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Crops around the world are facing a diversity of environmental problems, of which high temperatures are proving to be the most serious threat to crops. Polyamine putrescine (Put) acts as a master growth regulator that contributes to optimal plant growth and development and increased stress tolerance. Here, the current study aimed to elucidate how Put functions in regulating chlorophyll (Chl) metabolism, oxidative stress, and antioxidant defense, as well as to characterize the expression of genes related to heat stress in tomato seedlings under such stress. The results revealed that Put treatment significantly attenuates heat-induced damage by promoting biomass production, increasing photosynthetic efficiency, and inhibiting excessive production of oxidative stress markers. Heat stress markedly decreased the Chl content in the tomato leaf and accelerated the leaf yellowing process. However, Put-treated tomato seedlings showed a higher Chl content, which could be associated with the functions of Put in elevating PBGD activity (Chl biosynthesis enzyme) and suppressing the activity of the Chl catabolic enzyme (Chlase and MDCase). Under high-temperature stress, the expression levels of the gene encoding factors involved in Chl biosynthesis and Chl catabolism were significantly down- and upregulated, respectively, and this trend was reversed in Put-treated heat-stressed seedlings. In addition, exogenous application of Put boosted the activity of antioxidant enzymes, along with the levels of expression of their encoding genes, only in plants that were heat stressed. Furthermore, the expression levels of heat-shock-related genes (HSP90, HSP70, and HsfA1) were elevated in Put-treated, high-temperature-stressed tomato seedlings. Taken together, our results indicate that Put treatment significantly increases the heat tolerance of tomato seedlings, by elevating Chl concentrations and suppressing Chl catabolic enzyme activity, modulating endogenous free PA content, increasing antioxidant defense efficiency, and upregulating the expression of heat-shock-related genes. [ABSTRACT FROM AUTHOR]

Photosynthesis is a fundamental biosynthetic process in plants that can enhance carbon absorption and increase crop productivity. Heat stress severely inhibits photosynthetic efficiency. Melatonin is a bio-stimulator capable of regulating diverse abiotic stress tolerances. However, the underlying mechanisms of melatonin-mediated photosynthesis in plants exposed to heat stress largely remain elucidated. Our results revealed that melatonin treatment (100 μM) in tomato seedlings increased the endogenous melatonin levels and photosynthetic pigment content along with upregulated of their biosynthesis gene expression under high-temperature stress (42 °C for 24 h), whereas heat stress significantly decreased the values of gas exchange parameters. Under heat stress, melatonin boosted CO 2 assimilation, i.e., V c,max (maximum rate of ribulose-1,5-bisphosphate carboxylase, Rubisco), and J max (electron transport of Rubisco generation) and also enhanced the Rubisco and FBPase activities, which resulted in upregulated photosynthetic related gene expression. In addition, heat stress greatly reduced the photochemical chemistry of photosystem II (PSII) and photosystem I (PSI), particularly the maximum quantum efficiency of PSII (Fv/Fm) and PSI (Pm). Conversely, melatonin supplementation increased the chlorophyll a fluorescence parameters led to amplifying the electron transport efficiency. Moreover, heat stress decreased the actual PSII efficiency (ΦPSII), electron transport rate (ETR) and photochemical quenching coefficient (qP), while increasing nonphotochemical quenching (NPQ); however, melatonin reversed these values, which helps to fostering the dissipation of excess excitation energy. Taken together, our results provide a concrete insight into the efficacy of melatonin-mediated photosynthesis performance in a high-temperature regime. • Melatonin increased photosynthetic pigment contents and CO 2 assimilation of tomato seedlings exposed to heat stress. • Melatonin protected the photosystem II and photosystem I reaction center and reduced photoinhibition under thermal stress. • Melatonin enhanced photosynthesis efficiency by elevating the Rubisco and FBPase enzyme activities. • Melatonin treatment efficiently enhanced the heat-induced reduction of chlorophyll a fluorescence photochemistry. [ABSTRACT FROM AUTHOR]

