1600,1619]" class="Liliopsida" family="Orchidaceae" genus="Ionopsis" kingdom="Plantae" order="Asparagales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="species" species="utricularioides">
<emphasis id="62A62B736B50FB1BFCB9DD6CD7E0F99F" bold="true" box="[818,963,1600,1619]" italics="true" pageId="1" pageNumber="2">I. utricularioides</emphasis>
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. Floral volatile composition and concentrations of each compound varied between individuals (
<tableCitation id="1D50C2DA6B50FB1BFB3ADD70D0DAF9A3" box="[1201,1273,1628,1647]" captionStart="Table 1" captionStartId="2.[139,189,394,410]" captionTargetBox="[155,1952,482,1172]" captionTargetPageId="2" captionText="Table 1 Floral fragrance profile and color saturation according to bees visual discrimination of the orchid, Ionopsis utricularioides individuals. Values are ng.μl 1 per flower for the emission of floral compounds , and hexagon model units for floral color saturation. Average values are mean ±standard error. RI designates retention index for the compounds." pageId="1" pageNumber="2">Table 1</tableCitation>
), as also recorded for several other deceptive orchids (Delle-Vedove et al., 2011;
<bibRefCitation id="34438A906B50FB1BFAE8DD54D7A0F96B" author="Dormont, L. &amp; Delle-Vedove, R. &amp; Schatz, B." pageId="1" pageNumber="2" pagination="51 - 59" refId="ref9693" refString="Dormont, L., Delle-Vedove, R., Bessi`ere, J. M., Schatz, B., 2014. Floral scent emitted by white and coloured morphs in orchids. Phytochemistry 100, 51 - 59. https: // doi. org / 10.1016 / j. phytochem. 2014.01.009." type="journal article" year="2014">Dormont et al, 2014</bibRefCitation>
,
<bibRefCitation id="34438A906B50FB1BFC1FDDB8D7E7F96B" author="Dormont, L. &amp; Joffard, N. &amp; Schatz, B." box="[916,964,1684,1703]" pageId="1" pageNumber="2" pagination="1036 - 1058" refId="ref9750" refString="Dormont, L., Joffard, N., Schatz, B., 2019. Intraspecific variation in floral color and odor in orchids. Int. J. Plant Sci. 180, 1036 - 1058. https: // doi. org / 10.1086 / 705589." type="journal article" year="2019">2019</bibRefCitation>
;
<bibRefCitation id="34438A906B50FB1BFC5EDDB8D0F7F96B" author="Juillet, N. &amp; Scopece, G." box="[981,1236,1684,1703]" pageId="1" pageNumber="2" pagination="317 - 322" refId="ref10557" refString="Juillet, N., Scopece, G., 2010. Does floral trait variability enhance reproductive success in deceptive orchids? Perspect. Plant Ecol. Evol. Syst. 12, 317 - 322. https: // doi. org / 10.1016 / j. ppees. 2010.05.001." type="journal article" year="2010">Juillet and Scopece, 2010</bibRefCitation>
;
<bibRefCitation id="34438A906B50FB1BFB6EDDB8D1EAF96B" author="Salzmann, C. C. &amp; Cozzolino, S. &amp; Schiestl, F. P." box="[1253,1481,1684,1703]" pageId="1" pageNumber="2" pagination="720 - 729" refId="ref12783" refString="Salzmann, C. C., Cozzolino, S., Schiestl, F. P., 2007 a. Floral scent in food-deceptive orchids: species specificity and sources of variability. Plant Biol. 9, 720 - 729. https: // doi. org / 10.1055 / s- 2007 - 965614." type="journal article" year="2007">Salzmann et al., 2007a</bibRefCitation>
,
<bibRefCitation id="34438A906B50FB1BFCB9DD83D74DF90E" author="Salzmann, C. C. &amp; Nardella, A. M. &amp; Cozzolino, S. &amp; Schiestl, F. P." box="[818,878,1711,1730]" pageId="1" pageNumber="2" pagination="757 - 765" refId="ref12841" refString="Salzmann, C. C., Nardella, A. M., Cozzolino, S., Schiestl, F. P., 2007 b. Variability in floral scent in rewarding and deceptive orchids: the signature of pollinator-imposed selection? Ann. Bot. 100, 757 - 765. https: // doi. org / 10.1093 / aob / mcm 161." type="journal article" year="2007">2007b</bibRefCitation>
;
<bibRefCitation id="34438A906B50FB1BFC01DD83D0EAF90F" author="Salzmann, C. C. &amp; Schiestl, F. P." box="[906,1225,1711,1731]" pageId="1" pageNumber="2" pagination="37 - 45" refId="ref12727" refString="Salzmann, C. C., Schiestl, F. P., 2007. Odour and colour polymorphism in the food-deceptive orchid Dactylorhiza romana. Plant Systemat. Evol. 267, 37 - 45. https: // doi. org / 10.1007 / s 00606 - 007 - 0560 - z." type="journal article" year="2007">Salzmann and Schiestl, 2007</bibRefCitation>
