{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T05:16:25Z","timestamp":1788239785007,"version":"build-2803163510"},"reference-count":87,"publisher":"MDPI AG","issue":"17","license":[{"start":{"date-parts":[[2026,8,30]],"date-time":"2026-08-30T00:00:00Z","timestamp":1788048000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100004763","name":"Natural Science Foundation of Inner Mongolia","doi-asserted-by":"crossref","award":["2026QC0549"],"award-info":[{"award-number":["2026QC0549"]}],"id":[{"id":"10.13039\/501100004763","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>Cucurbitacins are highly oxygenated tetracyclic triterpenoids characterized by intense bitterness, substantial structural diversity, and important consequences for plant\u2013herbivore interactions. Although best known from Cucurbitaceae, cucurbitacins and related cucurbitane-type metabolites also occur in phylogenetically distant herbaceous and woody plants. Genetic and biochemical studies have validated several core biosynthetic steps, including cucurbitadienol formation by oxidosqualene cyclases and subsequent modification by cytochrome P450 monooxygenases, acyltransferases, and glycosyltransferases. Tissue-preferential basic helix\u2013loop\u2013helix transcription factors constitute the best-characterized regulatory layer, whereas the evidence supporting accessory regulators, transporters, and environmental responses varies from functional validation to transcriptomic or genomic prediction. From the plant perspective, cucurbitacins deter feeding or impair performance in many generalist and non-adapted herbivores. By contrast, their use as host-recognition cues and feeding stimulants by specialist diabroticite beetles reflects evolved herbivore adaptations involving perception, tolerance, metabolism, or sequestration rather than a second defensive function of the plant trait. Herbivore-induced cucurbitacin accumulation has been demonstrated in particular systems, although its regulatory mechanisms and ecological generality remain unresolved. Unlike previous reviews centered primarily on cucurbitacin chemistry, pharmacological activity, or individual biosynthetic pathways, this review integrates evidence-graded pathway reconstruction and molecular regulation with taxonomic distribution, insect adaptation, domestication, and agroecological consequences. Mechanistically, this review traces how scaffold formation, oxidative tailoring, conjugation, tissue-specific regulation, and transport give rise to contrasting ecological outcomes through herbivore-specific perception, tolerance, metabolism, and sequestration. We conclude that uniformly increasing or eliminating cucurbitacins is unlikely to provide broadly effective crop resistance because either direction may favor a different herbivore group. Future priorities include functional validation of candidate genes, spatially resolved metabolite analysis, comparative investigation of non-cucurbit lineages, and field evaluation involving generalist and specialist herbivores, crop quality, and non-target organisms. These advances will support context-specific fruit-quality improvement, behavioral pest control, and integrated pest management strategies rather than cucurbitacin manipulation as a stand-alone resistance approach.<\/jats:p>","DOI":"10.3390\/plants15172660","type":"journal-article","created":{"date-parts":[[2026,8,31]],"date-time":"2026-08-31T02:08:58Z","timestamp":1788142138000},"page":"2660","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Cucurbitacins in Plant\u2013Insect Interactions: Biosynthesis, Regulation, Ecological Functions, and Prospects for Crop Protection"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0009-0004-4748-9437","authenticated-orcid":false,"given":"Qi","family":"Zhang","sequence":"first","affiliation":[{"name":"College of Forestry, Inner Mongolia Agricultural University, Hohhot 010018, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yu-e","family":"Bai","sequence":"additional","affiliation":[{"name":"College of Forestry, Inner Mongolia Agricultural University, Hohhot 010018, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Aoga","family":"Li","sequence":"additional","affiliation":[{"name":"College of Forestry, Inner Mongolia Agricultural University, Hohhot 010018, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,30]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"386","DOI":"10.1039\/b418841c","article-title":"Cucurbitacins and cucurbitane glycosides: Structures and biological activities","volume":"22","author":"Chen","year":"2005","journal-title":"Nat. Prod. Rep."