{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T20:32:02Z","timestamp":1790195522041,"version":"4.1.0"},"reference-count":63,"publisher":"MDPI AG","issue":"2","license":[{"start":{"date-parts":[[2022,1,9]],"date-time":"2022-01-09T00:00:00Z","timestamp":1641686400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["31770752"],"award-info":[{"award-number":["31770752"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Jiangsu Science and Technology Support Program","award":["BY2015006\u201301"],"award-info":[{"award-number":["BY2015006\u201301"]}]},{"name":"the 333 Projects of Jiangsu Province","award":["BRA2018065"],"award-info":[{"award-number":["BRA2018065"]}]},{"name":"Natural Science Foundation for Universities of Jiangsu Province","award":["20KJB220006"],"award-info":[{"award-number":["20KJB220006"]}]},{"name":"Project of the 13th Five-Year Plan of Jiangsu Open University","award":["2020-D-01"],"award-info":[{"award-number":["2020-D-01"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Molecules"],"abstract":"<jats:p>Nymphaea hybrid, a precious water lily, is a widely-cultivated aquatic flower with high ornamental, economic, medicinal, and ecological value; it blooms recurrently and emits a strong fragrance. In the present study, in order to understand the volatile components of N. hybrid and its relationship with petals structure characteristics, the morphologies and anatomical structures of the flower petals of N. hybrid were investigated, and volatile compounds emitted from the petals were identified. Scanning and transmission electron microscopy were used to describe petal structures, and the volatile constituents were collected using headspace solid-phase microextraction (HS-SPME) fibers and analyzed using gas chromatography coupled with mass spectrometry (GC-MS). The results indicated that the density and degree of protrusion and the number of plastids and osmiophilic matrix granules in the petals play key roles in emitting the fragrance. There were distinct differences in the components and relative contents of volatile compounds among the different strains of N. hybrid. In total, 29, 34, 39, and 43 volatile compounds were detected in the cut flower petals of the blue-purple type (Nh-1), pink type (Nh-2), yellow type (Nh-3) and white type (Nh-4) of N. hybrid at the flowering stage, with total relative contents of 96.78%, 97.64%, 98.56%, and 96.15%, respectively. Analyses of these volatile components indicated that alkenes, alcohols, and alkanes were the three major types of volatile components in the flower petals of N. hybrid. The predominant volatile compounds were benzyl alcohol, pentadecane, trans-\u03b1-bergamotene, (E)-\u03b2-farnesene, and (6E,9E)-6,9-heptadecadiene, and some of these volatile compounds were terpenes, which varied among the different strains. Moreover, on the basis of hierarchical cluster analysis (HCA) and principal component analysis (PCA), the N. hybrid samples were divided into four groups: alcohols were the most important volatile compounds for Nh-4 samples; esters and aldehydes were the predominant volatiles in Nh-3 samples; and ketones and alkenes were important for Nh-2 samples. These compounds contribute to the unique flavors and aromas of the four strains of N. hybrid.<\/jats:p>","DOI":"10.3390\/molecules27020408","type":"journal-article","created":{"date-parts":[[2022,1,9]],"date-time":"2022-01-09T23:06:15Z","timestamp":1641769575000},"page":"408","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":19,"title":["Comparative Study of the Petal Structure and Fragrance Components of the Nymphaea hybrid, a Precious Water Lily"],"prefix":"10.3390","volume":"27","author":[{"given":"Qi","family":"Zhou","sequence":"first","affiliation":[{"name":"College of Environment and Ecology, Jiangsu Open University, Nanjing 210017, China"},{"name":"Co-Innovation Center for Sustainable Forestry in Southern China, Nanjing Forestry University, Nanjing 210037, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Man","family":"Shi","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Subtropical Silviculture, Zhejiang Agriculture and Forestry University, Hangzhou 311300, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8778-8122","authenticated-orcid":false,"given":"Huihui","family":"Zhang","sequence":"additional","affiliation":[{"name":"Co-Innovation Center for