). As expected by the bee-pollination deceptive mechanism of
<taxonomicName id="97D28CE26B50FB1BFB36DDE0D7EBF936" authority="(Aguiar and Pansarin, 2019)" baseAuthorityName="Aguiar and Pansarin" baseAuthorityYear="2019" class="Liliopsida" family="Orchidaceae" genus="Ionopsis" kingdom="Plantae" order="Asparagales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="species" species="utricularioides">
<emphasis id="62A62B736B50FB1BFB36DDE0D171F912" bold="true" box="[1213,1362,1739,1759]" italics="true" pageId="1" pageNumber="2">I. utricularioides</emphasis>
(
<bibRefCitation id="34438A906B50FB1BFAE8DDE7D79CF936" author="Aguiar, J. M. R. B. V. &amp; Pansarin, E. R." pageId="1" pageNumber="2" pagination="72 - 78" refId="ref8875" refString="Aguiar, J. M. R. B. V., Pansarin, E. R., 2019. Deceptive pollination of Ionopsis utricularioides (Oncidiinae: orchidaceae). Flora 250, 72 - 78. https: // doi. org / 10.1016 / j. flora. 2018.11.018." type="journal article" year="2019">Aguiar and Pansarin, 2019</bibRefCitation>
)
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, many compounds in the orchid s fragrance are known to be important in pollinators attraction in other bee-pollinated plants, such as phenylacetaldehyde (
<bibRefCitation id="34438A906B50FB1BFBC6DC33D169F8FE" author="Knauer, A. C. &amp; Schiestl, F. P." box="[1101,1354,1823,1842]" pageId="1" pageNumber="2" pagination="135 - 143" refId="ref10868" refString="Knauer, A. C., Schiestl, F. P., 2015. Bees use honest floral signals as indicators of reward when visiting flowers. Ecol. Lett. 18, 135 - 143. https: // doi. org / 10.1111 / ele. 12386." type="journal article" year="2015">Knauer and Schiestl, 2015</bibRefCitation>
; Theis, 2006; Waelti et al., 2008) and limonene, one of the most common volatiles emitted by flowers (
<bibRefCitation id="34438A906B50FB1BFC74DC7BD0E3F8A6" author="Benelli, G. &amp; Canale, A. &amp; Romano, D. &amp; Flamini, G. &amp; Tavarini, S. &amp; Martini, A. &amp; Ascrizzi, R. &amp; Conte, G. &amp; Mele, M. &amp; Angelini, L. G." box="[1023,1216,1879,1898]" pageId="1" pageNumber="2" pagination="381 - 388" refId="ref9146" refString="Benelli, G., Canale, A., Romano, D., Flamini, G., Tavarini, S., Martini, A., Ascrizzi, R., Conte, G., Mele, M., Angelini, L. G., 2017. Flower scent bouquet variation and bee pollinator visits in Stevia rebaudiana Bertoni (Asteraceae), a source of natural sweeteners. Arthropod. Plant. Interact. 11, 381 - 388. https: // doi. org / 10.1007 / s 11829 - 016 - 9488 - y." type="journal article" year="2017">Benelli et al., 2017</bibRefCitation>
;
<bibRefCitation id="34438A906B50FB1BFB5ADC7BD1EAF8A6" author="Erasto, P. &amp; Viljoen, A. M." box="[1233,1481,1879,1898]" pageId="1" pageNumber="2" pagination="1193 - 1202" refId="ref9872" refString="Erasto, P., Viljoen, A. M., 2008. Limonene - a review: biosynthetic, ecological and pharmacological relevance. Nat. Prod. Commun. 3, 1193 - 1202. https: // doi. org / 10.1177 / 1934578 x 0800300728." type="journal article" year="2008">Erasto and Viljoen, 2008</bibRefCitation>
;
<bibRefCitation id="34438A906B50FB1BFCB9DC5FD7DFF84A" author="Knudsen, J. T. &amp; Tollsten, L." box="[818,1020,1907,1926]" pageId="1" pageNumber="2" pagination="253 - 280" refId="ref10920" refString="Knudsen, J. T., Tollsten, L., Bergstr ¨ om, L. G., 1993. Floral scents-a checklist of volatile compounds isolated by head-space techniques. Phytochemistry 33, 253 - 280. https: // doi. org / 10.1016 / 0031 - 9422 (93) 85502 - I." type="journal article" year="1993">Knudsen et al., 1993</bibRefCitation>
). β- ocimene, the major compound found in the fragrance of
<taxonomicName id="97D28CE26B50FB1BFC20DCA3D01DF86D" baseAuthorityName="Aguiar and Pansarin" baseAuthorityYear="2019" box="[939,1086,1934,1954]" class="Liliopsida" family="Orchidaceae" genus="Ionopsis" kingdom="Plantae" order="Asparagales" pageId="1" pageNumber="2" phylum="Tracheophyta" rank="species" species="utricularioides">
<emphasis id="62A62B736B50FB1BFC20DCA3D01DF86D" bold="true" box="[939,1086,1934,1954]" italics="true" pageId="1" pageNumber="2">I. utricularioides</emphasis>
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, with an average emission of 52.6 ±