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"225","DOI":"10.1146\/annurev-arplant-050312-120229","article-title":"Triterpene biosynthesis in plants","volume":"65","author":"Thimmappa","year":"2014","journal-title":"Annu. Rev. Plant Biol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"1331","DOI":"10.1142\/S0192415X15500755","article-title":"Cucurbitacins: A systematic review of the phytochemistry and anticancer activity","volume":"43","author":"Cai","year":"2015","journal-title":"Am. J. Chin. Med."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"e9323","DOI":"10.1002\/rcm.9323","article-title":"Screening and identifying cucurbitacins and cucurbitacin glycosides in Cucumis sativus using high-performance liquid chromatography\/quadrupole-time-of-flight mass spectrometry combined with in-source fragmentation and alkali adduct ions","volume":"36","author":"Chen","year":"2022","journal-title":"Rapid Commun. Mass Spectrom."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"1000","DOI":"10.1016\/j.molp.2017.02.005","article-title":"Developmentally regulated glucosylation of bitter triterpenoid in cucumber by the UDP-glucosyltransferase UGT73AM3","volume":"10","author":"Zhong","year":"2017","journal-title":"Mol. Plant"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"995","DOI":"10.1111\/nph.18786","article-title":"Vacuolar MATE\/DTX protein-mediated cucurbitacin C transport is co-regulated with bitterness biosynthesis in cucumber","volume":"238","author":"Ma","year":"2023","journal-title":"New Phytol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"5534","DOI":"10.1021\/jf4002457","article-title":"Chemical modification produces species-specific changes in cucurbitacin antifeedant effect","volume":"61","author":"Lang","year":"2013","journal-title":"J. Agric. Food Chem."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"4659","DOI":"10.1093\/molbev\/msab213","article-title":"An independent evolutionary origin for insect-deterrent cucurbitacins in Iberis amara","volume":"38","author":"Dong","year":"2021","journal-title":"Mol. Biol. Evol."},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Chen, C.H., Kuo, T.C., Yang, M.H., Chien, T.Y., Chu, M.J., Huang, L.C., Chen, C.Y., Lo, H.F., Jeng, S.T., and Chen, L.F.O. (2014). Identification of cucurbitacins and assembly of a draft genome for Aquilaria agallocha. BMC Genom., 15.","DOI":"10.1186\/1471-2164-15-578"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"234","DOI":"10.3724\/SP.J.1009.2012.00234","article-title":"Cucurbitacins from fruits of Aquilaria sinensis","volume":"10","author":"Mei","year":"2012","journal-title":"Chin. J. Nat. Med."},{"key":"ref_11","first-page":"290","article-title":"Structural elucidation of two new megastigmane glycosides from the leaves of Aquilaria sinensis","volume":"13","author":"Li","year":"2015","journal-title":"Chin. J. Nat. Med."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1007\/s13659-022-00334-3","article-title":"Bioassay-guided isolation of cytotoxic constituents from the flowers of Aquilaria sinensis","volume":"12","author":"Yang","year":"2022","journal-title":"Nat. Prod. Bioprospect."},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Ding, X., Yang, Z., Wang, H., Zeng, J., Dai, H., and Mei, W. (2024). Aquilaria sinensis: An upstart resource for cucurbitacin production offers insights into the origin of plant bitter (Bi) gene clusters. Plants, 13.","DOI":"10.3390\/plants13020260"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"444","DOI":"10.1021\/np200879p","article-title":"Isolation, structure elucidation, and biological evaluation of 16,23-epoxycucurbitacin constituents from Elaeocarpus chinensis","volume":"75","author":"Pan","year":"2012","journal-title":"J. Nat. Prod."