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Plant Physiol. Biochem. [JOURNAL]

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Genome-wide analysis of the LAR gene family and the role of OvLAR71 in proanthocyanidin biosynthesis in Onobrychis viciifolia.

Liu L, Wang J, Jiang G … +5 more , He H, Du P, Meng Z, Li H, Xie Q

Plant Physiol Biochem · 2026 Jun · PMID 42323918 · Publisher ↗

Sainfoin (Onobrychis viciifolia Scop.) is a valuable perennial forage legume abundant in proanthocyanidins (PAs). As important secondary metabolites in plants, PAs not only contribute to plant stress tolerance and defens... Sainfoin (Onobrychis viciifolia Scop.) is a valuable perennial forage legume abundant in proanthocyanidins (PAs). As important secondary metabolites in plants, PAs not only contribute to plant stress tolerance and defense processes, but also significantly improve forage quality. Leucoanthocyanidin reductase (LAR) is a key enzyme in the biosynthesis and accumulation of PAs. However, systematic research on the LAR gene family in sainfoin remains limited. Therefore, we conducted a genome-wide identification and systematic characterization of the LAR gene family in the legume forage species sainfoin. A total of 83 OvLAR genes were identified in the sainfoin genome, which were unevenly distributed across 28 chromosomes. Phylogenetic analysis classified the 83 OvLAR proteins into five evolutionary clades. Furthermore, segmental duplication served as the major driving force underlying the expansion of the OvLAR gene family. Cis-acting element profiling indicated that OvLAR gene expression is potentially regulated by both endogenous cues and environmental signals, suggesting that these genes are involved in complex regulatory networks that coordinate secondary metabolism and adaptive development. Consistently, qRT-PCR results revealed tissue-specific expression patterns among OvLAR genes, as well as differential transcriptional responses to multiple abiotic stresses. Functional assays further demonstrated that overexpression of OvLAR71 in tobacco and sainfoin significantly promoted PA biosynthesis and accumulation in leaves and flowers, accompanied by distinct changes in floral pigmentation. Collectively, this work provides the first comprehensive insight into the LAR gene family in sainfoin, offers critical insights into their roles in PA biosynthesis, and establishes a theoretical foundation for the genetic improvement of forage quality.

Highway to stress: Proline dynamics in tomato plants under osmotic disturbances.

Konrad C, Spormann S, Nadais P … +3 more , Martins M, Soares C, Fidalgo F

Plant Physiol Biochem · 2026 Jun · PMID 42314389 · Publisher ↗

Upon exposure to osmotic stress, plants accumulate proline, an osmoprotectant and antioxidant, often associated with stress tolerance. However, recent studies suggest that proline buildup does not necessarily improve the... Upon exposure to osmotic stress, plants accumulate proline, an osmoprotectant and antioxidant, often associated with stress tolerance. However, recent studies suggest that proline buildup does not necessarily improve the redox status or stress resilience in plant tissues, being instead perceived as a sign of sensitivity. Therefore, this study aimed to elucidate how plants coordinate proline accumulation and metabolism during osmotic stress and subsequent recovery. A bifactorial experimental design was conducted using tomato plants in a semi-hydroponic system. Tissue samples were collected in three moments: t0: after 20 days of growth under control conditions; t1: after 4 days of exposure to 8% (w/v) polyethylene glycol (PEG); t2: after one week of recovery under control conditions. PEG-induced stress triggered proline accumulation. Despite this pronounced buildup, exposure to PEG delayed growth and caused oxidative damage. Following recovery, proline decreased in roots despite persistent biosynthetic activity, whereas in shoots, proline remained high despite a decline of the biosynthetic activity, coinciding with oxidative damage. Expression and activity of P5CS and ProDH illustrate that proline metabolism shows differential transcriptional and post-transcriptional regulation. Results also indicate the involvement of the ornithine pathway in proline accumulation in shoots, and regulation of proline content via distribution among organs, besides metabolic homeostasis. Overall, proline appears to act as a compatible solute in roots with rapid but transient accumulation, while its sustained and massive accumulation in shoots does not confer effective antioxidant protection or stress tolerance. This supports previous observations that proline may function as a sign of stress severity rather than a tolerance mechanism.

BolMYB122-BolTGG1 module enhances salt stress tolerance via accelerating the hydrolysis of glucosinolates to produce sulforaphane in broccoli.

