The HSF (heat shock factor) gene family contains highly conserved plant-specific transcription factors that play an important role in plant high-temperature stress responses. The present study aimed to characterize the HSF transcription factor genes in tomato (Solanum lycopersicum), which is an important vegetable crop worldwide and the model plant for fruit development studies. Twenty-six SlyHSF genes were identified in tomato, and the phylogenetic analysis showed the possible evolution profile of subgroups among in the plant kingdom. A new group O was identified that involved HSF genes in primitive plant species, like in the green algae, mosses and lycophytes. The gene structure and motifs of each SlyHSF were comprehensively analyzed. We identified orthologous, co-orthologous and paralogous HSF gene pairs in tomato, Arabidopsis and rice, and constructed a complex interaction network among these genes. The SlyHSF genes were expressed differentially in different species and at a higher level in mature fruits. The qPCR analysis was performed and showed SlyHSF genes greatly participate in plant heat tolerant pathways. Our comprehensive genome-wide analysis provided insights into the HSF gene family of tomatoes.

Abiotic stress adversely inhibits the growth and development of plants, by changing the expression of multiple genes. Circular RNAs (circRNAs), as a class of non-coding RNAs, function in transcriptional and posttranscriptional regulation. Yet, the involvement of circRNAs in abiotic stress response is rarely reported. In this study, the participation and function of circRNAs in low-temperature (LT)-induced stress response were investigated in tomato leaves. We generated genome-wide profiles of circRNAs and mRNAs in tomato leaves grown at 25°C room temperature (RT) and 12°C LT. Our results show that 1,830 circRNAs were identified in tomato leaves in both RT and LT treatments, among which 1,759 were differentially induced by the LT treatment. We find that the identified circRNAs are mainly located at exons of genes, but less distributed at introns of genes or intergenic regions. Our results suggest that there are 383 differentially expressed circRNAs predicted to function as putative sponges of 266 miRNAs to target 4,476 mRNAs in total. Moreover, Gene Ontology (GO) and Kyoto Encyclopedia of Genes and Genomes (KEGG) analysis assays indicate that multiple pathways were enriched in both differentially expressed genes induced by LT and parental genes of differentially expressed circRNAs induced by LT, revealing the key functions of circRNAs and the corresponding targeted genes in response to LT stress. Our results suggest that circRNAs may be involved in regulating metabolism (i.e., carbohydrate, amino acid, lipid, and energy), signal transduction, and environmental adaptation-related pathways and that these circRNAs were predicted to regulate the expression of transcription factors, genes in signal transduction pathways, and genes related to the Ca²⁺ channel through targeting the corresponding proteins, such as WRKY, NAC, cytochrome P450, and calmodulin binding protein. Taken together, our study uncovers that multiple circRNAs are isolated and differently regulated in response to LT stress and provides the resource and potential networks of circRNA–miRNA–mRNA under LT stress for further investigations in tomato leaves.

Low temperature is a type of abiotic stress affecting the tomato (Solanum lycopersicum) growth. Understanding the mechanisms and utilization of exogenous substances underlying plant tolerance to cold stress would lay the foundation for improving temperature resilience in this important crop. Our study is aiming to investigate the effect of exogenous glycine betaine (GB) on tomato seedlings to increase tolerance to low temperatures. By treating tomato seedlings with exogenous GB under low temperature stress, we found that 30 mmol/L exogenous GB can significantly improve the cold tolerance of tomato seedlings. Exogenous GB can influence the enzyme activity of antioxidant defense system and ROS levels in tomato leaves. The seedlings with GB treatment presented higher Fv/Fm value and photochemical activity under cold stress compared with the control. Moreover, analysis of high-throughput plant phenotyping of tomato seedlings also supported that exogenous GB can protect the photosynthetic system of tomato seedlings under cold stress. In addition, we proved that exogenous GB significantly increased the content of endogenous abscisic acid (ABA) and decreased endogenous gibberellin (GA) levels, which protected tomatoes from low temperatures. Meanwhile, transcriptional analysis showed that GB regulated the expression of genes involved in antioxidant capacity, calcium signaling, photosynthesis activity, energy metabolism-related and low temperature pathway-related genes in tomato plants. In conclusion, our findings indicated that exogenous GB, as a cryoprotectant, can enhance plant tolerance to low temperature by improving the antioxidant system, photosynthetic system, hormone signaling, and cold response pathway and so on.

