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Very-long-chain (VLC) alkanes are the main wax compounds of tomato fruit and leaf. ECERIFERUM1 (CER1) and ECERIFERUM3 (CER3) are the two key genes involved in VLC alkane biosynthesis in Arabidopsis thaliana. However, the CER1 and CER3 homologous genes in tomato have not been investigated and their exact biological function remains unknown. We analyzed the wax profiles in tomato leaves and fruits at different growth stages, and characterized the CER1 and CER3 homologous genes. VLC alkanes were the predominant wax compounds both in the leaf and fruit at all developmental stages. We identified five CER1 homologs and two CER3 homologs in tomato, which were designated as SlCER1–1 to SlCER1–5 and SlCER3–1 and SlCER3–2 respectively. The genes exhibited tissue- and organ-dependent expression patterns and were induced by abiotic stresses. SlCER1–1 was localized to the endoplasmic reticulum (ER), which is also the main site of wax biosynthesis. Silencing the SlCER1–1 gene in tomato significantly reduced the amounts of n-Alkanes and branched alkanes, whereas its overexpression in Arabidopsis had the opposite effect. Under drought stress, both n-Alkanes and branched alkanes increased significantly in wild-type but not the SlCER1–1 RNAi tomato plants. Furthermore, SlCER1–1 silencing also increased the cuticular permeabilities of the leaves and fruits. In conclusion, SlCER1–1 is involved in wax alkane biosynthesis in tomato and plays an important role in the drought tolerance and fruit storability.

Brassinosteroid (BR), a growth-promoting phytohormone, regulates many plant growth processes including cell development. However, the mechanism by which BR regulates fiber growth is poorly understood. Cotton (Gossypium hirsutum) fibers are an ideal single-cell model in which to study cell elongation due to their length. Here we report that BR controls cotton fiber elongation by modulating very-long-chain fatty acid (VLCFA) biosynthesis. BR deficiency reduces the expression of 3-ketoacyl-CoA synthases (GhKCSs), the rate-limiting enzymes involved in VLCFA biosynthesis, leading to lower saturated VLCFAs contents in pagoda1 (pag1) mutant fibers. In vitro ovule culture experiments show that BR acts upstream of VLCFAs. Silencing of BRI1-EMS-SUPPRESOR 1.4 (GhBES1.4), encoding a master transcription factor (TF) of the BR signaling pathway, significantly reduces fiber length, whereas GhBES1.4 over-expression produces longer fibers. GhBES1.4 regulates endogenous VLCFA contents and directly binds to BR RESPONSE ELEMENTS (BRREs) in the GhKCS10_At promoter region, which in turn regulates GhKCS10_At expression to increase endogenous VLCFA contents. GhKCS10_At overexpression promotes cotton fiber elongation, whereas GhKCS10_At silencing inhibits cotton fiber growth, supporting a positive regulatory role for GhKCS10_At in fiber elongation. Overall, these results uncover a mechanism of fiber elongation through crosstalk between BR and VLCFAs at the single-cell level.