Wang J, Sun W, Tao J … +7 more , Fang Y, Wang X, Li Y, Ma C, Deng L, Li H, Wang C

Plant Physiol Biochem · 2026 Jun · PMID 42314388 · Publisher ↗

Glucosinolates (GSLs), one type of secondary metabolites mainly enriched in cruciferous plants, play important roles in environmental stress responses. However, the function and regulatory mechanism of GSLs in combating... Glucosinolates (GSLs), one type of secondary metabolites mainly enriched in cruciferous plants, play important roles in environmental stress responses. However, the function and regulatory mechanism of GSLs in combating salt stress in broccoli are largely unclear. In this study, BolTGG1, a member of β-thioglucoside glucohydrolase (TGG) genes mainly functioning to hydrolyze GSLs, was identified in broccoli. BolTGG1 expression was upregulated under salt stress. Overexpression of BolTGG1 significantly improved the tolerance to salt stress in both broccoli and Arabidopsis. Subsequently, BolTGG1 was heterologously expressed in yeast and the BolTGG1 protein was isolated to evaluate its ability to hydrolyze GSLs. The results confirmed that exogenous addition of BolTGG1 could obviously increase the yield of sulforaphane (SFN), a major hydrolysis product of GSLs, in vitro. Meanwhile, the content of SFN was significantly increased in BolTGG1-overexpression transgenic broccoli plants. The role of SFN in the salt stress response was then explored. The results indicated that external application of SFN could directly enhance the salt resistance with the elevated activities of superoxide dismutase (SOD) and peroxidase (POD), and reduced malondialdehyde (MDA) levels in broccoli. Furthermore, BolMYB122 was confirmed to be the upstream regulatory transcription factor of BolTGG1. BolMYB122 positively regulated BolTGG1 transcription. In summary, these results indicated that BolMYB122-BolTGG1 module confers salt stress tolerance mainly by accelerating the generation of SFN. These findings provide new insights into the role and regulatory mechanism of GSL metabolites in plant salt stress response, and suggest a novel breeding strategy for stress resistance from the perspective of regulating plant secondary metabolism.

PHYTOCHROME INTERACTING FACTORS act upstream to coordinate microtubule dynamics and differential cell growth during Arabidopsis apical hook opening.

Burachik NB, Vacs P, Santin F … +2 more , Mazzella MA, González-Schain N

Plant Physiol Biochem · 2026 Jun · PMID 42314387 · Publisher ↗

Differential cell growth, a key evolutionary strategy, involves unequal cell expansion to bend organs like stems or roots, enabling directional growth to optimize access to resources and avoid harmful conditions. Apical... Differential cell growth, a key evolutionary strategy, involves unequal cell expansion to bend organs like stems or roots, enabling directional growth to optimize access to resources and avoid harmful conditions. Apical hook development in darkness is an excellent model to study asymmetric elongation between the convex and concave sides of the hypocotyl during its two distinctive phases: formation and opening. While PHYTOCHROME INTERACTING FACTORS (PIFs) are recognized as essential for proper apical hook development, the underlying cellular basis remains largely unknown. Here, we show that PIF activity is necessary to sustain the coordination of cortical microtubule (cMT) organization (involved in orchestrating cell growth anisotropy) with cell expansion during apical hook opening in darkness under our imaging conditions. In the pifq mutant this coordination is lost, as both apical hook and apical hypocotyl cells undergo excessive elongation despite predominantly longitudinal cMT alignment and significant alignment strength, resulting in defective differential growth. RNA-seq reanalysis revealed the reduced expression of genes associated with cell wall biosynthesis and remodeling, as well as the upregulation of cMT remodeling factors. This work expands the classical view of PIFs as transcriptional regulators of elongation-related genes, identifying them as central coordinators of cytoskeletal organization and directional cell growth that underpins tissue curvature in darkness. These findings establish an epidermis-centered framework for PIF-dependent coupling of cMT organization and anisotropic growth, positioning them as candidate central regulators providing a foundation for future cross-layer analyses to generalize these mechanical interactions across tissues.

HY5 enhances Arabidopsis tolerance to combined high light and heat stress by coordinating photoprotection and hormone signaling.

Balfagón D, Segarra-Medina C, Chávez-Jácome D … +4 more , Dos Reis de Oliveira T, Santa-Catarina C, Silveira V, Halliday KJ

Plant Physiol Biochem · 2026 Jun · PMID 42314386 · Publisher ↗

High light (HL) and heat stress (HS) are two major abiotic factors that commonly co-occur in nature and severely impair photosynthetic performance when combined. The bZIP transcription factor HY5 is a well-known integrat... High light (HL) and heat stress (HS) are two major abiotic factors that commonly co-occur in nature and severely impair photosynthetic performance when combined. The bZIP transcription factor HY5 is a well-known integrator of light and temperature cues, but its role under combined HL + HS stress remains largely unexplored. Here, we investigated the role of HY5 in Arabidopsis tolerance to HL + HS using wild-type (Col-0), HY5-deficient (hy5-215), and HY5-overexpressing (HY5OX) lines. Physiological and biochemical analyses revealed that HY5OX plants maintained higher photosynthetic efficiency, lower membrane damage, and improved leaf health under HL + HS, while hy5-215 mutants were hypersensitive. Proteomic profiling showed that HL + HS induced distinct HY5-dependent changes in the accumulation of photosynthesis-related proteins, particularly Photosystem II core subunits D1 and D2. NPQ4/PsbS, a key component of non-photochemical quenching (NPQ), was related to the presence of HY5, with impaired NPQ activation in hy5-215 correlating with lower F/F and higher increased oxidative damage. Hormonal profiling further revealed that HY5 is required for ABA and JA signaling under HL + HS. Our findings highlight HY5 as a central regulator of tolerance to combined HL + HS stress, acting through the transcriptional coordination of photoprotective proteins and hormonal signaling networks.