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μl
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(mean ± S.E.) and occurring in 80% of the sampled individuals (
<tableCitation id="1D50C2DA6B52FB19FFE7DBB8D497FF6B" box="[108,180,148,167]" captionStart="Table 1" captionStartId="2.[139,189,394,410]" captionTargetBox="[155,1952,482,1172]" captionTargetPageId="2" captionText="Table 1 Floral fragrance profile and color saturation according to bees visual discrimination of the orchid, Ionopsis utricularioides individuals. Values are ng.μl 1 per flower for the emission of floral compounds , and hexagon model units for floral color saturation. Average values are mean ±standard error. RI designates retention index for the compounds." pageId="3" pageNumber="4">Table 1</tableCitation>
), is also very common in rewarding flowers pollinated by insects (Farr´e-Armengol et al., 2017;
<bibRefCitation id="34438A906B52FB19FD89DB9CD6D7FF0F" author="Knudsen, J. T. &amp; Eriksson, R. &amp; Gershenzon, J. &amp; Stahl, B." box="[514,756,176,195]" pageId="3" pageNumber="4" pagination="1 - 120" refId="ref10980" refString="Knudsen, J. T., Eriksson, R., Gershenzon, J., Stahl, B., 2006. Diversity and distribution of floral scent. Bot. Rev. 72, 1 - 120. https: // doi. org / 10.1663 / 0006 - 8101 (2006) 72 [1: dadofs] 2.0. co; 2." type="journal article" year="2006">Knudsen et al., 2006</bibRefCitation>
). Farr´e-Armengol et al. (2017) proposed an interesting hypothesis that, given its recurrence and abundance in floral fragrances, β- ocimene would act as a general global floral signal to many plants in a community. In this context, other minor compounds could ensure the scent specificity needed for pollination (Farr´e-Armengol et al., 2017), as pollinators, especially bees, can detect minor changes in floral fragrance composition (
<bibRefCitation id="34438A906B52FB19FF63DA5FD58CFE4A" author="Locatelli, F. F. &amp; Fernandez, P. C. &amp; Smith, B. H." box="[232,431,371,390]" pageId="3" pageNumber="4" pagination="2752 - 2762" refId="ref11396" refString="Locatelli, F. F., Fernandez, P. C., Smith, B. H., 2016. Learning about natural variation of odor mixtures enhances categorization in early olfactory processing. J. Exp. Biol. 219, 2752 - 2762. https: // doi. org / 10.1242 / jeb. 141465." type="journal article" year="2016">Locatelli et al., 2016</bibRefCitation>
;
<bibRefCitation id="34438A906B52FB19FE36DA5FD61CFE4A" author="Raguso, R. A." box="[445,575,371,390]" pageId="3" pageNumber="4" pagination="549 - 569" refId="ref12549" refString="Raguso, R. A., 2008. Wake up and Smell the Roses: the ecology and evolution of floral scent. Annu. Rev. Ecol. Evol. Syst. 39, 549 - 569. https: // doi. org / 10.1146 / annurev. ecolsys. 38.091206.095601." type="journal article" year="2008">Raguso, 2008</bibRefCitation>
). In the generalized food deceptive pollination context of
<taxonomicName id="97D28CE26B52FB19FE43DAA3D4FBFE72" authority="(Aguiar and Pansarin, 2019)" baseAuthorityName="Aguiar and Pansarin" baseAuthorityYear="2019" class="Liliopsida" family="Orchidaceae" genus="Ionopsis" kingdom="Plantae" order="Asparagales" pageId="3" pageNumber="4" phylum="Tracheophyta" rank="species" species="utricularioides">
<emphasis id="62A62B736B52FB19FE43DAA3D679FE6D" bold="true" box="[456,602,398,418]" italics="true" pageId="3" pageNumber="4">I. utricularioides</emphasis>
(
<bibRefCitation id="34438A906B52FB19FDE3DAA3D4ECFE72" author="Aguiar, J. M. R. B. V. &amp; Pansarin, E. R." pageId="3" pageNumber="4" pagination="72 - 78" refId="ref8875" refString="Aguiar, J. M. R. B. V., Pansarin, E. R., 2019. Deceptive pollination of Ionopsis utricularioides (Oncidiinae: orchidaceae). Flora 250, 72 - 78. https: // doi. org / 10.1016 / j. flora. 2018.11.018." type="journal article" year="2019">Aguiar and Pansarin, 2019</bibRefCitation>
)
</taxonomicName>
, β- ocimene could, thus, act as a global sign to attract pollinators, and bees could also perceive the variability derived from the other minor compounds of this orchid s scent. However, further behavioral or electrophysiological studies are needed to corroborate this hypothesis.
</paragraph>
</subSubSection>
</treatment>
</document>