Heat stress and abscisic acid (ABA) induce leaf senescence, whereas melatonin (MT) and gibberellins (GA) play critical roles in inhibiting leaf senescence. Recent research findings confirm that plant tolerance to diverse stresses is closely associated with foliage lifespan. However, the molecular mechanism underlying the signaling interaction of MT with GA and ABA regarding heat-induced leaf senescence largely remains undetermined. Herein, we investigated putative functions of melatonin in suppressing heat-induced leaf senescence in tomato and how ABA and GA coordinate with each other in the presence of MT. Tomato seedlings were pretreated with 100 μM MT or water and exposed to high temperature (38/28°C) for 5 days (d). Heat stress significantly accelerated senescence, damage to the photosystem and upregulation of reactive oxygen species (ROS), generating RBOH gene expression. Melatonin treatment markedly attenuated heat-induced leaf senescence, as reflected by reduced leaf yellowing, an increased Fv/Fm ratio, and reduced ROS production. The Rbohs gene, chlorophyll catabolic genes, and senescence-associated gene expression levels were significantly suppressed by MT addition. Exogenous application of MT elevated the endogenous MT and GA contents but reduced the ABA content in high-temperature-exposed plants. However, the GA and ABA contents were inhibited by paclobutrazol (PCB, a GA biosynthesis inhibitor) and sodium tungstate (ST, an ABA biosynthesis inhibitor) treatment. MT-induced heat tolerance was compromised in both inhibitor-treated plants. The transcript abundance of ABA biosynthesis and signaling genes was repressed; however, the biosynthesis genes MT and GA were upregulated in MT-treated plants. Moreover, GA signaling suppressor and catabolic gene expression was inhibited, while ABA catabolic gene expression was upregulated by MT application. Taken together, MT-mediated suppression of heat-induced leaf senescence has collaborated with the activation of MT and GA biosynthesis and inhibition of ABA biosynthesis pathways in tomato. [ABSTRACT FROM AUTHOR]

Tomato is an important vegetable that is highly sensitive to drought (DR) stress which impairs the development of tomato seedlings. Recently, melatonin (ME) has emerged as a nontoxic, regulatory biomolecule that regulates plant growth and enhances the DR tolerance mechanism in plants. The present study was conducted to examine the defensive role of ME in photosynthesis, root architecture, and the antioxidant enzymes' activities of tomato seedlings subjected to DR stress. Our results indicated that DR stress strongly suppressed growth and biomass production, inhibited photosynthesis, negatively affected root morphology, and reduced photosynthetic pigments in tomato seedlings. Per contra, soluble sugars, proline, and ROS (reactive oxygen species) were suggested to be improved in seedlings under DR stress. Conversely, ME (100 µM) pretreatment improved the detrimental-effect of DR by restoring chlorophyll content, root architecture, gas exchange parameters and plant growth attributes compared with DR-group only. Moreover, ME supplementation also mitigated the antioxidant enzymes [APX (ascorbate peroxidase), CAT (catalase), DHAR (dehydroascorbate reductase), GST (glutathione S-transferase), GR (glutathione reductase), MDHAR (monodehydroascorbate reductase), POD (peroxidase), and SOD (superoxide dismutase)], non-enzymatic antioxidant [AsA (ascorbate), DHA (dehydroascorbic acid), GSH (glutathione), and GSSG, (oxidized glutathione)] activities, reduced oxidative damage [EL (electrolyte leakage), H2O2 (hydrogen peroxide), MDA (malondialdehyde), and O2•− (superoxide ion)] and osmoregulation (soluble sugars and proline) of tomato seedlings, by regulating gene expression for SOD, CAT, APX, GR, POD, GST, DHAR, and MDHAR. These findings determine that ME pretreatment could efficiently improve the seedlings growth, root characteristics, leaf photosynthesis and antioxidant machinery under DR stress and thereby increasing the seedlings' adaptability to DR stress. [ABSTRACT FROM AUTHOR]

