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Large-scale analysis of MYB genes in Cucurbitaceae identifies a novel gene regulating plant height.

The MYB transcription factor (TF) family, which is involved in plant growth and development, is large and diverse. Previous studies on MYB family in Cucurbitaceae were mostly based on a single genome or focused on the R2R3 subfamily. Here, we analyzed 91 genomes of 11 Cucurbitaceae species and identified a total of 15 858 MYB genes. According to phylogenetic relationships, these genes were divided into 27 subgroups. The identified MYB genes were further classified into 121 MYB orthologous gene groups (OGGs), including 25 core, 57 softcore, 19 shell and 20 line-specific/cloud groups. Whole-genome duplication was the most common mechanism of MYB genes expansion. In core group, the higher proportions of MYB genes were found to be in the coexpression network constructed by the RNA-seq data. Through the comprehensive analysis including phylogeny and gene expression profile of cucumber MYB genes, as well as genetic variations in 103 cucumber germplasms, we identified a MYB gene CsRAX5, which may be related to cucumber plant height. We used gene editing technology to knockout and overexpress CsRAX5. In the knockout lines, Csrax5, the height was significantly increased compared with wild type (WT), whereas after overexpression the height of CsRAX5-OE plants was significantly decreased compared with WT. These results indicated that MYB gene CsRAX5 negatively regulated cucumber plant height. The large-scale analysis of MYB genes in Cucurbitaceae in this study provides insights for further investigating the evolution and function of MYB genes in Cucurbitaceae crops.

Journal Article

Tandem Duplication-Driven Neofunctionalization of UDP-Glycosyltransferases Shapes the Diversification of Triterpenoid Saponins in the Cucurbitaceae.

Tandem duplication of tailoring enzymes allows evolutionary innovation that diversifies plant specialized metabolism. Here, we present an interesting example of how tandem duplicated UDP-glycosyltransferases undergo neofunctionalization and shape the chemical diversity of triterpenoid saponins in the Cucurbitaceae family. A chromosome-level genome of Siraitia grosvenorii was assembled and aligned with multiple cucurbit genomes, revealing a specific UGT73AM tandem duplication responsible for regio-selective glycosylation (e.g. the rare 1,4-linked disaccharide) of diverse saponins such as mogrosides, ginsenosides, and momordicines. Comparative genomics depicted the evolutionary trajectory of a universal saponin-biosynthesizing UGT73 tandem arrays syntenously preserved across core eudicots, where lineage-specific UGT copies contribute to distinct metabolic phenotypes. A crystal structure of SgUGT73AM30 (mogrol 25-O-glycosyltransferase) in complex with UDP and mogrol was obtained to elucidate the molecular basis of the regio-specific decoration on vicinal diol of the substrates. Altogether, these findings provide insights into tandem duplication-driven diversification of glycosyltransferases and lay the foundation for engineered glycosylation of valuable triterpenoid saponins.

Saponins

Cucurbit Leaf Crumple Virus: An Important Pathogen of Cucurbit and Snap Bean Crops.

TAXONOMY: Cucurbit leaf crumple virus (CuLCrV); Begomovirus cucurbitae; Geminiviridae; Geplafuvirales. GEOGRAPHICAL DISTRIBUTION: The presence of CuLCrV is exclusively limited to North America, mainly Mexico and the United States. PHYSICAL PROPERTIES: CuLCrV is a bipartite begomovirus comprising two circular single-stranded DNA molecules (DNA-A and DNA-B), encapsidated within geminate icosahedral particles. GENOME AND ORGANIZATION: CuLCrV possesses a bipartite genome of DNA-A (2632 nucleotides) and DNA-B (2600 nucleotides). DNA-A contains five open reading frames (ORFs): AV1 (coat protein), AC1 (replication-associated protein), AC2 (transcriptional activator protein), AC3 (replication enhancer protein) and AC4. DNA-B contains two ORFs: BV1 (nuclear shuttle protein) and BC1 (movement protein). TRANSMISSION: CuLCrV is transmitted by the sweetpotato whitefly, Bemisia tabaci, in a persistent, circulative and non-propagative manner. HOSTS: CuLCrV primarily infects crop members of the Cucurbitaceae and snap bean (Phaseolus vulgaris, Fabaceae). Multiple weed species belonging to Brassicaceae, Convolvulaceae, Cucurbitaceae and Verbenaceae act as persistent virus reservoir hosts. SYMPTOMS: Symptom expression varies with host and infection timing. In cucurbits, infection induces leaf crumpling, thickening and downward curling of leaves, with green streaks and distortion of fruits. In snap bean, symptoms include leaf distortion, chlorosis and malformed pods. CONTROL: No commercial cultivars with resistance to CuLCrV are available for cucurbit crops, although some resistance has been reported in snap bean cultivars. Therefore, management relies primarily on integrated disease management.