The microtubule-associated protein RIC1 forms biomolecular condensates to promote the polymerization of microtubule bundles in vitro.

Bai W, Chen Y, Chen Y … +13 more , Zhang S, Wu T, Wu X, Chen M, Wu J, Sun D, Dang X, Liao J, Yamamuro C, Wang Q, Pan X, Chen B, Lin D

Plant Physiol Biochem · 2026 Jun · PMID 42314385 · Publisher ↗

The Arabidopsis thaliana RIC1, a key member of the ROP (Rho GTPase of Plants)-Interactive CRIB motif-containing protein family, has been characterized as a plant-specific microtubule-associated protein. RIC1 is thought t... The Arabidopsis thaliana RIC1, a key member of the ROP (Rho GTPase of Plants)-Interactive CRIB motif-containing protein family, has been characterized as a plant-specific microtubule-associated protein. RIC1 is thought to function as a key downstream effector of the small GTPase Rho-of-Plant 6 (ROP6), orchestrating the formation of well-ordered cortical microtubule arrays and promoting the interdigitated growth pattern of cotyledon or leaf epidermal pavement cells. However, the precise physicochemical properties governing RIC1's function in microtubule organization regulation remain known. Here, we demonstrate that RIC1 is a predicted intrinsically disordered protein that forms biomolecular condensates, which promote the polymerization of microtubule bundles in vitro. In Arabidopsis cotyledon pavement cells, RIC1 forms distinct condensate-like puncta, with a specific subset associating directly with cortical microtubules. RIC1 self-assembles into distinct spherical condensates under in vitro macromolecular crowding conditions. Remarkably, these RIC1 condensates co-condense with tubulin in vitro. We further demonstrate that RIC1 condensates elevate the local tubulin concentration, correlate with enhanced microtubule nucleation, and promote the polymerization of microtubule bundles in vitro. Moreover, the growing microtubule bundles actively reshape these condensates. Thus, RIC1 condensates may provide a specialized microenvironment that enables tubulin sequestration while simultaneously regulating microtubule nucleation and polymerization in vitro. This study provides crucial insights into the biomolecular condensation properties of the microtubule-associated protein RIC1, establishing a groundwork for future investigations into its in vivo functional mechanisms.

Integrated physiological and transcriptomic analyses reveal the role of polyamines in blueberry fruit coloration.

Zhao T, Zhang X, Li Q … +4 more , Yan T, Wang Q, Lu J, Wang D

Plant Physiol Biochem · 2026 Jun · PMID 42309044 · Publisher ↗

Polyamines (PAs) play important roles in regulating fruit ripening and coloration. This study investigated the effects of exogenous polyamines including spermidine (Spd), spermine (Spm) and putrescine (Put) and the polya... Polyamines (PAs) play important roles in regulating fruit ripening and coloration. This study investigated the effects of exogenous polyamines including spermidine (Spd), spermine (Spm) and putrescine (Put) and the polyamine synthesis inhibitor, methylglyoxal-bis(guanylhydrazone) (MGBG), on blueberry fruit ripening and coloration. We treated blueberry fruits at the green-stage and subsequently measured the levels of Spd, Spm, Put, abscisic acid (ABA), ethylene (ETH), anthocyanins, and the activities of enzymes associated with PA synthesis and metabolism. The results showed that SPD treatment significantly increased anthocyanin content and PA levels in blueberry fruits, promoting fruit coloration. To understand the molecular basis of fruit ripening and coloration, we further performed transcriptome sequencing on blueberry fruits treated with Spd and MGBG and analyzed differentially expressed genes (DEGs) involved in the anthocyanin and hormone biosynthesis pathways. The transcriptome data revealed significant up-regulation of several key genes which were involved in the anthocyanin biosynthesis and hormone synthesis. Additionally, heterologous over-expression of the blueberry spermidine synthase gene (VcSPDS) in tobacco significantly increased anthocyanin accumulation and upregulated the expression of both PA biosynthetic genes (VcSPDS, VcADC, and VcSPMS) and anthocyanin biosynthetic structural genes (VcPAL, VcCHS, VcCHI, VcF3H, VcF3'5'H, VcDFR, VcANS, and VcUFGT), preliminary providing molecular evidence that supports the role of Spd as a positive regulator of anthocyanin synthesis. These results suggest that PAs promote blueberry fruit coloration by affecting the anthocyanin synthesis pathway, providing a new molecular strategy for improving fruit quality.

Streptomyces violaceoruber induces root morphological changes in Solanum lycopersicum via early epigenetic and transcriptional reprogramming.

