Super-pangenome analyses highlight genomic diversity and structural variation across wild and cultivated tomato species.

Effective utilization of wild relatives is key to overcoming challenges in genetic improvement of cultivated tomato, which has a narrow genetic basis; however, current efforts to decipher high-quality genomes for tomato wild species are insufficient. Here, we report chromosome-scale tomato genomes from nine wild species and two cultivated accessions, representative of Solanum section Lycopersicon, the tomato clade. Together with two previously released genomes, we elucidate the phylogeny of Lycopersicon and construct a section-wide gene repertoire. We reveal the landscape of structural variants and provide entry to the genomic diversity among tomato wild relatives, enabling the discovery of a wild tomato gene with the potential to increase yields of modern cultivated tomatoes. Construction of a graph-based genome enables structural-variant-based genome-wide association studies, identifying numerous signals associated with tomato flavor-related traits and fruit metabolites. The tomato super-pangenome resources will expedite biological studies and breeding of this globally important crop.

Genome-wide identification and analysis of the evolution and expression pattern of the HVA22 gene family in three wild species of tomatoes.

Wild tomato germplasm is a valuable resource for improving biotic and abiotic stresses in tomato breeding. The HVA22 is widely present in eukaryotes and involved in growth and development as well as stress response, such as cold, salt, drought, and biotic stress. In the present study, we identified 45 HVA22 genes in three wild species of tomatoes. The phylogenetic relationships, gene localization to chromosomes, gene structure, gene collinearity, protein interactions, and cis-acting element prediction of all 45 HVA22 genes (14 in Solanum pennellii, 15 in S. pimpinellifolium, and 16 in S. lycopersicoides) were analyzed. The phylogenetic analysis showed that the all HVA22 proteins from the family Solanaceae were divided into three branches. The identified 45 HVA22 genes were grouped into four subfamilies, which displayed similar number of exons and expanded in a fragmentary replication manner. The distribution of HVA22 genes on the chromosomes of the three wild tomato species was also highly similar. RNA-seq and qRT-PCR revealed that HVA22 genes were expressed in different tissues and induced by drought, salt, and phytohormone treatments. These results might be useful for explaining the evolution, expression patterns, and functional divergence of HVA22 genes in Lycopersicon.

The LEA gene family in tomato and its wild relatives: genome-wide identification, structural characterization, expression profiling, and role of SlLEA6 in drought stress.

BACKGROUND: Late embryogenesis abundant (LEA) proteins are widely distributed in higher plants and play crucial roles in regulating plant growth and development processes and resisting abiotic stress. Cultivated tomato (Solanum lycopersicum) is an important vegetable crop worldwide; however, its growth, development, yield, and quality are currently severely constrained by abiotic stressors. In contrast, wild tomato species are more tolerant to abiotic stress and can grow normally in extreme environments. The main objective of this study was to identify, characterize, and perform gene expression analysis of LEA protein families from cultivated and wild tomato species to mine candidate genes and determine their potential role in abiotic stress tolerance in tomatoes. RESULTS: Total 60, 69, 65, and 60 LEA genes were identified in S. lycopersicum, Solanum pimpinellifolium, Solanum pennellii, and Solanum lycopersicoides, respectively. Characterization results showed that these genes could be divided into eight clusters, with the LEA_2 cluster having the most members. Most LEA genes had few introns and were non-randomly distributed on chromosomes; the promoter regions contained numerous cis-acting regulatory elements related to abiotic stress tolerance and phytohormone responses. Evolutionary analysis showed that LEA genes were highly conserved and that the segmental duplication event played an important role in evolution of the LEA gene family. Transcription and expression pattern analyses revealed different regulatory patterns of LEA genes between cultivated and wild tomato species under normal conditions. Certain S. lycopersicum LEA (SlLEA) genes showed similar expression patterns and played specific roles under different abiotic stress and phytohormone treatments. Gene ontology and protein interaction analyses showed that most LEA genes acted in response to abiotic stimuli and water deficit. Five SlLEA proteins were found to interact with 11 S. lycopersicum WRKY proteins involved in development or resistance to stress. Virus-induced gene silencing of SlLEA6 affected the antioxidant and reactive oxygen species defense systems, increased the degree of cellular damage, and reduced drought resistance in S. lycopersicum. CONCLUSION: These findings provide comprehensive information on LEA proteins in cultivated and wild tomato species and their possible functions under different abiotic and phytohormone stresses. The study systematically broadens our current understanding of LEA proteins and candidate genes and provides a theoretical basis for future functional studies aimed at improving stress resistance in tomato.

