Exogenous Spermidine Research Articles

Exogenous spermidine enhances salt-stressed rice photosynthetic performance by stabilizing structure and function of chloroplast and thylakoid membranes

Spermidine (Spd) is a ubiquitous low-molecular-mass aliphatic amine that acts in abiotic stress tolerance in plants. We investigated how the treatment with exogenous Spd contributed to the protection against salt stress in rice chloroplasts. Analysis of the chlorophyll (Chl) fluorescence showed that there were many negative effects of salinity on different sites of the photosynthetic machinery; however, these were alleviated by adding Spd. Spd prevented the damage of structure and function of chloroplasts under salt stress; thylakoid membrane protein components and photosynthetic pigments were not severely affected by salinity in Spd-treated plants. Spd enhanced the activities of antioxidant enzymes and decreased contents of reactive oxygen species and malondialdehyde accumulation, suggesting that Spd may participate in the redox homeostasis in chloroplasts under salt stress. These results highlighted the positive effects of Spd on rice chloroplasts under salt stress through maintaining chloroplast structure stability.

Effects of Exogenous Spermidine on Seed Germination and As Uptake and Accumulation of Rice Under As5+ Stress

As pollution in farmland has a toxic effect on the growth of crops, which reduces their yield and quality. The effects of exogenous spermidine (Spd) on rice seed germination and seedling growth under As5+ stress were studied. The results showed that exogenous Spd could promote the germination of rice seeds under As5+ stress, improve the germination potential and germination rate of seeds, and promote the growth of seedling roots. The addition of Spd could increase the activity of catalase (CAT), peroxidase (POD), and superoxide dismutase (SOD) of rice seedlings and roots under As5+ stress, and reduce the content of malondialdehyde (MDA) in rice buds and roots. When As5+ concentration was 25 μmol·L-1, adding 500 μmol·L-1 and 1000 μmol·L-1 Spd, MDA content in rice roots was decreased by 12.3% and 31.3% and CAT activity of rice shoots was increased by 105.1% and 101.4%, and CAT activity of rice roots was increased by 29.9% and 57.1%, respectively. The addition of Spd also affected the uptake and accumulation of As in rice. When the concentration of As5+ was 25 μmol·L-1, adding 500 μmol·L-1 and 1000 μmol·L-1 Spd, the concentration of As in rice shoots decreased by 69.4% and 75.1%, and As concentration in rice roots decreased by 7.6% and 24.4%, respectively. Spd could therefore effectively alleviate the toxic effect of As5+ on rice.

Exogenous Spermidine Promotes γ-Aminobutyric Acid Accumulation and Alleviates the Negative Effect of NaCl Stress in Germinating Soybean (Glycine max L.).

We investigated the effects of exogenous spermidine (Spd) on the physiological status, γ-aminobutyric acid (GABA) synthase activity, and gene expressions in germinating soybeans under NaCl stress. The results show that Spd significantly increases sprout growth and biomass, decreases malonaldehyde and H2O2 contents, and markedly promotes the activities of superoxide dismutase, catalase, peroxidase, and glutathione peroxidase of germinating soybeans. The harmful effect of NaCl stress was alleviated by exogenous Spd. GABA accumulation in germinating soybeans was caused by the activation of diamine oxidase, polyamine oxidase, aminoaldehyde dehydrogenase, and glutamate decarboxylase activities and by up-regulating their gene expression under Spd-NaCl treatment. The GABA content decreased by 57% and 46% in germinating soybeans with the application of aminoguanidine under Spd and Spd-NaCl treatments, respectively. In conclusion, spermidine induces the accumulation of GABA and increases sprouts biomass, thereby enhancing the functional quality of germinating soybeans.