Ricciardi V, Puccio G, Abbate L … +8 more , Faddetta T, Lo Pinto M, Caldiero C, Polito G, Palumbo Piccionello A, Gallo G, Mercati F, Cavalieri V

Plant Physiol Biochem · 2026 May · PMID 42309043 · Publisher ↗

The increasing demand for sustainable agricultural practices has highlighted the potential of Plant Growth-Promoting Bacteria (PGPB) as eco-friendly tools to enhance crop productivity while minimizing environmental impac... The increasing demand for sustainable agricultural practices has highlighted the potential of Plant Growth-Promoting Bacteria (PGPB) as eco-friendly tools to enhance crop productivity while minimizing environmental impact. Among PGPBs, members of the Actinomycetota phylum (formerly known as actinobacteria), and particularly Streptomyces violaceoruber, have emerged as promising candidates due to their ability to produce bioactive metabolites, promote plant growth, and modulate plant physiological responses. In this work, we investigated the effects of S. violaceoruber on tomato (Solanum lycopersicum) in vitro - grown seedlings using an integrated phenotypic, volatilomic, transcriptomic, and epigenetic approach. Seedlings were analyzed at seven (T1) and fifteen days (T2) post-inoculation with S. violaceoruber. Phenotypic assessment of inoculated seedlings revealed no significant alterations in shoot length or biomass, while a remarkable increase in seedling root diameter and the formation of aerial roots was observed. Transcriptomic analyses showed substantial transcriptional reprogramming, with a greater number of differentially expressed genes (DEGs) at T1, in particular involved in the regulation of biological processes, metabolic pathways, and responses to external stimuli, such as light. Co-expression network analysis of four root-associated bait genes further confirmed that these pathways are primary targets of S. violaceoruber effects. Epigenetic profiling of treated plant roots revealed an increase in global DNA methylation (5-methylcytosine) levels, along with a significant enrichment of histone post-translational modifications associated with permissive chromatin at the bait gene loci. Overall, S. violaceoruber inoculation induced notable molecular and developmental changes in tomato seedlings, reinforcing its potential as a sustainable biofertilizer. These findings provide new insights into PGPB-plant interactions and contribute to the development of environmentally-friendly strategies for crop improvement.

Regulation of light-induced anthocyanin biosynthesis in sweet cherry by the WRKY transcription factor PavWRKY18.

Chen Y, Tang W, Chu Y … +11 more , Dong Y, Liao J, Chen J, Li M, He R, Huang Y, Li H, Niu W, Du Q, Pei Y, Gong R

Plant Physiol Biochem · 2026 Jun · PMID 42302561 · Publisher ↗

Anthocyanin production and storage are crucial for determining the color and quality of sweet cherry (Prunus avium) fruits. Although WRKY transcription factors are recognized as critical regulators of secondary metabolis... Anthocyanin production and storage are crucial for determining the color and quality of sweet cherry (Prunus avium) fruits. Although WRKY transcription factors are recognized as critical regulators of secondary metabolism, their specific roles and underlying mechanisms regarding coloration in sweet cherry remain elusive. Through two comparative transcriptomic analysis of sweet cherry in different ripening stages and lighting treatments, we identified the candidate gene PavWRKY18. The expression patterns of PavWRKY18 exhibits specifically high expression during late fruit ripening and showed a strong positive correlation with light. With overexpression and sliencing of WRKY18 in sweet cherry fruits, functional characterization demonstrated that PavWRKY18 serves as a light-induced positive role of fruit coloration and accumulation of anthocyanin. Integrated transcriptomic analysis further revealed that fruit light-mediated related genes are significantly altered in the PavWRKY18-OE2 fruits. Molecular assays demonstrated that the PavWRKY18 protein directly binds to W-box cis-elements in the promoters of key structural genes involved in anthocyanin biosynthesis (PavCHS, PavF3H, PavDFR, and PavUFGT), thus activating their transcription. Our findings elucidate a molecular mechanism wherein light-induced PavWRKY18 positively regulates anthocyanin biosynthesis and accumulation through the direct targeting multiple structural genes, which might provide target for the breeding of enhancing the coloration of sweet cherry fruits.

Lipidomic analysis reveals the potentially critical roles of very-long-chain fatty acids (VLCFAs) in Antarctic moss under low temperature adaptation.

Wang H, Kong Q, Xu F … +3 more , Wang J, Liu S, Zhang P

Plant Physiol Biochem · 2026 Jun · PMID 42302560 · Publisher ↗

The Antarctic continent possesses extremely harsh environment with low temperature and short growing seasons. Antarctic mosses, as one of the dominant terrestrial vegetation, have evolved molecular strategies to survive... The Antarctic continent possesses extremely harsh environment with low temperature and short growing seasons. Antarctic mosses, as one of the dominant terrestrial vegetation, have evolved molecular strategies to survive in harsh conditions. However, the specific mechanisms underlying their adaptation to cold stress remain largely unknown. Here, we performed a comprehensive lipidomic analysis of Antarctic moss (Pohlia nutans) across a temperature gradient (4°C-23°C). The results underscored the potentially critical role of membrane lipid remodeling, fatty acid unsaturation and very-long-chain fatty acids (VLCFAs) in low temperature adaptation. Furthermore, the β-ketoacyl-CoA synthase (PnKCS) genes family, encoding the rate-limiting enzyme for VLCFA biosynthesis, comprised 19 members in P. nutans and exhibited differential expression under different temperature treatments. Heterologous expression of PnKCS10/11 promoted the resistance to low temperature in Physcomitrium patens by increasing VLCFAs accumulation. Collectively, these findings elucidate the molecular survival strategies of Antarctic moss in polar habitats, highlighting the potential involvement of VLCFAs in adaptation to extreme temperature.