Plant Diseases

Deciphering differential mRNA and lncRNA expression profiles in response to PEG simulated drought stress in cucumber (Cucumis sativus L.).

Cucumber (Cucumis sativus L.), a vital fruit vegetable of the Cucurbitaceae family, originated in India ∼ 3000 years ago. It is widely used in the culinary, therapeutic, and cosmetic sectors. Cucumber cultivation is significantly impacted by drought stress, especially in arid and semi-arid regions. This study investigates the molecular response to drought using two contrasting cucumber lines: WBC-23-2 (drought-tolerant) and DGPC-59 (drought-sensitive). Drought was simulated using polyethylene glycol (PEG), and effects on physiological and biochemical traits were evaluated. The tolerant line exhibited reduced leaf wilting and higher relative water content (RWC). Based on these physiological markers, transcriptomic profiling was employed to identify the underlying regulatory networks. Analysis identified 4,736 DEGs, suggesting that the tolerant line's superior resilience is driven by preferential activation of genes involved in photosynthesis and glutathione metabolism. Conversely, the sensitive genotype showed enrichment in organonitrogen compound catabolism and water deprivation response. This divergence is further reflected in the regulation of 155 transcription factors (TFs) across various families, indicating distinct regulatory architectures between the two lines. Additionally, 774 drought-responsive long non-coding RNAs (lncRNAs) were identified, acting via cis, trans, and competing endogenous RNA (ceRNA) mechanisms to modulate gene expression. Key candidate genes associated with drought tolerance included WAT1-related protein At5g64700, thaumatin-like protein, berberine bridge enzyme-like 18, probable WRKY transcription factor, and pathogenesis-related protein 1. This study reveals a complex regulatory network of mRNAs, lncRNAs, and TFs underlying drought response and provides a valuable foundation for breeding drought-resilient cucumber cultivars. A web-based genomic resource, CsDTDb, has been developed and made publicly available to facilitate future functional genomics studies related to drought tolerance in cucumber.

DEGs

Host Range and Chemical Control of Cercospora citrullina, the Causal Agent of Watermelon Spot Disease.

This study systematically evaluated cultivation requirements, host range, and chemical control options for Cercospora citrullina causing watermelon spot disease. Among 11 chemically defined media tested, corn meal agar medium supported optimal mycelial growth of C. citrullina, with an average radial growth rate of 56.72 ± 1.45 mm under controlled conditions (25°C, darkness). Host range determination via artificial inoculation of 19 plant species confirmed that the host range of the strain UNL090101 is limited to the Cucurbitaceae species tested, with watermelon (Citrullus lanatus) exhibiting the highest susceptibility, followed by melon (Cucumis melo) and cucumber (Cucumis sativus). Fungicide screening of 17 commercial formulations identified 40% iminoctadine tris (albesilate) WP (EC50 = 2.82 μg·liter-1) and 64% mancozeb + 8% cymoxanil (WS) (EC50 = 48.75 μg·liter-1) as the most effective treatments, achieving control efficacies of 74.47 and 58.62%, respectively. These findings provide actionable guidelines for optimizing crop rotation, intercropping strategies, and fungicide selection in watermelon production systems.

Cercospora citrullina

Development of a highly efficient prime editing platform for cucurbits enables breeding of multi-disease-resistant cucumber.