Spermidine application affects the adventitious root formation and root morphology of apple rootstock by altering the hormonal profile and regulating the gene expression pattern

Adventitious root (AR) formation is a critical process for the clonal propagation of horticultural crops. Polyamines are involved in several processes associated with root growth such as cell division and cell development. In this study, Malus prunifolia var. ringo stem cuttings were treated with exogenous spermidine (Spd) 20 μmol/L in order to investigate the physiological and molecular mechanism of Spd to promote ARs formation. The results of the present study indicated an involvement of Spd in AR formation in apple rootstock by interacting with IAA. Morphological and anatomical observation revealed that the Spd application promotes AR formation. According to our results exogenous Spd application promoted AR formation by improving the concentration of endogenous IAA and GA3 at 1, 3 and 1, 8 days respectively as well as by decreasing the ABA and JA contents at all time points except JA at 16 day in response to Spd treatment. Furthermore no significant difference was observed in BR and ZR levels at all time points except at 8 day in both groups. Spd application obviously up-regulated the expression of Spd related genes (MdSAMDC1, MdPAO and MdSPDS6) and auxin biosynthesis related genes (MdIAA7, MdIAA14 and MdIAA23) in the basal parts of apple rootstock. On the other hand MdWOX11 up-regulated the expression of MdLBD16 and MdLBD29, which induced the expression of cell cycle related genes MdCYCD1;1 and MdCYCP4;1. Collectively, our work laid the foundation for applied research and could be useful in future crop improvement programs.

Polyamines produced by Sinorhizobium meliloti Rm8530 contribute to symbiotically relevant phenotypes ex planta and to nodulation efficiency on alfalfa.

In nitrogen-fixing rhizobia, emerging evidence shows significant roles for polyamines in growth and abiotic stress resistance. In this work we show that a polyamine-deficient ornithine decarboxylase null mutant (odc2) derived from Sinorhizobium meliloti Rm8530 had significant phenotypic differences from the wild-type, including greatly reduced production of exopolysaccharides (EPS; ostensibly both succinoglycan and galactoglucan), increased sensitivity to oxidative stress and decreased swimming motility. The introduction of the odc2 gene borne on a plasmid into the odc2 mutant restored wild-type phenotypes for EPS production, growth under oxidative stress and swimming. The production of calcofluor-binding EPS (succinoglycan) by the odc2 mutant was also completely or mostly restored in the presence of exogenous spermidine (Spd), norspermidine (NSpd) or spermine (Spm). The odc2 mutant formed about 25 % more biofilm than the wild-type, and its ability to form biofilm was significantly inhibited by exogenous Spd, NSpd or Spm. The odc2 mutant formed a less efficient symbiosis with alfalfa, resulting in plants with significantly less biomass and height, more nodules but less nodule biomass, and 25 % less nitrogen-fixing activity. Exogenously supplied Put was not able to revert these phenotypes and caused a similar increase in plant height and dry weight in uninoculated plants and in those inoculated with the wild-type or odc2 mutant. We discuss ways in which polyamines might affect the phenotypes of the odc2 mutant.

Exogenous Spermidine Promotes Somatic Embryogenesis of Cunninghamia lanceolata by Altering the Endogenous Phytohormone Content

In order to study how exogenous hormones in C. lanceolata (gymnosperm) regulate somatic embryogenesis, we measured the endogenous phytohormones of two genotypes with different somatic embryogenesis efficiency and found that an increase in endogenous concentrations of IAA and ABA may be correlated to more efficient somatic embryogenesis. By applying exogenous spermidine, we found that exogenous hormones may affect somatic embryogenesis efficiency through affecting the endogenous phytohormone content. Based on these results, further studies can be conducted whereby the concentration of exogenous hormones or the levels of endogenous phytohormones by molecular methods are regulated to promote somatic embryogenesis. Our research may benefit the long-term economic output of the forestry industry and lays the foundation to studying the molecular mechanism that controls somatic embryogenesis efficiency.

Prooxidant-antioxidant balance in the methylotrophic yeast Ogataea polymorpha exoposed to spermidine