Alternative transcription initiation and termination and its control through epigenetic marks in rice revealed by nanopore RNA sequencing.

Li H, Wang G, Li X … +4 more , Zhang J, Li X, Shen W, Wang G

Plant Physiol Biochem · 2026 Jun · PMID 42302559 · Publisher ↗

Gene transcription is a complex process. The alternative transcription-initiation (ATI) and termination (ATT) contribute to the complexity of transcriptomes and subsequently cause structural and functional diversity of p... Gene transcription is a complex process. The alternative transcription-initiation (ATI) and termination (ATT) contribute to the complexity of transcriptomes and subsequently cause structural and functional diversity of proteomes. Investigations on ATI and ATT were mainly explored in animal, while it was rarely investigated in plants. Here, with the assay of full-length RNA sequencing by nanopore, we identified new- and tissue specific transcripts, ATI and ATT events during rice floret developmental stages, revealing the rice transcriptome diversity. Differentially expressed ATI among the three floret developmental stages highlighted the importance of TI sites selection during developmental switches. By integrating DNA methylation data, we found that the selection of alternative transcription initiation (ATI) and termination (ATT) sites was significantly correlated with low DNA methylation levels. In addition, genes exhibiting ATI/ATT events were significantly associated with a highly open chromatin state. Taken together, nanopore sequencing offers advantages in obtaining the full-length new transcripts, which improve the annotation and accuracy of the rice genome. The integrative analysis sheds light on those epigenetic marks regulates the selection of TI and TT sites, which resulted in transcriptome complexity in rice.

Kibdelosporangium sp. and chitosan nanoparticles differentially mitigated mercury stress in two Phaseolus vulgaris L. cultivars: Improved photosynthetic efficiency, primary metabolism and redox homeostasis.

Korany SM, Mohamed MYA, Sonbol H … +5 more , Alkhateeb MA, Aloufi AS, Maridueña-Zavala MG, Hagagy N, Tan U

Plant Physiol Biochem · 2026 Jun · PMID 42302558 · Publisher ↗

Mercury contamination poses a significant risk to worldwide crop yields, negatively impacting their development and metabolic processes. Effective mitigation strategies can enhance plant resilience under such stress cond... Mercury contamination poses a significant risk to worldwide crop yields, negatively impacting their development and metabolic processes. Effective mitigation strategies can enhance plant resilience under such stress conditions. This study investigates the combined mitigation potential of Kibdelosporangium sp. and chitosan nanoparticles (CSNPs) in mitigating Hg toxicity in two Phaseolus vulgaris L. cultivars: the sensitive 'Nebraska' and the tolerant 'Giza 6'. Hg exposure reduced biomass and impaired photosynthesis, with fresh weight and photosynthetic rates declining by up to 57% and 65%, respectively, in Nebraska cultivar. However, the combined application of Kibdelosporangium sp. and CSNPs alleviated these stress effects, increasing fresh weight up to 84.7% and reducing Hg accumulation from 86.58 to 48.23 μg g DW in Nebraska. The combined application restored photosynthesis and increasing the bioavailability of primary sugars. These served as essential carbon skeletons and energy precursors for the biosynthesis of organic, amino, and fatty acids. Consequently, the combined treatment reduced HO accumulation and lipid peroxidation by approximately 38-45% in Nebraska compared with the Hg-only treatment. Antioxidant defences (e.g., CAT, DHAR, SOD, TAC) were increased by 34-111% in Nebraska and 52-160% in Giza 6. under Hg stress, and combined treatments further increase this defence response. Hg significantly increased total soluble sugars in Nebraska (85%) and Giza 6 (100%), while proline rose by 33% in Nebraska. However, combined treatment mitigated these effects, normalizing sugar levels and balancing carbon allocation, especially in Giza 6. By stabilizing antioxidant systems and metabolic balance, the intervention effectively maintained high tocopherol levels. Particularly, the sensitive Nebraska cultivar showed greater recovery, pointing out higher responsiveness. This study demonstrates an innovative approach to enhancing legume resilience through microbial and nano-mitigants that optimize photosynthesis, metabolic accumulation, and redox homeostasis in heavy-metal-polluted environments.

Humic substances for abiotic stress resilience: current knowledge and perspectives.