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1016\/S0031-9422(02)00232-7","article-title":"Ellagic acid derivatives and cytotoxic cucurbitacins from Elaeocarpus mastersii","volume":"61","author":"Ito","year":"2002","journal-title":"Phytochemistry"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"112988","DOI":"10.1016\/j.phytochem.2021.112988","article-title":"Pentacyclic and hexacyclic cucurbitacins from Elaeocarpus petiolatus","volume":"193","author":"Cho","year":"2022","journal-title":"Phytochemistry"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"113581","DOI":"10.1016\/j.phytochem.2022.113581","article-title":"Heterodimers with a cucurbitane-type triterpenoid skeleton from the branches of Elaeocarpus dubius","volume":"207","author":"Hu","year":"2023","journal-title":"Phytochemistry"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1741","DOI":"10.1055\/s-2008-1081356","article-title":"Cytotoxic cucurbitane-type triterpenoids from Elaeocarpus hainanensis","volume":"74","author":"Meng","year":"2008","journal-title":"Planta Med."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"1084","DOI":"10.1126\/science.1259215","article-title":"Biosynthesis, regulation, and domestication of bitterness in cucumber","volume":"346","author":"Shang","year":"2014","journal-title":"Science"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"16183","DOI":"10.1038\/nplants.2016.183","article-title":"Convergence and divergence of bitterness biosynthesis and regulation in Cucurbitaceae","volume":"2","author":"Zhou","year":"2016","journal-title":"Nat. Plants"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"2724","DOI":"10.1111\/pbi.14399","article-title":"CmBr confers fruit bitterness under CPPU treatment in melon","volume":"22","author":"Wang","year":"2024","journal-title":"Plant Biotechnol. J."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"24697","DOI":"10.1021\/acs.jafc.4c04608","article-title":"Transcription factor CmWRKY13 regulates cucurbitacin B biosynthesis leading to bitterness in oriental melon fruit (Cucumis melo var. makuwa Makino)","volume":"72","author":"Luo","year":"2024","journal-title":"J. Agric. Food Chem."},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Zhao, G., Wang, M., Luo, C., Li, J., Gong, H., Zheng, X., Liu, X., Luo, J., and Wu, H. (2022). Metabolome and transcriptome analyses of cucurbitacin biosynthesis in luffa (Luffa acutangula). Front. Plant Sci., 13.","DOI":"10.3389\/fpls.2022.886870"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"314","DOI":"10.1016\/j.plaphy.2022.06.014","article-title":"Comparative transcriptome analysis and identification of candidate genes involved in cucurbitacin IIa biosynthesis in Hemsleya macrosperma","volume":"185","author":"Zhang","year":"2022","journal-title":"Plant Physiol. Biochem."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"100835","DOI":"10.1016\/j.xplc.2024.100835","article-title":"Biosynthetic pathway of prescription cucurbitacin IIa and high-level production of key triterpenoid intermediates in engineered yeast and tobacco","volume":"5","author":"Chen","year":"2024","journal-title":"Plant Commun."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"uhac038","DOI":"10.1093\/hr\/uhac038","article-title":"Comparative genomics analysis of bHLH genes in cucurbits identifies a novel gene regulating cucurbitacin biosynthesis","volume":"9","author":"Xu","year":"2022","journal-title":"Hortic. Res."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"225","DOI":"10.1023\/A:1021945101308","article-title":"Role of cucurbitacin C in resistance to spider mite (Tetranychus urticae) in cucumber (Cucumis sativus L.)","volume":"29","author":"Zijlstra","year":"2003","journal-title":"J. Chem. Ecol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"16473","DOI":"10.1038\/s41598-018-34821-w","article-title":"Impact of the secondary plant metabolite cucurbitacin B on the demographical traits of the melon aphid, Aphis gossypii","volume":"8","author":"Yousaf","year":"2018","journal-title":"Sci. Rep."},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Zhao, C., Ma, C., Luo, J., Niu, L., Hua, H., Zhang, S., and Cui, J. (2021). Potential of Cucurbitacin B and Epigallocatechin Gallate as Biopesticides against Aphis gossypii. Insects, 12.","DOI":"10.3390\/insects12010032"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"23390","DOI":"10.1007\/s11356-019-04438-1","article-title":"Toxicity and developmental effect of cucurbitacin E from Citrullus colocynthis L. against Spodoptera litura and the non-target earthworm Eisenia fetida","volume":"27","author":"Ponsankar","year":"2020","journal-title":"Environ. Sci. Pollut. Res."