A calcium-dependent protein kinase gene SpCPK33 from Solanum pennellii associated with increased cold tolerance in tomato.

Calcium-dependent protein kinases (CDPKs, CPKs) represent a vital class of calcium sensors, which play a crucial role in plant growth, development and adaption to complex environmental stresses. Wild species tend to exhibit greater tolerance than cultivated species under environmental stress. Here, we isolated a calcium-dependent protein kinase gene SpCPK33 located primarily on the plasma membrane of abiotic-resistant species (Solanum pennellii LA0716). It was highly expressed in stems and leaves and was also induced by cold stress. Compared with WT plants, the overexpression of SpCPK33 in cultivated tomato (cv M82) enhanced its tolerance to cold stress. Transgenic lines demonstrated strong vitality under low temperature treatment. Moreover, the levels of malondialdehyde (MDA) and reactive oxygen species (ROS) were decreased in SpCPK33-overexpressing plants. The activities of antioxidant enzymes and the levels of osmotic regulatory substances were higher. The transcript levels of cold stress-related genes were up-regulated. In summary, the results indicate that SpCPK33-overexpressing transgenic plants experience less severe chilling injury under cold stress, and improved tomato cold tolerance by scavenging ROS accumulation and modulating the expression of stress-related genes.

Effects of vegetation and terrain changes on spatial heterogeneity of soil C-N-P in the coastal zone protected forests at northern China.

Soil organic carbon (SOC), total nitrogen (TN) and total phosphorus (TP) are important indicators reflecting soil quality, and they can be used to effectively evaluate the effect of soil remediation. Many studies have evaluated the content of SOC, TN and TP in different ecosystems. However, after constructing protected forests for ecological restoration in the ecologically fragile coastal zone, the spatial distribution and influencing mechanism of SOC, TN and TP content is still uncertain. In this study, the spatial heterogeneity and influencing factors of SOC, TN and TP in surface (0-20 cm) soil were analyzed by traditional analysis and geostatistics. A total of 39 soil samples were collected under the coastal zone protected forest types including Quercus acutissima Carruth (QAC), Pinus thunbergii Parl (PTP), mixed PTP and QAC (QP) and Castanea mollissima BL (CMB) in the coastal zone protected forests in northern China. The results show that SOC, TN and TP content were defined as moderate variation, and they also show significant changes under different protected forest types (P < 0.05). The semivariance results indicate that SOC, TN and TP all exhibited strong spatial dependence class, with Range of 224 m, 229 m and 282 m respectively, which were more than the sampling scale of 200 m. The spatial prediction results showed that SOC, TN and TP content all appear in large areas of extremely low value in CMB, and its cross validation results showed that using vegetation and terrain factors as covariates in the spatial prediction of SOC, TN and TP can improve the prediction accuracy. The results of correlation analysis showed that the influencing factor for SOC and TN, and TP were NDVI and topographical changes, respectively. In general, vegetation and terrain factors as auxiliary factors can improved the accuracy of soil C-N-P spatial distribution prediction after afforestation in coastal zone.

Genome-Wide Characterization of Salt-Responsive miRNAs, circRNAs and Associated ceRNA Networks in Tomatoes.

Soil salinization is a major environmental stress that causes crop yield reductions worldwide. Therefore, the cultivation of salt-tolerant crops is an effective way to sustain crop yield. Tomatoes are one of the vegetable crops that are moderately sensitive to salt stress. Global market demand for tomatoes is huge and growing. In recent years, the mechanisms of salt tolerance in tomatoes have been extensively investigated; however, the molecular mechanism through which non-coding RNAs (ncRNAs) respond to salt stress is not well understood. In this study, we utilized small RNA sequencing and whole transcriptome sequencing technology to identify salt-responsive microRNAs (miRNAs), messenger RNAs (mRNAs), and circular RNAs (circRNAs) in roots of M82 cultivated tomato and Solanum pennellii (S. pennellii) wild tomato under salt stress. Based on the theory of competitive endogenous RNA (ceRNA), we also established several salt-responsive ceRNA networks. The results showed that circRNAs could act as miRNA sponges in the regulation of target mRNAs of miRNAs, thus participating in the response to salt stress. This study provides insights into the mechanisms of salt tolerance in tomatoes and serves as an effective reference for improving the salt tolerance of salt-sensitive cultivars.