Salehi H, Savy D, Rao MA … +1 more , Lucini L

Plant Physiol Biochem · 2026 Jun · PMID 42302557 · Publisher ↗

Humic substances (HS), encompassing humic and fulvic fractions as well as humic-like materials, are among the most adopted biostimulants to enhance plant resilience under abiotic stress. Yet their efficacy remains highly... Humic substances (HS), encompassing humic and fulvic fractions as well as humic-like materials, are among the most adopted biostimulants to enhance plant resilience under abiotic stress. Yet their efficacy remains highly variable depending on the source, chemical composition, application dose, and experimental conditions. This review provides an overview of scientific studies addressing the application of HS to plants under different abiotic stresses. The chemical characterization of HS, including aromaticity, functional group distribution, and molecular size, affects their biological activity, though a direct and predictive structure-activity relationship has not yet been established. A convergent view across stress types indicates that HS influence plasma membrane function, ion transport, and redox balance, with the magnitude and direction of these effects depending on the chemical composition of the product, the application dose, and the stress conditions. However, the current review, considering studies evaluating the mode of action of HS during the years 2021-2025, reflects insufficient reporting of HS characterization and dose normalization and their corresponding biological effects. We also found a lack of high-throughput omics technologies to provide a deeper understanding of the mode of action of HS. Future studies should prioritize reporting clear standards for HS characterization and harmonized dosage recommendations. It is also critical to link the chemical properties of HS with their behaviors at the root-soil interface, the specific phase of stress being addressed, and the responses at various scales.

Physiological and transcriptomic insights into the alleviating effects of dopamine on drought stress in loquat (Eriobotrya japonica) seedlings.

Luo X, Wang X, Xu A … +8 more , Kong X, Luo Y, Zheng J, Li Y, Zhang H, Lin L, Deng Q, Jia Y

Plant Physiol Biochem · 2026 Jun · PMID 42302556 · Publisher ↗

Drought stress impairs metabolic processes and suppresses growth in loquat (Eriobotrya japonica (Thunb.) Lindl.) seedlings, resulting in compromised physiological function. Although previous studies have indicated that d... Drought stress impairs metabolic processes and suppresses growth in loquat (Eriobotrya japonica (Thunb.) Lindl.) seedlings, resulting in compromised physiological function. Although previous studies have indicated that dopamine can improve plant resistance to abiotic stresses, its regulatory role and molecular mechanisms underlying drought tolerance in loquat seedlings remain unclear. This study investigated the effects of exogenous dopamine on the drought resistance of loquat seedlings and its underlying mechanisms by integrating morphological, physiological, and transcriptomic analyses. The results revealed that dopamine pretreatment increased photosynthetic pigment content and stomatal aperture, thereby enhancing photosynthetic capacity. Concurrently, the activities of antioxidant enzymes and levels of antioxidants were significantly increased, which effectively scavenged excess reactive oxygen species (ROS), reduced membrane lipid peroxidation, and alleviated wilting symptoms. Transcriptomic analysis identified a total of 377 differentially expressed genes (DEGs) involved in the dopamine-mediated response to drought stress. These genes were primarily enriched in glutathione metabolism, MAPK signaling pathway, plant hormone signal transduction, and plant-pathogen interaction pathways. The improvement of photosynthetic efficiency and antioxidant capacity was mainly associated with CaM4, MKK9, PYR/PYL, and SnRK2 genes, as well as WRKY, AP2/ERF, and HD-Zip transcription factors.

ClWRKY75 regulates lignin biosynthesis to confer Colletotrichum fructicola resistance in watermelon.

Gong S, Wang D, Yang Z … +10 more , Liu G, He C, Sun L, Chen Z, Li S, Klein J, Sun X, Dai S, Bu F, Tang B

Plant Physiol Biochem · 2026 Jun · PMID 42296584 · Publisher ↗

Watermelon, a significant horticultural crop, is threatened by various fungal diseases throughout its growth cycle, with Colletotrichum fructicola, a member of the Colletotrichum spp., being notorious for causing anthrac... Watermelon, a significant horticultural crop, is threatened by various fungal diseases throughout its growth cycle, with Colletotrichum fructicola, a member of the Colletotrichum spp., being notorious for causing anthracnose, which damages both seedlings and fruits. Anthracnose control mainly depends on fungicides, while resistant watermelon breeding is hindered by scarce resistance genes and unclear molecular mechanisms. Here we show that the WRKY transcription factor ClWRKY75 was strongly induced in the resistant line M10 following Colletotrichum fructicola infection by comparative transcriptomic analysis. Silencing ClWRKY75 through virus-induced gene silencing (VIGS) converted resistant M10 plants into a susceptible phenotype, accompanied by reduced lignin accumulation. We further demonstrate that ClWRKY75 directly binds to and activates the promoter of ClPAL20, a key gene in the lignin biosynthetic pathway. These results reveal that ClWRKY75 enhances anthracnose resistance by promoting lignin synthesis, and offer useful gene resources for resistant watermelon breeding.

Pumpkin rootstock-driven variation in mineral nutrient allocation determines differential melon fruit quality.