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"1061","DOI":"10.1007\/s10340-022-01568-3","article-title":"Sequestration of cucurbitacins from cucumber plants by Diabrotica balteata larvae provides little protection against biological control agents","volume":"96","author":"Bruno","year":"2023","journal-title":"J. Pest Sci."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"106","DOI":"10.1007\/s00425-023-04139-7","article-title":"Consequences of squash (Cucurbita argyrosperma) domestication for plant defence and herbivore interactions","volume":"257","author":"Jaccard","year":"2023","journal-title":"Planta"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"2812","DOI":"10.1111\/pce.13844","article-title":"Divergence of defensive cucurbitacins in independent Cucurbita pepo domestication events leads to differences in specialist herbivore preference","volume":"43","author":"Brzozowski","year":"2020","journal-title":"Plant Cell Environ."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1392","DOI":"10.1126\/science.153.3742.1392","article-title":"Cucurbitacins: Specific insect attractants in Cucurbitaceae","volume":"153","author":"Chambliss","year":"1966","journal-title":"Science"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"3769","DOI":"10.1073\/pnas.77.7.3769","article-title":"Cucurbitacins as kairomones for diabroticite beetles","volume":"77","author":"Metcalf","year":"1980","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"1307","DOI":"10.1007\/BF01024118","article-title":"Disposition and fate of cucurbitacin B in five species of diabroticites","volume":"11","author":"Ferguson","year":"1985","journal-title":"J. Chem. Ecol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1146\/annurev.ento.54.110807.090434","article-title":"Adaptation and invasiveness of western corn rootworm: Intensifying research on a worsening pest","volume":"54","author":"Gray","year":"2009","journal-title":"Annu. Rev. Entomol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"15992","DOI":"10.1038\/s41598-019-52404-1","article-title":"Metabolite profiling and quantitation of cucurbitacins in Cucurbitaceae plants by liquid chromatography coupled to tandem mass spectrometry","volume":"9","author":"Ali","year":"2019","journal-title":"Sci. Rep."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"135006","DOI":"10.1016\/j.foodchem.2022.135006","article-title":"Cucurbitacin and volatile compound profiling reveals independent domestication of cucumber (Cucumis sativus L.) fruit","volume":"405","author":"Min","year":"2023","journal-title":"Food Chem."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"1165","DOI":"10.1080\/14786419.2014.921787","article-title":"23,24-Dihydrocucurbitacin C: A new compound regarded as the next metabolite of cucurbitacin C","volume":"28","author":"Qing","year":"2014","journal-title":"Nat. Prod. Res."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"106065","DOI":"10.1016\/j.jfca.2024.106065","article-title":"Quantification of cucurbitacin C in bitter cucumber and its reduction by fermentation and acidification","volume":"129","author":"Fan","year":"2024","journal-title":"J. Food Compos. Anal."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"554","DOI":"10.21273\/HORTSCI.11.6.554","article-title":"Genes of the Cucurbitaceae","volume":"11","author":"Robinson","year":"1976","journal-title":"HortScience"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"2285","DOI":"10.1023\/A:1020821823794","article-title":"Polymorphism in plant defense against herbivory: Constitutive and induced resistance in Cucumis sativus","volume":"25","author":"Agrawal","year":"1999","journal-title":"J. Chem. Ecol."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"6995","DOI":"10.1016\/j.tet.2004.04.088","article-title":"Cucurbitadienol synthase, the first committed enzyme for cucurbitacin biosynthesis, is a distinct enzyme from cycloartenol synthase for phytosterol biosynthesis","volume":"60","author":"Shibuya","year":"2004","journal-title":"Tetrahedron"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"1172","DOI":"10.1093\/pcp\/pcv043","article-title":"Functional characterization of cucurbitadienol synthase and triterpene glycosyltransferase involved in biosynthesis of mogrosides from Siraitia grosvenorii","volume":"56","author":"Dai","year":"2015","journal-title":"Plant Cell Physiol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"18662","DOI":"10.1074\/jbc.RA119.009944","article-title":"Allylic hydroxylation of triterpenoids by a plant cytochrome P450 triggers key chemical transformations that produce a variety of bitter compounds","volume":"294","author":"Takase","year":"2019","journal-title":"J. Biol. Chem."