Sustainable Forestry in Southern China, Nanjing Forestry University, Nanjing 210037, China"},{"name":"College of Landscape Architecture, Nanjing Forestry University, Nanjing 210037, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zunling","family":"Zhu","sequence":"additional","affiliation":[{"name":"Co-Innovation Center for Sustainable Forestry in Southern China, Nanjing Forestry University, Nanjing 210037, China"},{"name":"College of Landscape Architecture, Nanjing Forestry University, Nanjing 210037, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2022,1,9]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"1359","DOI":"10.1039\/c1np00021g","article-title":"Plant volatiles: Production, function and pharmacology","volume":"28","author":"Maffei","year":"2011","journal-title":"Nat. Prod. Rep."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"667809","DOI":"10.3389\/fpls.2021.667809","article-title":"Comparative study of fig volatile compounds using headspace solid-phase microextraction-gas chromatography\/mass spectrometry: Effects of cultivars and ripening stages","volume":"12","author":"Zidi","year":"2021","journal-title":"Front. Plant Sci."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"32","DOI":"10.1016\/j.ppees.2013.11.002","article-title":"Floral volatiles play a key role in specialized ant pollination","volume":"16","author":"Vega","year":"2014","journal-title":"Perspect. Plant Ecol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1007\/s10265-018-1073-2","article-title":"Pollination features and floral volatiles of Gymnospermium scipetarum (barberidaceae)","volume":"132","author":"Rosati","year":"2019","journal-title":"J. Plant Res."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"712","DOI":"10.1111\/j.1365-313X.2008.03446.x","article-title":"Biosynthesis of plant-derived flavor compounds","volume":"54","author":"Schwab","year":"2008","journal-title":"Plant J."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"777","DOI":"10.1093\/aob\/mcq045","article-title":"Floral scents repel facultative flower visitors, but attract obligate ones","volume":"105","author":"Junker","year":"2010","journal-title":"Ann. Bot."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"1173","DOI":"10.1023\/A:1016229616845","article-title":"Allelopathic potential and chemical constituents of volatiles from Ageratum conyzoides under stress","volume":"28","author":"Kong","year":"2002","journal-title":"J. Chem. Ecol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"845","DOI":"10.1111\/j.1365-2745.2010.01665.x","article-title":"Flower-scent mimicry masks a deadly trap in the carnivorous plant Nepenthes rafflesiana","volume":"98","author":"Giusto","year":"2010","journal-title":"J. Ecol."},{"key":"ref_9","first-page":"13","article-title":"Optimized method to analyze rose plant volatile organic compounds by HS-SPME-GC-FID\/MSD","volume":"5","author":"Ibrahim","year":"2017","journal-title":"J. Biosci. Med."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"2020","DOI":"10.1016\/j.jksus.2020.02.001","article-title":"Function-specific volatiles and volatilization characteristics of Dendrobium officinale","volume":"32","author":"Dong","year":"2020","journal-title":"J. King Saud Univ. Sci."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"1601","DOI":"10.1021\/jf0306237","article-title":"Volatile constituents and key odorants in leaves, buds, flowers, and fruits of Laurus nobilis L.","volume":"52","author":"Kilic","year":"2004","journal-title":"J. Agric. Food Chem."},{"key":"ref_12","first-page":"855","article-title":"The impact of greenhouse tomato (Solanales: Solanaceae) floral volatiles on bumble bee (Hymenoptera: Apidae) pollination","volume":"4","author":"Andrew","year":"2012","journal-title":"Environ. Entomol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"1612","DOI":"10.1105\/tpc.104.028837","article-title":"ODORANT1 regulates fragrance biosynthesis in Petunia flowers","volume":"17","author":"Verdonk","year":"2005","journal-title":"Plant Cell"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"144","DOI":"10.1007\/s11032-018-0902-2","article-title":"An R2R3-MYB transcription factor ODORANT1 regulates fragrance biosynthesis in lilies (Lilium spp.)","volume":"38","author":"Yoshida","year":"2018","journal-title":"Mol. Breed."