Effects of Drought Stress on Leaf Micromorphology, Glandular Trichomes, and the Accumulation of Essential Oils and Flavonoids in Four Lamiaceae Species

The effects of progressive drought stress were examined in four economically important plant species belonging to the Lamiaceae family: catnip (Nepeta cataria L.), lavender (Lavandula angustifolia Mill.), holy basil (Ocimum tenuiflorum L.), and perilla mint (Perilla frutescens (L.) Britton). Plants were grown in a controlled pot experiment under three soil water capacity levels: 70% (control), 50% (moderate stress), and 30% (severe stress), and the drought stress lasted for 30 days. The study evaluated a comprehensive set of leaf micromorphological parameters, including the density and diameter of glandular trichomes, stomatal density and size, and the thickness of the lamina, mesophyll, epidermis, cuticle, and parenchymal layers. In addition, essential oil (EO) content, total flavonoid content (TFC), and elemental composition were analyzed. Drought responses were strongly species-specific. O. tenuiflorum, P. frutescens, and N. cataria showed high sensitivity characterized by reduced biomass and thinning of leaf tissues. These changes were accompanied by typical xeromorphic adaptations, such as increased stomatal and glandular trichome density, and reduced stomatal size. L. angustifolia exhibited pronounced cuticle thickening, suggesting an effective structural mechanism to minimize water loss. Secondary metabolism also responded differently among species. In some cases, drought shifted metabolic allocation toward flavonoid accumulation at the expense of essential oils, whereas in others, moderate stress promoted the co-accumulation of both compounds. These patterns indicate distinct adaptive strategies linking anatomical plasticity with metabolic regulation. Overall, moderate drought supported adaptive responses, while severe water limitation impaired growth and metabolic production. From a practical perspective, maintaining moderate soil water availability appears critical to optimize both plant performance and the accumulation of valuable secondary metabolites in Lamiaceae species.