Trichomes are universal biological structures originating from the aerial epidermis, which serve as an excellent model to study plant differentiation at the cell level. Although the pathway regulating trichome formation in the Rosids has been well characterized, only very recently a few genes were identified for trichome initiation in the Asterids. In this study, we clonedWoolly(Wo), essential for trichome formation in tomato. Transgenic experiments revealed that the woolly phenotype is caused by the mutation inWowhich encodes a homeodomain protein containing a bZIP motif and a START domain. We identified three alleles ofWoand found that each allele contains a missense mutation, which respectively results in an amino acid substitution at the C terminus. Microarray and expression analysis showed that the expression of a B-type cyclin gene,SlCycB2, is possibly regulated byWo, which also participates in trichome formation. Suppression ofWoorSlCycB2expression by RNAi decreased the number of type I trichomes, and direct protein–protein interaction was detected between them, implying that both proteins may work together in the regulation of this type of trichome formation. Cytological observation andWotranscript analysis in the developing seeds showed that embryo development was also correlated withWo.

Plants have evolved and adapted to different environments. Dwarfism is an adaptive trait of plants that helps them avoid high-energy costs under unfavourable conditions. The role of gibberellin (GA) in plant development has been well established. Several plant dehydration-responsive element-binding proteins (DREBs) have been identified and reported to be induced under abiotic and biotic stress conditions. A tomato DREB gene named SlDREB, which is a transcription factor and was cloned from cultivated tomato M82, was found to play a negative role in tomato plant architecture and enhances drought tolerance. Tissue expression profiles indicated that SlDREB was expressed mainly in the stem and leaf and could be induced by abscisic acid (ABA) but suppressed by GA and ethylene. SlDREB altered plant morphology by restricting leaf expansion and internode elongation when overexpressed, and the resulting dwarfism of tomato plants could be recovered by application of exogenous gibberellic acid (GA₃). Transcriptional analysis of transgenic plants revealed that overexpression of SlDREB caused the dwarf phenotype by downregulating key genes involved in GA biosynthesis such as ent-copalyl diphosphate synthase (SlCPS) and GA 20-oxidases (SlGA20ox1, -2, and -4), thereby decreasing endogenous GA levels in transgenic plants. A yeast activity assay demonstrated that SlDREB specifically bound to dehydration-responsive element/C-repeat (DRE/CRT) elements of the SlCPS promoter region. Taken together, these data demonstrated that SlDREB can downregulate the expression of key genes required for GA biosynthesis and that it acts as a positive regulator in drought stress responses by restricting leaf expansion and internode elongation.

The B-BOX (BBX) proteins encode a class of zinc-finger transcription factors possessing one or two B-BOX domains and in some cases an additional CCT (CO, CO-like, and TOC1) motif, which play important roles in regulating plant growth, development and stress response. Nevertheless, no systematic study of BBX genes has been undertaken in tomato (Solanum lycopersicum). Here we present the results of a genome-wide analysis of the 29 BBX genes in this important vegetable species. Their structures, conserved domains, phylogenetic relationships, subcellular localizations, and promoter cis-regulatory elements were analyzed; their tissue expression profiles and expression patterns under various hormones and stress treatments were also investigated in detail. Tomato BBX genes can be divided into five subfamilies, and thirteen of them were found to be segmentally duplicated. Real-time quantitative PCR analysis showed that most BBX genes exhibited different temporal and spatial expression patterns, such as SlBBX24, which was constitutively expressed with high abundance in nearly all tissues studied, and eight genes (SlBBX4, 5, 6, 11, 16, 18, 19, and 22) showed relatively higher expression levels in vegetative tissues and non-ripening fruits. The expression of most BBX genes can be induced by drought, polyethylene glycol-6000 or heat stress. Some BBX genes were induced strongly by phytohormones such as abscisic acid, gibberellic acid or ethephon. The majority of tomato BBX proteins was predicted to be located in nuclei, and the transient expression assay using Arabidopsis mesophyll protoplasts demonstrated that all the seven BBX members tested (SlBBX5, 7, 15, 17, 20, 22 and 24) were localized in nucleus. Our analysis of tomato BBX genes on the genome scale would provide valuable information for future functional characterization of specific genes in this family.

The basic leucine zipper (bZIP) transcription factors have crucial roles in plant stress responses. In this study, the bZIP family gene SlbZIP38 (GenBank accession No: XM004239373) was isolated from a tomato (Solanum lycopersicum cv. Ailsa Craig) mature leaf cDNA library. The DNA sequence of SlbZIP38 encodes a protein of 484 amino acids, including a highly conserved bZIP DNA-binding domain in the C-terminal region. We found that SlbZIP38 was differentially expressed in various organs of the tomato plant and was downregulated by drought, salt stress, and abscisic acid (ABA). However, overexpression of SlbZIP38 significantly decreased drought and salt stress tolerance in tomatoes (Ailsa Craig). The findings that SlbZIP38 overexpression reduced the chlorophyll and free proline content in leaves but increased the malondialdehyde content may explain the reduced drought and salt tolerance observed in these lines. These results suggest that SlbZIP38 is a negative regulator of drought and salt resistance that acts by modulating ABA signaling.

Drought stress is the major abiotic factor that can seriously affect plant growth and crop production. The functions of flavin-containing monooxygenases (FMOs) are known in animals. They add molecular oxygen to lipophilic compounds or produce reactive oxygen species (ROS). However, little information on FMOs in plants is available. Here, we characterized a tomato drought-responsive gene that showed homology to FMO, and it was designated as FMO1. FMO1 was downregulated promptly by drought and ABA treatments. Transgenic functional analysis indicated that RNAi suppression of the expression of FMO1 (FMO1-Ri) improved drought tolerance relative to wild-type (WT) plants, whereas overexpression of FMO1 (FMO1-OE) reduced drought tolerance. The FMO1-Ri plants exhibited lower ABA accumulation, higher levels of antioxidant enzyme activities, and less ROS generation compared with the WT and FMO1-OE plants under drought stress. RNA-seq transcriptional analysis revealed the differential expression levels of many drought-responsive genes that were co-expressed with FMO1, including PP2Cs, PYLs, WRKY, and LEA. Using Y2H screening, we found that FMO1 physically interacted with catalase 2 (CAT2), which is an antioxidant enzyme and confers drought resistance. Our findings suggest that tomato FMO1 negatively regulates tomato drought tolerance in the ABA-dependent pathway and modulates ROS homeostasis by directly binding to SlCAT2.