The Bcl-2-associated athanogene (BAG) family is a group of evolutionarily conserved cochaperones involved in diverse cellular functions. Here, ten putative SlBAG genes were identified in tomato. SlBAG2 and SlBAG5b have the same gene structure and conserved domains, along with highly similar identity to their homologs in Arabidopsis thaliana, Oryza sativa, and Triticum aestivum. The qPCR data showed that BAG2 and BAG5b were highly expressed in stems and flowers. Moreover, both genes were differentially expressed under diverse abiotic stimuli, including cold stress, heat stress, salt treatment, and UV irradiation, and treatments with phytohormones, namely, ABA, SA, MeJA, and ETH. Subcellular localization showed that SlBAG2 and SlBAG5b were located in the cell membrane and nucleus. To elucidate the functions in leaf senescence of BAG2 and BAG5b, the full-length CDSs of BAG2 and BAG5b were cloned, and transgenic tomatoes were developed. Compared with WT plants, those overexpressing BAG2 and BAG5b had significantly increased chlorophyll contents, chlorophyll fluorescence parameters and photosynthetic rates but obviously decreased ROS levels, chlorophyll degradation and leaf senescence related gene expression under dark stress. Conclusively, overexpression SlBAG2 and SlBAG5b could improve the tolerance of tomato leaves to dark stress and delay leaf senescence. [ABSTRACT FROM AUTHOR]

Arable land contamination with nickel (Ni) has become a major threat to worldwide crop production. Recently, melatonin has appeared as a promising stress-relief substance that can alleviate heavy metal-induced phytotoxicity in plants. However, the plausible underlying mechanism of melatonin function under Ni stress has not been fully substantiated in plants. Herein, we conducted an experiment that unveiled critical mechanisms in favor of melatonin-mediated Ni-stress tolerance in tomato. Ni stress markedly inhibited growth and biomass by impairing the photosynthesis, photosystem function, mineral homeostasis, root activity, and osmotic balance. In contrast, melatonin application notably reinforced the plant growth traits, increased photosynthesis efficiency in terms of chlorophyll content, upregulation of chlorophyll synthesis genes, i.e. POR , CAO , CHL G , gas exchange parameters, and PSII maximum efficiency (Fv/Fm), decreased Ni accumulation and increased mineral nutrient homeostasis. Moreover, melatonin efficiently restricted the hydrogen peroxide (H 2 O 2) and superoxide radical production and increased RBOH expression and restored cellular integrity (less malondialdehyde and electrolyte leakage) through triggering the antioxidant enzyme activities and modulating AsA-GSH pools. Notably, oxidative stress was effectively mitigated by upregulation of several defense genes (SOD , CAT , APX , GR , GST , MDHAR , DHAR) and melatonin biosynthesis-related genes (TDC , T5S , SNAT , ASMT). Besides, melatonin treatment enhanced secondary metabolites (phenols, flavonoids, and anthocyanin) contents along with their encoding genes (PAL , CHS) expression, and these metabolites potentially restricted excess H 2 O 2 accumulation. In conclusion, our findings deciphered the potential functions of melatonin in alleviating Ni-induced phytotoxicity in tomato through boosting the biomass production, photosynthesis, nutrient uptake, redox balance, and secondary metabolism. • Melatonin treatment inhibits Ni uptake both in roots and shoots and enhances plant growth and biomass production. • Under Ni-stress, melatonin increases photosynthetic efficiency, nutrients uptake and root activity and reduces osmotic damage. • Melatonin supplementation modulates antioxidant defense mechanism and redox homeostasis under Ni stress. • Melatonin triggers higher accumulation of secondary metabolites and upregualtion of their biosynthesis genes (PAL , CHS) expression and restrict ROS production. [ABSTRACT FROM AUTHOR]

