Defense Mechanisms Induced by Celery Seed Essential Oil against Powdery Mildew Incited by Podosphaera fusca in Cucumber

This study aimed to evaluate the effectiveness of essential oil extracted from celery (Apium graveolens) seeds (CSEO) for the control of powdery mildew of cucumber (Cucumis sativus) incited by Podosphaera fusca and to investigate the metabolic and genetic defense mechanisms triggered by the treatment with this essential oil in cucumber seedlings. The main compounds in the CSEO as determined by gas chromatography-mass spectrometry (GC-MS) analysis were d-limonene, 3-butyl phthalide, beta-selinene, and mandelic acid. The treatment with CSEO led to an increase in the content of both chlorophyll and phenolic/flavonoid compounds in cucumber leaves. In greenhouse tests, the application of CSEO reduced by 60% the disease severity on leaves of cucumber plants and stimulated the activity of defense-related enzymes such as beta-1,3-glucanase, chitinase, phenylalanine ammonia-lyase, peroxidase, and polyphenol oxidase. Moreover, treatment with CSEO induced overexpression of beta-1,3-glucanase, chitinase, and phenylalanine ammonia-lyase genes. A highly significant correlation was found between the beta-1,3-glucanase, chitinase, and phenylalanine ammonia-lyase enzymatic activities and the relative expression of the corresponding encoding genes in both inoculated and non-inoculated cucumber seedlings treated with the essential oil. Overall, this study showed that CSEO is a promising eco-friendly candidate fungicide that can be exploited to control cucumber powdery mildew.