Kaleem MM, Chen J, Song Y … +8 more , Nawaz MA, Ateeq M, Mu X, Ashraf MA, Ahamd M, Huang Y, Cheng J, Bie Z

Plant Physiol Biochem · 2026 Jun · PMID 42296583 · Publisher ↗

Rootstocks are commonly used in melon cultivation to mitigate biotic and abiotic stresses and improve fruit quality. However, the extent to which rootstocks influence nutrient allocation, growth dynamics, and fruit quali... Rootstocks are commonly used in melon cultivation to mitigate biotic and abiotic stresses and improve fruit quality. However, the extent to which rootstocks influence nutrient allocation, growth dynamics, and fruit quality formation in the scion remains insufficiently understood. This study compared the effects of two pumpkin rootstocks (Tianzhen No. 1 and Sizhuang No. 12) on melon fruit quality by examining nutrient partitioning, sugar accumulation, hormonal balance, and expression of sugar metabolism-related genes. Compared with non-grafted plants, rootstock grafting enhanced plant growth and increased total N, P, and K uptake (mg plant) as well as tissue nutrient concentrations (mg g DW) during vine growth and fruit maturation stages. Tianzhen No. 1 showed higher K distribution to the scion (97.6%) and greater fruit K accumulation (45.88 mg/g) than Sizhuang No. 12. Fruits grafted onto Tianzhen No. 1 also exhibited higher total soluble solids (15.96 °Brix) and soluble sugar contents, including glucose (18.28 mg g), fructose (20.20 mg g), and sucrose (144.25 mg g), whereas fruits grafted onto Sizhuang No. 12 showed higher firmness (4.18 kg cm) compared with Tianzhen No. 1-grafted fruits (3.07 kg cm). At fruit maturation, Tianzhen No. 1-grafted melons had higher ABA content (14.49 ng/g) and upregulated expression of sugar transporter genes (HT2, HT7, SWEET4, SWEET7) and genes encoding key metabolic enzymes (SPS1, NIN3) compared to Sizhuang No. 12. These observed associations suggest that rootstock-mediated differences in nutrient uptake may contribute to altered scion fruit quality metabolism. Overall, the findings provide insights into the physiological and molecular basis of rootstock-dependent variation in melon fruit quality and may contribute to the development of targeted rootstock selection strategies in melon breeding programs.

Sunflower NAC84 confers plant salt and drought tolerance and yield-increasing potential.

Jiang R, Liu Y, Li W … +3 more , Huang Q, Lei Z, Zeng Y

Plant Physiol Biochem · 2026 Jun · PMID 42296582 · Publisher ↗

Drought and salinity pose serious threats to global food security. However, enhancing crop stress tolerance through the overexpression of an individual gene often incurs trade-offs with other agronomic traits. Therefore,... Drought and salinity pose serious threats to global food security. However, enhancing crop stress tolerance through the overexpression of an individual gene often incurs trade-offs with other agronomic traits. Therefore, identifying alleles that simultaneously enhance stress tolerance and yield is a critical priority. Sunflower (Helianthus annuus L.) is often chosen for saline-alkali soil reclamation because of its significant salt and drought tolerance. As transcriptional controllers found only in plants, NAC proteins are central to orchestrating morphogenesis and mediating tolerance to various stresses. Here, we report that HaNAC84 confers salt and drought stress tolerance and exhibits yield-promoting potential. Stable over-expression of HaNAC84 in Arabidopsis and transient expression in sunflower both markedly increased tolerance to salt and drought. Mechanistically, HaNAC84 enhanced the activities of key antioxidant enzymes (SOD, POD, CAT, GR) and enhanced the overall antioxidant capacity, thereby limiting reactive oxygen accumulation and protecting membrane integrity. In Arabidopsis, HaNAC84 up-regulated POD, CAT and P5CS gene transcription, activated ABA biosynthesis and signaling to accelerate stomatal closure, and induced the expression of downstream defense genes (NHX1, DREB2A, LEA3, RD29A). Besides stress protection, HaNAC84 promoted root elongation and increased bolting branches, enhanced photosynthetic efficiency, and ultimately increased seed yield under control and stress conditions. Collectively, our findings identify HaNAC84 as a promising target for engineering stress-resilient crops that maintain productivity under abiotic stress.

High-temperature stress alters grain quality, starch granule structure, and gene expression in basmati rice cultivars.