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"319","DOI":"10.3389\/fpls.2017.00319","article-title":"Comparative transcriptome reveals benzenoid biosynthesis regulation as inducer of floral scent in the woody plant Prunus mume","volume":"8","author":"Zhao","year":"2017","journal-title":"Front. Plant Sci."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1867","DOI":"10.3732\/ajb.89.12.1867","article-title":"Floral anatomy and systematics of Alliaceae with particular reference to Gilliesia, a presumed insect mimic with strongly zygomorphic flowers","volume":"89","author":"Rudall","year":"2002","journal-title":"Am. J. Bot."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1016\/j.flora.2009.01.002","article-title":"Can a stench be beautiful? Osmophores in stem-succulent stapeliads (Apocynaceae-Asclepiadoideae-Ceropegieae-Stapeliinae)","volume":"205","author":"Plachno","year":"2010","journal-title":"Flora"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"499","DOI":"10.1093\/aob\/mch072","article-title":"Lip anatomy and its implications for the pollination mechanisms of Bulbophyllum species (Orchidaceae)","volume":"93","author":"Teixeira","year":"2004","journal-title":"Ann. Bot."},{"key":"ref_19","first-page":"710","article-title":"Research advances on the osmophores in plants","volume":"23","author":"Cui","year":"2015","journal-title":"J. Trop. Subtrop. Bot."},{"key":"ref_20","first-page":"1443","article-title":"Sem studies of petal structure of corolla of the species Sibbaldia L. (Rosaceae)","volume":"42","author":"Tahir","year":"2010","journal-title":"Pak. J. Bot."},{"key":"ref_21","first-page":"62","article-title":"Osmophores and floral fragrance in Anacardium humile and Mangifera indica (anacardiaceae): An overlooked secretory structure in sapindales","volume":"10","author":"Tlke","year":"2018","journal-title":"AoB Plants"},{"key":"ref_22","first-page":"34","article-title":"Variation in perianth structure during the blooming of Michelia alba DC. and its aroma emission mechanism","volume":"27","author":"Guo","year":"2006","journal-title":"Chin. J. Trop. Crop."},{"key":"ref_23","first-page":"33","article-title":"Aromatic constituents of Chunlan and its maternal variety Tieguanying","volume":"14","author":"Zhang","year":"1999","journal-title":"Fujian J. Agricul. Sci."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"213","DOI":"10.1002\/ffj.1875","article-title":"Factors affecting secondary metabolite production in plants: Volatile components and essential oils","volume":"23","author":"Figuieredo","year":"2008","journal-title":"Flavour Fragr. J."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1663\/0006-8101(2006)72[1:DADOFS]2.0.CO;2","article-title":"Diversity and distribution of floral scent","volume":"72","author":"Knudsen","year":"2006","journal-title":"Bot. Rev."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1080\/07352680600899973","article-title":"Plant volatiles: Recent advances and future perspectives","volume":"25","author":"Dudareva","year":"2006","journal-title":"Crit. Rev. Plant Sci."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"1893","DOI":"10.1104\/pp.104.049981","article-title":"Biochemistry of plant volatiles","volume":"135","author":"Dudareva","year":"2004","journal-title":"Plant Physiol."},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Fu, J., Dan, H., Chao, Z., Bao, Z., Zhao, H., and Hu, S. (2017). The emission of the floral scent of four Osmanthus fragrans cultivars in response to different temperatures. Molecules, 22.","DOI":"10.3390\/molecules22030430"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"1541","DOI":"10.1080\/00032719.2013.878840","article-title":"Determination of volatiles in water lily flowers using gas chromatography\u2013mass spectrometry","volume":"47","author":"Yuan","year":"2014","journal-title":"Anal. Lett."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"211","DOI":"10.1093\/aob\/mcu235","article-title":"Floral biology and ovule and seed ontogeny of Nymphaea thermarum, water lily at the brink of extinction with potential as a model system for basal angiosperms","volume":"115","author":"Povilus","year":"2015","journal-title":"Ann. Bot."},{"key":"ref_31","unstructured":"Henry, S.C. (1905). The Waterlilies, The Carnegie Institution of Washington."