},{"key":"ref_47","doi-asserted-by":"crossref","unstructured":"Benka, M., G\u00f6rlitz, K., Sch\u00f6ttgen, M.C., Lagies, S., Mohl, D.A., Kather, M., Du Preez-Bruwer, I., Mumbengegwi, D., Teufel, R., and Kowarschik, S. (2023). Mass spectrometric analysis of cucurbitacins and dihydrocucurbitacins from the tuber of Citrullus naudinianus. Biomolecules, 13.","DOI":"10.3390\/biom13081168"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"113123","DOI":"10.1016\/j.phytochem.2022.113123","article-title":"Identification of seven undescribed cucurbitacins in Cucumis sativus and their cytotoxic activity","volume":"197","author":"Qing","year":"2022","journal-title":"Phytochemistry"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"559","DOI":"10.1055\/s-2006-962095","article-title":"New cucurbitacin glycosides from Picrorhiza kurrooa","volume":"55","author":"Stuppner","year":"1989","journal-title":"Planta Med."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1007\/BF00987468","article-title":"Cucurbitacins from Cucurbita texana: Evidence for the role of isocucurbitacins","volume":"19","author":"Halaweish","year":"1993","journal-title":"J. Chem. Ecol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"253","DOI":"10.2503\/hortj.UTD-004","article-title":"Change in bitterness, accumulation of cucurbitacin B and expression patterns of CuB biosynthesis-related genes in melon during fruit development","volume":"88","author":"Hua","year":"2019","journal-title":"Hortic. J."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"1574","DOI":"10.2307\/1938019","article-title":"Squash beetle feeding behavior: An adaptation against induced cucurbit defenses","volume":"66","author":"Tallamy","year":"1985","journal-title":"Ecology"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1111\/j.1365-2311.1995.tb00428.x","article-title":"Squash beetle trenching behaviour: Avoidance of cucurbitacin induction or mucilaginous plant sap?","volume":"20","author":"McCloud","year":"1995","journal-title":"Ecol. Entomol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"kiag294","DOI":"10.1093\/plphys\/kiag294","article-title":"CmBt and CmBr synergistically regulate cucurbitacin B biosynthesis in melon","volume":"201","author":"Wang","year":"2026","journal-title":"Plant Physiol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"114207","DOI":"10.1016\/j.scienta.2025.114207","article-title":"Integrated transcriptomic and metabolomic analysis reveals spatiotemporal regulation of cucurbitacin B biosynthesis in melon (Cucumis melo L.)","volume":"348","author":"Tian","year":"2025","journal-title":"Sci. Hortic."},{"key":"ref_56","doi-asserted-by":"crossref","unstructured":"Kuo, T.C.Y., Chen, C.H., Chen, S.H., Lu, I.H., Chu, M.J., Huang, L.C., Lin, C.Y., Chen, C.Y., Lo, H.F., and Jeng, S.T. (2015). The effect of red light and far-red light conditions on secondary metabolism in agarwood. BMC Plant Biol., 15.","DOI":"10.1186\/s12870-015-0537-y"},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"1845","DOI":"10.1007\/s00299-021-02748-8","article-title":"De novo transcriptome analysis and identification of candidate genes associated with triterpenoid biosynthesis in Trichosanthes cucumerina L","volume":"40","author":"Lertphadungkit","year":"2021","journal-title":"Plant Cell Rep."},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Li, X., Chen, G., Gao, Q.Q., Xiang, C.F., Yuan, C.X., Li, X.N., Shu, Y.Y., Zhang, G.H., Liang, Y.L., and Yang, S.C. (2023). Site-directed mutagenesis identified the key active-site residues of 2,3-oxidosqualene cyclase HcOSC6 responsible for cucurbitacin biosynthesis in Hemsleya chinensis. Front. Plant Sci., 14.","DOI":"10.3389\/fpls.2023.1138893"},{"key":"ref_59","doi-asserted-by":"crossref","unstructured":"Almeida, A., Dong, L., Thorsen, T.H., Raadam, M.H., Khakimov, B., Carreno-Quintero, N., Kampranis, S.C., and Bak, S. (2022). Metabolic engineering of cucurbitacins in Cucurbita pepo hairy roots. Front. Plant Sci., 13.","DOI":"10.3389\/fpls.2022.1021907"},{"key":"ref_60","first-page":"3326","article-title":"Study of heterologous efficient synthesis of cucurbitadienol","volume":"42","author":"Li","year":"2017","journal-title":"China J. Chin. Mater. Med."