The prime editing (PE) system is a precise genome editing technology that works efficiently in monocots; however, its application is limited by low editing efficiency in dicots, particularly Cucurbitaceae and Solanaceae plants. Here, we first significantly improved the transformation efficiency by introducing spectinomycin in cucurbits, then used the tomato elongation factor 1-alpha (SlEF1α) promoter to enhance PE protein expression, and incorporated the Csy4 ribonuclease to process pegRNAs, collectively addressing multiple constraints limiting PE efficiency in cucurbits. The optimized PE systems, particularly Csy4-PE6d, achieved an average desired editing frequency of 80.83% at targeted loci in cucumber via stable genetic transformation, with frequencies reaching up to 100% at certain sites. Moreover, Csy4-PE6d generated homozygous edits in 36.43% of transgenic lines and demonstrated robust editing activity in melon, pumpkin, and potato. Using the Csy4-PE6d tool, we generated heritable edited cucumber lines with dual resistance to bacterial angular leaf spot and downy mildew by targeting the CsSGR gene. Collectively, this optimized system substantially enhances PE efficiency in Cucurbit crops, providing an effective solution to common challenges such as low editing efficiency and limited heritability in these species.

Disease Resistance

Sechium edule: Phytochemistry, Biological Activities, Potential Health Effects, Food-Industry Applications, and Future Perspectives.

Sechium edule (Jacq.) Sw. (chayote), a neglected and underutilized Cucurbitaceae crop widely cultivated across tropical and subtropical regions, has drawn growing interest as a source of health-promoting food components. This review critically synthesizes studies published between 2000 and 2026 on its botanical features, genome characterization, nutritional value, phytochemistry, bioactivities, safety, and food-industry applications, based on literature retrieved from PubMed, Scopus, Web of Science, ScienceDirect, and the Cochrane Library. Different plant parts (fruits, leaves, seeds, tuberous roots, and peels) contain diverse bioactive compounds, including flavonoids, phenolic acids, cucurbitacins, pectin polysaccharides, and carotenoids. Reported bioactivities include antioxidant, anti-inflammatory, hypoglycemic, cardioprotective, antiproliferative, and geroprotective effects, mediated in part through Nrf2-mediated antioxidant signaling and sirtuin (SIRT1/3/5/6) upregulation. Notably, a systematic meta-analysis demonstrated a significant reduction in serum glucose (MD = -20.56; 95% CI: -29.35 to -11.77) and HbA1c following three months of chayote intake in patients with metabolic syndrome and type 2 diabetes. Industrially, chayote has been developed into fermented products, starch- and peel-derived bioactive films, ultrasound-extracted pectin, α-amylase inhibitory seed protein isolates, and probiotic encapsulation systems. Recent genomic work has further revealed a chromosome-level genome assembly, whole-genome duplication events, and a domestication history tracing to Mexico's Oaxaca region. Collectively, this evidence positions chayote as a promising underutilized resource for food, nutraceutical, and biomedical use, while highlighting key gaps: the need for standardized clinical trials, bioavailability studies, and comprehensive safety evaluation.

Sechium edule

Cucurbitacins in Plant-Insect Interactions: Biosynthesis, Regulation, Ecological Functions, and Prospects for Crop Protection.

Cucurbitacins are highly oxygenated tetracyclic triterpenoids characterized by intense bitterness, substantial structural diversity, and important consequences for plant-herbivore interactions. Although best known from Cucurbitaceae, cucurbitacins and related cucurbitane-type metabolites also occur in phylogenetically distant herbaceous and woody plants. Genetic and biochemical studies have validated several core biosynthetic steps, including cucurbitadienol formation by oxidosqualene cyclases and subsequent modification by cytochrome P450 monooxygenases, acyltransferases, and glycosyltransferases. Tissue-preferential basic helix-loop-helix transcription factors constitute the best-characterized regulatory layer, whereas the evidence supporting accessory regulators, transporters, and environmental responses varies from functional validation to transcriptomic or genomic prediction. From the plant perspective, cucurbitacins deter feeding or impair performance in many generalist and non-adapted herbivores. By contrast, their use as host-recognition cues and feeding stimulants by specialist diabroticite beetles reflects evolved herbivore adaptations involving perception, tolerance, metabolism, or sequestration rather than a second defensive function of the plant trait. Herbivore-induced cucurbitacin accumulation has been demonstrated in particular systems, although its regulatory mechanisms and ecological generality remain unresolved. Unlike previous reviews centered primarily on cucurbitacin chemistry, pharmacological activity, or individual biosynthetic pathways, this review integrates evidence-graded pathway reconstruction and molecular regulation with taxonomic distribution, insect adaptation, domestication, and agroecological consequences. Mechanistically, this review traces how scaffold formation, oxidative tailoring, conjugation, tissue-specific regulation, and transport give rise to contrasting ecological outcomes through herbivore-specific perception, tolerance, metabolism, and sequestration. We conclude that uniformly increasing or eliminating cucurbitacins is unlikely to provide broadly effective crop resistance because either direction may favor a different herbivore group. Future priorities include functional validation of candidate genes, spatially resolved metabolite analysis, comparative investigation of non-cucurbit lineages, and field evaluation involving generalist and specialist herbivores, crop quality, and non-target organisms. These advances will support context-specific fruit-quality improvement, behavioral pest control, and integrated pest management strategies rather than cucurbitacin manipulation as a stand-alone resistance approach.