Background: Melatonin is a pleiotropic signaling molecule that plays multifarious roles in plants stress tolerance. The polyamine (PAs) metabolic pathway has been suggested to eliminate the effects of environmental stresses. However, the underlying mechanism of how melatonin and PAs function together under heat stress largely remains unknown. In this study, we investigated the potential role of melatonin in regulating PAs and nitric oxide (NO) biosynthesis, and counterbalancing oxidative damage induced by heat stress in tomato seedlings. Results: Heat stress enhanced the overproduction of reactive oxygen species (ROS) and damaged inherent defense system, thus reduced plant growth. However, pretreatment with 100 μM melatonin (7 days) followed by exposure to heat stress (24 h) effectively reduced the oxidative stress by controlling the overaccumulation of superoxide (O2•−) and hydrogen peroxide (H2O2), lowering the lipid peroxidation content (as inferred based on malondialdehyde content) and less membrane injury index (MII). This was associated with increased the enzymatic and non-enzymatic antioxidants activities by regulating their related gene expression and modulating the ascorbate–glutathione cycle. The presence of melatonin induced respiratory burst oxidase (RBOH), heat shock transcription factors A2 (HsfA2), heat shock protein 90 (HSP90), and delta 1-pyrroline-5-carboxylate synthetase (P5CS) gene expression, which helped detoxify excess ROS via the hydrogen peroxide-mediated signaling pathway. In addition, heat stress boosted the endogenous levels of putrescine, spermidine and spermine, and increased the PAs contents, indicating higher metabolic gene expression. Moreover, melatonin-pretreated seedlings had further increased PAs levels and upregulated transcript abundance, which coincided with suppression of catabolic-related genes expression. Under heat stress, exogenous melatonin increased endogenous NO content along with nitrate reductase- and NO synthase-related activities, and expression of their related genes were also elevated. Conclusions: Melatonin pretreatment positively increased the heat tolerance of tomato seedlings by improving their antioxidant defense mechanism, inducing ascorbate–glutathione cycle, and reprogramming the PAs metabolic and NO biosynthesis pathways. These attributes facilitated the scavenging of excess ROS and increased stability of the cellular membrane, which mitigated heat-induced oxidative stress. [ABSTRACT FROM AUTHOR]

Salt stress negatively affects plant growth, development, and crop productivity causing serious economic loss to agricultural production. Here, we investigated the exogenous application of spermidine (Spd) on tomato seedlings grown under salt stress. Salt stress reduced plant growth, biomass accumulation and chlorophyll contents, thus negatively affecting photosynthesis. Alternatively, Spd application effectively reduced the salinity-induced adverse effects in tomato seedlings by activating the H 2 O 2 mediated signaling involving the enhanced expression of RBOH1 and salt stress-responsive genes SlMYB102, SlHKT1, SlWRKY1 and SlDREB2, and improving detoxification through higher antioxidative activity and osmolyte (proline) accumulation under salt stress. It was further confirmed by significantly lower amount of H 2 O 2 , malondialdehyde and electrolyte leakage, and better ion homeostasis (Na+/K+ ratio) and photosynthetic performance of Spd-treated seedlings under salt stress. Furthermore, Spd application modulated endogenous polyamines and enhanced the biosynthesis of endogenous Spd and spermine from putrescine. Altogether, these results confirm the important role of Spd against salt stress and suggest that the increased endogenous Spd content in plants could regulate a number of stress-responsive mechanisms to protect tomato seedlings against salt stress. These results provide a good direction for further elucidation of the detailed interplay between polyamine metabolism and H 2 O 2 -mediated signaling, which would help to improve abiotic stress tolerance in plants. • Spermidine (Spd) application effectively reduced salinity-induced adverse effects in tomato. • Spd application upregulated the SlMYB102, SlHKT1, SlWRKY1 and SlDREB2 and enhanced ROS scavenging mechanisms. • Spd-treated seedlings maintained better ion homeostasis (Na+/K+ ratio) under salt stress. • Spd application modulated the endogenous polyamine content and improved the biosynthesis of endogenous Spd from putrescine. [ABSTRACT FROM AUTHOR]