},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"Wang, S., Xu, X., Lv, X., Liu, Y., Li, J., Du, G., and Liu, L. (2022). Construction and optimization of the de novo biosynthesis pathway of mogrol in Saccharomyces cerevisiae. Front. Bioeng. Biotechnol., 10.","DOI":"10.3389\/fbioe.2022.919526"},{"key":"ref_62","doi-asserted-by":"crossref","unstructured":"Wang, J., Li, C.H., Xiang, C.F., Zhou, P.H., Li, L.S., Li, X., Yang, S.C., Zhang, G.H., and Zhao, Y. (2024). Establishment and application of a highly efficient regeneration, genetic transformation, and genome-editing system for cucurbitacin biosynthesis in Hemsleya chinensis. BMC Plant Biol., 24.","DOI":"10.1186\/s12870-024-05717-x"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"107797","DOI":"10.1016\/j.plaphy.2023.107797","article-title":"Lessons learned from metabolic engineering in hairy roots: Transcriptome and metabolic profile changes caused by Rhizobium-mediated plant transformation in Cucurbitaceae species","volume":"201","author":"Almeida","year":"2023","journal-title":"Plant Physiol. Biochem."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"678","DOI":"10.1093\/ee\/26.3.678","article-title":"Cucurbitacins as feeding and oviposition deterrents to insects","volume":"26","author":"Tallamy","year":"1997","journal-title":"Environ. Entomol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"931","DOI":"10.1093\/ee\/11.4.931","article-title":"Cucurbitacin contents and diabroticite feeding upon Cucurbita spp","volume":"11","author":"Metcalf","year":"1982","journal-title":"Environ. Entomol."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1093\/jee\/76.1.47","article-title":"Influence of cucurbitacin content in cotyledons of Cucurbitaceae cultivars upon feeding behavior of diabroticine beetles","volume":"76","author":"Ferguson","year":"1983","journal-title":"J. Econ. Entomol."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"313","DOI":"10.1007\/BF02036720","article-title":"Identification of sensilla involved in taste mediation in adult western corn rootworm, Diabrotica virgifera virgifera","volume":"21","author":"Chyb","year":"1995","journal-title":"J. Chem. Ecol."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1016\/0020-1790(88)90038-8","article-title":"Metabolic transformations of cucurbitacins by Diabrotica virgifera virgifera and D. undecimpunctata howardi","volume":"18","author":"Andersen","year":"1988","journal-title":"Insect Biochem."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"738","DOI":"10.1007\/s10886-024-01523-9","article-title":"Cultivar-specific defense responses in wild and cultivated squash induced by belowground and aboveground herbivory","volume":"50","author":"Ye","year":"2024","journal-title":"J. Chem. Ecol."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"317","DOI":"10.1093\/ee\/nvv003","article-title":"Conspecific and heterospecific aboveground herbivory both reduce preference by a belowground herbivore","volume":"44","author":"Milano","year":"2015","journal-title":"Environ. Entomol."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"28","DOI":"10.1002\/pei3.10071","article-title":"The effect of squash domestication on a belowground tritrophic interaction","volume":"3","author":"Jaccard","year":"2022","journal-title":"Plant-Environ. Interact."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"2451","DOI":"10.1093\/jee\/toaf172","article-title":"Multicomponent attract-and-kill system for pest insects attacking cucurbit crops","volume":"118","author":"Wallingford","year":"2025","journal-title":"J. Econ. Entomol."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"1294","DOI":"10.1603\/EC10213","article-title":"Evaluation of cucurbitacin-based gustatory stimulant to facilitate cucumber beetle management with foliar insecticides in melons","volume":"104","author":"Pedersen","year":"2011","journal-title":"J. Econ. Entomol."