},{"key":"ref_32","first-page":"200","article-title":"Extraction optimization through response surface methodology and antioxidant activity of polyphenols of Nymphaea Hybrid","volume":"34","author":"Zhou","year":"2018","journal-title":"Mod. Food Sci. Technol."},{"key":"ref_33","first-page":"101","article-title":"Analysis of essential oil from the flowers of Nymphaea hybrida by GC-MS","volume":"29","author":"Xu","year":"2008","journal-title":"Food Res. Dev."},{"key":"ref_34","first-page":"19","article-title":"Extraction, isolation and structural determination of the polysaccharides from Nymphaea hybrid","volume":"36","author":"Jiang","year":"2017","journal-title":"Chin. Wild Plant Res."},{"key":"ref_35","first-page":"5","article-title":"Acute and chronic toxicity of Nymphaea hybrida","volume":"34","author":"Jin","year":"2015","journal-title":"Chin. Wild Plant Res."},{"key":"ref_36","first-page":"28","article-title":"Effect of Nymphaea hybrid on prostatic hyperplasia in rats","volume":"15","author":"Che","year":"2015","journal-title":"J. Chin. Inst. Food Sci. Technol."},{"key":"ref_37","first-page":"227","article-title":"Inhibitory effect of extracts from Nymphaea hybrid flowers on pancreatic lipase","volume":"42","author":"Zhang","year":"2017","journal-title":"Food Sci. Technol."},{"key":"ref_38","first-page":"776","article-title":"Study on whitening effect of Nymphaea hybrid extracts","volume":"37","author":"Sun","year":"2018","journal-title":"J. Food Sci. Biotechnol."},{"key":"ref_39","first-page":"19","article-title":"Study on inhibition of ethanol extract from Nymphaea hybrid on the activity of tyrosinase","volume":"12","author":"Ren","year":"2009","journal-title":"Deterg. Cosmet."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"110268","DOI":"10.1016\/j.scienta.2021.110268","article-title":"Flowering characteristics and reproductive biology of Nymphaea hybrid, a precious water lily","volume":"287","author":"Zhang","year":"2021","journal-title":"Sci. Hortic."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"506","DOI":"10.1111\/j.1438-8677.2008.00140.x","article-title":"A simple floral fragrance and unusual osmophore structure in Cyclopogon elatus (Orchidaceae)","volume":"11","author":"Wiemer","year":"2010","journal-title":"Plant Biol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"52","DOI":"10.1016\/j.flora.2016.02.003","article-title":"Protection and attraction: Bracts and secretory structures in reduced inflorescences of Malpighiales","volume":"220","author":"Gagliardi","year":"2016","journal-title":"Flora"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1111\/plb.12000","article-title":"Scent glands in legume flowers","volume":"16","author":"Marinho","year":"2014","journal-title":"Plant Biol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"146","DOI":"10.1111\/plb.13061","article-title":"Unravelling the structure and function of the petal appendages in the tribe Schwenckieae (Solanaceae)","volume":"22","author":"Paucar","year":"2020","journal-title":"Plant Biol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"1127","DOI":"10.1007\/s00468-015-1193-4","article-title":"Corolla structure and fragrance components in Styrax tonkinensis","volume":"29","author":"Xu","year":"2015","journal-title":"Trees"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1093\/botlinnean\/boy002","article-title":"Chemical composition and secretion biology of the floral bouquet in legume trees (Fabaceae)","volume":"187","author":"Marinho","year":"2018","journal-title":"Bot. J. Linn. Soc."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"338","DOI":"10.1016\/j.tplants.2016.12.009","article-title":"The sexual advantage of looking, smelling, and tasting good: The metabolic network that produces signals for pollinators","volume":"22","author":"Borghi","year":"2017","journal-title":"Trends Plant Sci."},{"key":"ref_48","first-page":"97","article-title":"Study on the aroma component emission pattern of Osmanthus fragrans \u2018Boye Jingui\u2019","volume":"42","author":"Shi","year":"2018","journal-title":"J. Nanjing For. Univ."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"929","DOI":"10.1007\/s10886-006-9044-9","article-title":"Analysis of the volatiles emitted by whole flowers and isolated flower organs of the carob tree using HS-SPME-GC\/MS","volume":"32","author":"Serra","year":"2006","journal-title":"J. Chem. Ecol."