The conserved histone variant H2A.Z is essential for transcriptional regulation; defense responses; and various biological processes in plants, such as growth, development, and flowering. However, little is known about how H2A.Z affects the developmental process and ripening of tomato fruits. Here, we utilized the CRISPR/Cas9 gene-editing system to generate a sl_hta9 sl_hta11 double-mutant, designated sl_h2a.z, and found that these two mutations led to a significant reduction in the fresh weight of tomato fruits. Subsequent messenger RNA (mRNA)-seq results showed that dysfunction of Sl_H2A.Z has profound effects on the reprogramming of genome-wide gene expression at different developmental stages of tomato fruits, indicating a ripening-dependent correlation between Sl_H2A.Z and gene expression regulation in tomato fruits. In addition, the expression of three genes, SlPSY1, SlPDS, and SlVDE, encoding the key enzymes in the biosynthesis pathway of carotenoids, was significantly upregulated in the later ripening stages, which was consistent with the increased contents of carotenoids in sl_h2a.z double-mutant fruits. Overall, our study reveals a role of Sl_H2A.Z in the regulation of carotenoids and provides a resource for the study of Sl_H2A.Z-dependent gene expression regulation. Hence, our results provide a link between epigenetic regulation via histone variants and fruit development, suggesting a conceptual framework to understand how histone variants regulate tomato fruit quality.

Heat stress (HS) has been considered as a severe threat to crop yields in recent years. Sucrose, as a major product of photosynthesis, plays an important role in plant growth and stress response. Sucrose phosphate synthase (SPS) is a key rate-limiting enzyme in the sucrose synthesis pathway in plants. However, its molecular mechanism and signaling pathway remain unclear. In this study, we identified a novel SPS gene (SlSPS) in tomato and generated over-expression and knock-out of SlSPS gene transgenic tomato plants to investigate its biological functions related to the growth and thermotolerance of tomato. Over-expression of SlSPS gene increased the growth and biomass of transgenic tomato plants, such as fresh weight, dry weight, plant height, fruit weight and root length. In contrast, knock-out of SlSPS gene decreased the growth and biomass of transgenic tomato plants. Under heat stress, the survival rates were positively correlated with the expression level of SlSPS gene in different tomato varieties. Furthermore, SlSPS-overexpressing tomato plants showed higher SPS activity and sucrose content and heat stress resistant phenotypes. By comparison, knock-out tomato plants showed lower SPS activity and sucrose content and susceptible to heat stress. The determination of several reference values of oxidative stress parameters were also consistent with their heat resistance of these transgenic plants. In summary, SlSPS gene could positively mediate the growth and thermotolerance in tomato plants.

Waterlogging and heavy mental (e.g., cadmium) stress are two primary threats to crop growth. The combination of abiotic stresses was common and frequent, especially in the field condition. Even though the effects of individual waterlogging and cadmium on tomato plants have been widely investigated, the response of tomatoes under combined waterlogging and cadmium stress remains unclear. This study aimed to clarify and compare physiological, biochemical characteristics and plant growth of two tomato genotypes under individual and combined stress. Two tomato genotypes (‘MIX-002’ and ‘LA4440’) were treated under control, waterlogging, cadmium stress and their combination. The results showed that chloroplast ultrastructure of tomatoes under individual and combined stress was damaged with disordered stroma and grana lamellae. The H₂O₂ (hydrogen peroxide) content and O₂^·− (superoxide anion radical) production rate of plants under all the three stresses was not significantly higher than the control except for ‘LA4440’ under the combined stress. Antioxidant enzymes actively responded in the two tomato genotypes, as shown by significant increase in SOD activity from ‘MIX-002’ under waterlogging and combined stress and from ‘LA4440’ under cadmium. Meanwhile, CAT activity of ‘MIX-002’ under waterlogging and ‘LA4440′ under combined stress significantly decreased, and the POD activity of ‘MIX-002’ under combined stress significantly increased as compared with the respective control. The APX activity of ‘MIX-002’ and ‘LA4440’ under combined stress was significantly lower and higher than the respective controls. This indicated that tomato plants were able to secure redox homeostasis and protect plants from oxidative damage through the synergetic regulation of antioxidant enzymes. Plant height and biomass of the two genotypes under individual and combined stress significantly decreased, which could be a direct result from the chloroplast alteration and resource re-allocation. Overall, the effects of combined waterlogging and cadmium stress were not simply the sum of individual effects on two tomato genotypes. Distinct ROS (reactive oxygen species) scavenging systems of two tomato genotypes under stresses suggest a genotype-dependent antioxidant enzymes regulation.