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"1021","DOI":"10.1086\/505609","article-title":"Effects of the spatial pattern of leaf damage on growth and reproduction: Whole plants","volume":"167","author":"Stephenson","year":"2006","journal-title":"Int. J. Plant Sci."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1016\/j.baae.2025.04.002","article-title":"Cucurbit plant defenses against aboveground or belowground insect herbivores are distinct and shaped by eco-evolutionary factors","volume":"86","author":"Thompson","year":"2025","journal-title":"Basic Appl. Ecol."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"7897","DOI":"10.1038\/s41467-022-35621-7","article-title":"Graph-based pangenomics maximizes genotyping density and reveals structural impacts on fungal resistance in melon","volume":"13","author":"Vaughn","year":"2022","journal-title":"Nat. Commun."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"1649","DOI":"10.1038\/s41588-024-01823-6","article-title":"Telomere-to-telomere Citrullus super-pangenome provides direction for watermelon breeding","volume":"56","author":"Zhang","year":"2024","journal-title":"Nat. Genet."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"176","DOI":"10.1038\/s41477-024-01899-2","article-title":"Molecular regulation and domestication of parthenocarpy in cucumber","volume":"11","author":"Nie","year":"2025","journal-title":"Nat. Plants"},{"key":"ref_79","doi-asserted-by":"crossref","unstructured":"Uranga, M., Mart\u00edn-Hern\u00e1ndez, A.M., De Storme, N., and Pasin, F. (2024). CRISPR\u2013Cas systems and applications for crop bioengineering. Front. Bioeng. Biotechnol., 12.","DOI":"10.3389\/fbioe.2024.1483857"},{"key":"ref_80","doi-asserted-by":"crossref","unstructured":"Meinke, L.J., Souza, D., and Siegfried, B.D. (2021). The use of insecticides to manage the western corn rootworm, Diabrotica virgifera virgifera, LeConte: History, field-evolved resistance, and associated mechanisms. Insects, 12.","DOI":"10.3390\/insects12020112"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"100061","DOI":"10.1016\/j.ncrops.2024.100061","article-title":"The development and prospects of insect-resistant crops expressing double-strand RNAs","volume":"2","author":"Zhang","year":"2025","journal-title":"New Crops"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"100019","DOI":"10.1016\/j.ncrops.2024.100019","article-title":"Risk assessment of RNAi-based biopesticides","volume":"1","author":"Luo","year":"2024","journal-title":"New Crops"},{"key":"ref_83","doi-asserted-by":"crossref","unstructured":"Demurtas, O.C., Nicolia, A., and Diretto, G. (2023). Terpenoid transport in plants: How far from the final picture?. Plants, 12.","DOI":"10.3390\/plants12030634"},{"key":"ref_84","doi-asserted-by":"crossref","unstructured":"Zhang, J., Yu, Y., Qian, X., Zhang, X., Li, X., and Sun, X. (2024). Recent advances in the specialized metabolites mediating resistance to insect pests and pathogens in tea plants (Camellia sinensis). Plants, 13.","DOI":"10.3390\/plants13020323"},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"2168","DOI":"10.1111\/tpj.16924","article-title":"Advances in mass spectrometry imaging for plant metabolomics\u2014Expanding the analytical toolbox","volume":"119","author":"Yin","year":"2024","journal-title":"Plant J."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"563","DOI":"10.1038\/s41576-024-00691-4","article-title":"Plant pangenomes for crop improvement, biodiversity and evolution","volume":"25","author":"Schreiber","year":"2024","journal-title":"Nat. Rev. Genet."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"643","DOI":"10.1038\/s41588-026-02506-0","article-title":"Graph-based pangenome reveals structural variation dynamics during cucumber breeding","volume":"58","author":"Zhao","year":"2026","journal-title":"Nat. Genet."}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/17\/2660\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T04:24:49Z","timestamp":1788236689000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/17\/2660"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,30]]},"references-count":87,"journal-issue":{"issue":"17","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["plants15172660"],"URL":"https:\/\/doi.org\/10.3390\/plants15172660","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,30]]}}}