Cuticle is the major transpiration barrier that restricts non-stomatal water loss and is closely associated with plant drought tolerance. Although multiple efforts have been made, it remains controversial what factors shape up the cuticular transpiration barrier. Previously, we found that the cuticle from the tender tea leaf was mainly constituted by very-long-chain-fatty-acids and their derivatives while alicyclic compounds dominate the mature tea leaf cuticle. The presence of two contrasting cuticle within same branch offered a unique system to investigate this question. In this study, tea seedlings were subjected to water deprivation treatment, cuticle structures and wax compositions from the tender leaf and the mature leaf were extensively measured and compared. We found that cuticle wax coverage, thickness, and osmiophilicity were commonly increased from both leaves. New waxes species were specifically induced by drought; the composition of existing waxes was remodeled; the chain length distributions of alkanes, esters, glycols, and terpenoids were altered in complex manners. Drought treatment significantly reduced leaf water loss rates. Wax biosynthesis-related gene expression analysis revealed dynamic expression patterns dependent on leaf maturity and the severity of drought. These data suggested that drought stress-induced structural and compositional cuticular modifications improve cuticle water barrier property. In addition, we demonstrated that cuticle from the tender leaf and the mature leaf were modified through both common and distinct modes.

The cuticle is regarded as a non-living tissue; it remains unknown whether the cuticle could be reversibly modified and what are the potential mechanisms. In this study, three tea germplasms (Wuniuzao, 0202-10, and 0306A) were subjected to water deprivation followed by rehydration. The epicuticular waxes and intracuticular waxes from both leaf surfaces were quantified from the mature 5th leaf. Cuticular transpiration rates were then measured from leaf drying curves, and the correlations between cuticular transpiration rates and cuticular wax coverage were analyzed. We found that the cuticular transpiration barriers were reinforced by drought and reversed by rehydration treatment; the initial weak cuticular transpiration barriers were preferentially reinforced by drought stress, while the original major cuticular transpiration barriers were either strengthened or unaltered. Correlation analysis suggests that cuticle modifications could be realized by selective deposition of specific wax compounds into individual cuticular compartments through multiple mechanisms, including in vivo wax synthesis or transport, dynamic phase separation between epicuticular waxes and the intracuticular waxes, in vitro polymerization, and retro transportation into epidermal cell wall or protoplast for further transformation. Our data suggest that modifications of a limited set of specific wax components from individual cuticular compartments are sufficient to alter cuticular transpiration barrier properties.

Camellia sinensis (tea tree) is a perennial evergreen woody crop that has been planted in more than 50 countries worldwide; its leaves are harvested to make tea, which is one of the most popular nonalcoholic beverages. The cuticle is the major transpiration barrier to restrict nonstomatal water loss and it affects the drought tolerance of tea plants. The cuticle may also provide molecular cues for the interaction with herbivores and pathogens. The tea-making process almost always includes a postharvest withering treatment to reduce leaf water content, and many studies have demonstrated that withering treatment-induced metabolite transformation is essential to shape the quality of the tea made. Tea leaf cuticle is expected to affect its withering properties and the dynamics of postharvest metabolome remodeling. In addition, it has long been speculated that the cuticle may contribute to the aroma quality of tea. However, concrete experimental evidence is lacking to prove or refute this hypothesis. Even though its relevance to the abiotic and biotic stress tolerance and postharvest processing properties of tea tree, tea cuticle has long been neglected. Recently, there are several studies on the tea cuticle regarding its structure, wax composition, transpiration barrier organization, environmental stresses-induced wax modification, and structure–function relations. This review is devoted to tea cuticle, the recent research progresses were summarized and unresolved questions and future research directions were also discussed.

Fatty acids (FAs) are important precursors of oolong tea volatile substances, and their famous derivatives have been shown to be the key aroma components. However, the relationship between fatty acids and their derivatives during oolong tea production remains unclear. In this study, fresh Tieguanyin leaves were manufactured into oolong tea and green tea (control), and fatty acids and fatty acid-derived volatiles (FADV) were extracted from processed samples by the sulfuric acid–methanol method and solvent-assisted flavor evaporation (SAFE), respectively. The results showed that unsaturated fatty acids were more abundant than saturated fatty acids in fresh leaves and decreased significantly during tea making. Relative to that in green tea, fatty acids showed larger variations in oolong tea, especially at the green-making stage. Unlike fatty acids, the FADV content first increased and then decreased. During oolong tea manufacture, FADV contents were significantly and negatively correlated with total fatty acids; during the green-making stage, methyl jasmonate (MeJA) content was significantly and negatively correlated with abundant fatty acids except steric acid. Our data suggest that the aroma quality of oolong tea can be improved by manipulating fatty acid transformation.

The cuticle plays a major role in restricting nonstomatal water transpiration in plants. There is therefore a long-standing interest to understand the structure and function of the plant cuticle. Although many efforts have been devoted, it remains controversial to what degree the various cuticular parameters contribute to the water transpiration barrier. In this study, eight tea germplasms were grown under normal conditions; cuticle thickness, wax coverage, and compositions were analyzed from the epicuticular waxes and the intracuticular waxes of both leaf surfaces. The cuticular transpiration rates were measured from the individual leaf surface as well as the intracuticular wax layer. Epicuticular wax resistances were also calculated from both leaf surfaces. The correlation analysis between the cuticular transpiration rates (or resistances) and various cuticle parameters was conducted. We found that the abaxial cuticular transpiration rates accounted for 64–78% of total cuticular transpiration and were the dominant factor in the variations for the total cuticular transpiration. On the adaxial surface, the major cuticular transpiration barrier was located on the intracuticular waxes; however, on the abaxial surface, the major cuticular transpiration barrier was located on the epicuticular waxes. Cuticle thickness was not a factor affecting cuticular transpiration. However, the abaxial epicuticular wax coverage was found to be significantly and positively correlated with the abaxial epicuticular resistance. Correlation analysis suggested that the very-long-chain aliphatic compounds and glycol esters play major roles in the cuticular transpiration barrier in tea trees grown under normal conditions. Our results provided novel insights about the complex structure–functional relationships in the tea cuticle.