agroecology

Ancient DNA reveals early use of melons in China's Song dynasty.

Melon (Cucumis melo L.) domestication is thought to have occurred independently once in Northeast Africa and twice in India, but archaeobotanical seed remains point to a possible additional domestication event in China. Because Cucumis seeds are difficult to diagnose morphologically, genomic data from archaeological material are needed to evaluate these scenarios and reconstruct ancient melon traits. We sequenced two Song Dynasty (960-1279 CE) melon seeds from Shuomen Gugang (China), recovering 5.5× and 2.1× nuclear genome coverage. Nuclear and chloroplast analyses place both seeds within cultivated C. melo from China, within the "agrestis" East Asian gene pool. To assess whether these seeds carried traits associated with sweet dessert melons, we examined loci underlying fruit phenotypes. Neither seed carried alleles for orange flesh; one harbored an allele linked to yellow/orange peel, the other possessed alleles associated with green flesh and reduced acidity. Since wild melons are monoecious, the presence of the derived andromonoecy allele in one seed, associated with rounder fruit shape, suggests early selection on fruit morphology. Together, these findings indicate that Song Dynasty melons were likely consumed as fresh or culinary fruits rather than sweet dessert melons. Their flesh coloration resonates with Song-period aesthetic sensibilities, exemplified by jade-green celadon ceramics frequently crafted in melon-shaped forms. By anchoring East Asian archaeobotanical remains within modern melon genomic variation, this study provides a temporal framework for melon cultivation in China and shows how ancient genomics can illuminate past crop use.

China

Pleiotropic mutation in a tendril TCP gene underlies the yield-enhancing multiple-flowering trait in summer squash (Cucurbita pepo).

Crop yield is a focal point in plant breeding. Regulation of lateral budding through apical dominance was a central target of crop domestication, directly affecting crop production. The young fruits of Cucurbita pepo, summer squash, are produced on plants characterized by apical dominance and differentiation of a single flower bud per leaf axil. A single recessive mutation, mf, results in differentiation of more than one flower per leaf axil, thereby directly increasing production because of the continual day-to-day harvest of the summer squash crop. Positional cloning of the Cucurbita pepo mf (Cpmf) gene denoted a frameshift mutation in a TCP transcription factor, Cp4.1LG13g07780, as causative for the increase in axillary flowering. Cpmf is an ortholog of a tendril-development TCP gene in other cucurbits, and likewise, the recessive allele of Cpmf is associated with distorted tendril development. Gene function is context dependent, and we propose that multiple flowering is a unique pleiotropic attribute of mutation in a tendril-development gene of C. pepo. Characterization of a C. pepo collection confirmed a significant association of the Cpmf mutation with multiple flowering and showed that the mutant allele is absent in ancestral C. pepo and one of its two cultivated subspecies. The beneficial mutation occurred and was selected after the domestication of the other subspecies, during its cultivation for young fruit production. We demonstrate the discovery of a causative yield-increasing sequence variant and its practical utilization in breeding. Our findings provide a molecular target for creation of high-yielding, multiple-flowering summer squash cultivars through marker-assisted breeding or precise genome editing.

Cucurbita