},{"key":"ref_50","first-page":"1130","article-title":"Study of volatiles in Lysimachia parvifolia flower using HS-SPME-GC-MS","volume":"50","author":"Yang","year":"2014","journal-title":"Chem. Nat. Compd."},{"key":"ref_51","doi-asserted-by":"crossref","unstructured":"Ma, X.K., Li, X.F., Zhang, J.Y., Lei, J., Li, W.W., and Wang, G. (2020). Analysis of the volatile components in Selaginella doederleinii by headspace solid phase microextraction-gas chromatography-mass spectrometry. Molecules, 25.","DOI":"10.3390\/molecules25010115"},{"key":"ref_52","doi-asserted-by":"crossref","unstructured":"Tsai, F.J., Liu, H.J., Lee, M.Y., and Lin, C.C. (2019). Determination of volatile components from live water lily flowers by an orthogonal-array-design-assisted trapping cell. Appl. Sci., 9.","DOI":"10.3390\/app9071269"},{"key":"ref_53","first-page":"104","article-title":"GC-MS analysis on components of essential oil from fresh flowers of tropical water lily","volume":"26","author":"Shi","year":"2017","journal-title":"J. Plant Res. Environ."},{"key":"ref_54","first-page":"108","article-title":"Research on the cell microstructure of Jasminum sambac\u2019s petal","volume":"25","author":"Zhang","year":"1999","journal-title":"J. Hunan Agric. Univ."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"1357","DOI":"10.1351\/pac199062071357","article-title":"The chemistry of flowers, fruits and spices: Live vs. dead\u2014A new dimension in fragrance research","volume":"62","author":"Mookherjee","year":"1990","journal-title":"Pure Appl. Chem."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"117","DOI":"10.17660\/ActaHortic.2012.943.12","article-title":"Volatile compounds in four species of thai waterlily (Nymphaea spp.)","volume":"943","author":"Jirapong","year":"2012","journal-title":"Acta Hortic."},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Francesco, S.R.D.C., Jacopo, C., Elia, B., Annalisa, G., Cinzia, B., and Aldo, T. (2019). Characterization and antioxidant activity of essential oil of four sympatric orchid species. Molecules, 24.","DOI":"10.3390\/molecules24213878"},{"key":"ref_58","first-page":"1366","article-title":"Characterisation of volatile compounds in Cilento (Italy) figs (Ficus carica L.) cv. Dottato as affected by the drying process","volume":"20","author":"Russo","year":"2017","journal-title":"Int. J. Food Prop."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"74","DOI":"10.1016\/j.foodchem.2016.03.034","article-title":"Benzaldehyde in cherry flavour as a precursor of benzene formation in beverages","volume":"206","author":"Loch","year":"2016","journal-title":"Food Chem."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"043101","DOI":"10.1063\/1.3653552","article-title":"Retention indices for frequently reported compounds of plant essential oils","volume":"40","author":"Babushok","year":"2011","journal-title":"J. Phys. Chem. Ref. Data"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"182","DOI":"10.1016\/j.jchromb.2008.04.042","article-title":"Retention index thresholds for compound matching in GC-MS metabolite profiling","volume":"871","author":"Strehmel","year":"2018","journal-title":"J. Chromatogr. B Anal. Technol. Biomed. Life Sci."},{"key":"ref_62","doi-asserted-by":"crossref","unstructured":"Zhu, X., Li, Q., Li, J., Luo, J., Chen, W., and Li, X. (2018). Comparative study of volatile compounds in the fruit of two banana cultivars at different ripening stages. Molecules, 23.","DOI":"10.3390\/molecules23102456"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1016\/j.foodchem.2015.04.032","article-title":"Assessment of flavonoids and volatile compounds in tea infusions of water lily flowers and their antioxidant activities","volume":"187","author":"Yin","year":"2015","journal-title":"Food Chem."}],"container-title":["Molecules"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1420-3049\/27\/2\/408\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,13]],"date-time":"2025-10-13T13:38:23Z","timestamp":1760362703000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1420-3049\/27\/2\/408"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,1,9]]},"references-count":63,"journal-issue":{"issue":"2","published-online":{"date-parts":[[2022,1]]}},"alternative-id":["molecules27020408"],"URL":"https:\/\/doi.org\/10.3390\/molecules27020408","relation":{},"ISSN":["1420-3049"],"issn-type":[{"value":"1420-3049","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,1,9]]}}}