Mishra A, Chaudhary R, Fatima P … +8 more , Sharma V, Srivastava D, Yadav P, Shamim M, Kumar P, Siddiqui MW, Siddiqui MH, Irfan M

Plant Physiol Biochem · 2026 May · PMID 42296581 · Publisher ↗

High-temperature stress is a serious issue for rice production, yet the molecular and biochemical foundation of heat-induced deterioration of grain quality is remains poorly understood, especially in the case of aromatic... High-temperature stress is a serious issue for rice production, yet the molecular and biochemical foundation of heat-induced deterioration of grain quality is remains poorly understood, especially in the case of aromatic rice. In the current study, 15 rice cultivars including Basmati rice, Non-Basmati aromatic rice and non-aromatic heat-tolerant cultivar Nagina 22 as a check were evaluated for high temperature stress. Physiological, biochemical, and molecular characteristics related to grain quality characteristics including starch parameters, amino acid composition, starch granule structure, and gene expression of starch synthesis related genes viz. GBSSI, Wx and Pro were studied. High-temperature stress during flowering and grain filling stage significantly lowered photosynthetic rate, stomatal conductance, transpiration rate, amylose content and essential amino acids such as lysine, methionine, and glutamine. Scanning electron microscopy indicated abnormal and deformed starch granules in Basmati genotypes, while Nagina 22 was affected the least. Gene expression analysis indicated down-regulation of GBSSI Wx and Pro 7 genes leads to inhibition of starch synthesis and accelerating grain deterioration. Collectively, these results prove that rice that is inherently aromatic is especially sensitive to heat-induced deterioration of grain quality and reveal new knowledge of the physiological, biochemical, and molecular basis of heat-induced sensitivity of rice. The results also indicate possible molecular targets for developing thermotolerant Basmati and Non-Basmati aromatic rice.

Apoplastic retention over vacuolar sequestration: a potential more efficient strategy limiting root-to-shoot cadmium translocation in leafy vegetable crops.

Huang Y, Fu H, Huang B … +2 more , Xin J, Shen C

Plant Physiol Biochem · 2026 Jun · PMID 42296580 · Publisher ↗

Cadmium (Cd) accumulation in plants is coordinated by uptake, long-distance transport, and intracellular detoxification, yet the relative contributions of these processes to Cd allocation remain unclear. Using leafy vege... Cadmium (Cd) accumulation in plants is coordinated by uptake, long-distance transport, and intracellular detoxification, yet the relative contributions of these processes to Cd allocation remain unclear. Using leafy vegetables as a representative model for rapid heavy metal translocation, we conducted a global meta-analysis of 86 studies to identify the major physiological control points associated with Cd allocation. Our results showed that restricted root-to-shoot translocation represented the dominant regulatory bottleneck, with an effect size (-0.37) more than double that of root Cd reduction (-0.17). Mechanistically, this transport restriction was strongly associated with enhanced apoplastic retention within root cell walls. Increased pectin content and PME-mediated demethylation may increase Cd binding in the cell wall, thereby potentially limiting Cd availability for symplastic loading and xylem transport. In contrast, high-Cd accumulation triggers the synthesis of high-molecular-weight phytochelatins (e.g., PC4), consistent with the activation of intracellular detoxification pathways. Together, these findings suggest that cell wall immobilization may contribute more strongly than downstream vacuolar sequestration to limiting Cd translocation in leafy vegetables. This study provides a fundamental framework for understanding physiological trade-offs under Cd stress and identifies cell wall remodeling as a primary target for developing low-Cd leafy vegetable culticvars.

Biochemical trade-offs in basil: Photovoltaic shading and alcoholic foliar sprays in a climate-smart greenhouse.

Batebi S, Moradi H, Motevali A

Plant Physiol Biochem · 2026 Jun · PMID 42296579 · Publisher ↗

This study evaluated the effects of photovoltaic (PV)-induced shading and alcoholic foliar application on photosynthetic pigments, biochemical characteristics, oxidative stress, and essential oil composition of Italian b... This study evaluated the effects of photovoltaic (PV)-induced shading and alcoholic foliar application on photosynthetic pigments, biochemical characteristics, oxidative stress, and essential oil composition of Italian basil (Ocimum basilicum L.) grown under greenhouse conditions during two cultivation seasons. Three PV roof coverage levels, 0% (C0), 50% (C50), and 100% (C100), were combined with ethanol-methanol foliar spray treatments at different ratios. Moderate shading (C50) sustained most biochemical traits without significant decline while directing favorable shifts in essential oil composition, notably enhancing the accumulation of eucalyptol and linalool. In contrast, full roof coverage (C100) significantly increased chlorophyll a, chlorophyll b, and total chlorophyll contents by 29%, 27.5%, and 34%, respectively, indicating physiological acclimation to reduced irradiance, but reduced total phenolic content, flavonoids, and antioxidant activity by 51%, 25%, and 28%, respectively. Plants grown under full-light conditions exhibited significantly greater malondialdehyde accumulation, with MDA levels 19.7% and 28.3% higher than shaded treatments across the two cultivation seasons, reflecting increased oxidative stress. Alcoholic foliar application reduced lipid peroxidation and improved biochemical performance, with the 30% ethanol +30% methanol treatment producing the strongest response. Alcohol treatments also modified the volatile composition through increased neophytadiene abundance. The results indicate that moderate PV-induced shading, particularly at 50% roof coverage, combined with optimized alcoholic foliar application, can improve physiological performance and favorably modulate phytochemical composition in greenhouse-grown basil.
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