Background. Many molecular compounds used by eukaryotic cells as antioxidant agents can directly interact with reactive oxygen/nitrogen species. Among them are polyamines – natural substances with pronounced antioxidant properties. The main polyamines are putrescine, spermidine, spermine. Spermidine is a positively charged polyamine that is synthesized from putrescine and serves a precursor of spermine. In particular, spermidine is distinguished by its anti-aging properties. When administered to several model organisms, it prolonged their lifespan and provided resistance to stress. Spermidine also limits overproduction of reactive oxigene species and reduces age-related oxidative protein damage, which accounts for its antioxidant activity. The aim of our study was to investigate the effects of exogenous spermidine in different concentrations on the prooxidant-antioxidant balance in the methylotrophic yeast Ogataea polymorpha naturally adapted to stressful conditions such as growth on toxic methanol-containing medium and cultivation at high temperatures (42–45 °C). Materials and Methods. In this work, O. polymorpha wild-type strain was used. Yeast cells were cultivated in the mineral medium with 0.1 % glucose for creating stressful conditions. Spermidine was added in 1 and 2 mM concentrations immediately before the start of yeast cultivation. Reactive oxygen species, the level of oxidative modifications of proteins and lipids, the concentration of reduced gluthatione, superoxide dismutase and catalase activities were assayed to examine the prooxidant-antioxidant state of the yeast cells. Results. In this study, we analyzed the effects of exogenous spermidine in different concentrations on the enzymatic (superoxide dismutase and catalase activity) and non-enzymatic (reduced glutathione) antioxidant defense systems and markers of oxidative injury (products of proteins and lipids oxidation) in the methylotrophic yeast O. polymorpha starved for glucose. It was revealed that 1 mM spermidine had a protective effect on O. polymorpha cells and decreased the content of products of the oxidative modification of proteins. At the same time, the superoxide dismutase and catalase activities and content of the reduced glutathione remained almost unchanged in the cells cultivated in the glucose-depleted medium with 1 mM spermidine compared to the medium without spermidine. Adverse effects of 2 mM spermidine (increased levels of carbonyl groups of proteins, lipid peroxidation products, disregulated superoxide dismutase and catalase activities, reduced glutathione levels, growth inhibition and cells vacuolization) were observed in the exponential growth phase of the yeast culture. During a long-term cultivation, these effects diminished, and the corresponding values approximated those of the cells grown in the control medium with the low concentration of glucose without spermidine. Conclusions. The data suggest a concentration-dependent effect of spermidine on the physiology of O. polymorpha that can be used in further studies on compounds able to mitigate negative effects of the oxidative stress in this yeast and other model organisms . At 1 mM concentration, spermidine had an apparent protective effect, whereas at the elevated 2 mM concentration this polyamine exacerbated stress load in this yeast. Keywords: methylotrophic yeast, Ogataea polymorpha , spermidine, oxidative stress

Spermidine improves antioxidant activity and energy metabolism in mung bean sprouts

The effects of exogenous spermidine and dicyclohexylamine (DCHA, spermidine synthesis inhibitor) on the antioxidative system and energy status of germinating mung bean were investigated. Results showed that exogenous spermidine increased the content of total phenolic and ascorbic acid and the antioxidative activity, but reduced activities and gene expressions of peroxidase (POD) and catalase (CAT). These changes might be explained by increased H2O2 content and activities of succinic dehydrogenase (SDH), adenosine triphosphatases (ATPases), and cytochrome c oxidase (CCO), resulting in higher adenosine triphosphate (ATP) and energy charge (EC). Interestingly, spermidine down-regulated expressions of SDH, H+-ATPase, Ca2+-ATPase and CCO whilst DCHA reduced energy metabolism and induced the opposite effects to spermidine, except for ascorbic acid content. Inhibition was reversed by exogenous spermidine. In conclusion, spermidine induced the accumulation of H2O2, enhanced the antioxidative system and improved the energy metabolism to enhance the functional quality of mung bean sprouts.

Effect of exogenous spermidine on floral induction, endogenous polyamine and hormone production, and expression of related genes in \u2018Fuji\u2019 apple (Malus domestica Borkh.)

Flower bud formation in ‘Fuji’ apple (Malus domestica Borkh.) is difficult, which severely constrains commercial production. Spermidine (Spd) plays an important role in floral induction, but the mechanism of its action is incompletely understood. To investigate the effect of Spd on flowering, 6-year-old ‘Fuji’ apple trees were treated with 1 × 10−5 mol L−1 Spd to study the responses of polyamines [putrescine (Put), Spd and spermine (Spm)], hormones [gibberellins (GA3) and abscisic acid (ABA)], and polyamine-, hormone- and flowering-related genes. Spd application promoted flowering during floral induction by increasing MdGA2ox2 (gibberellin 2-oxidase) through GA3 reduction and increasing MdNCED1 and MdNCED3 (9-cis-epoxycarotenoid dioxygenase) through ABA enrichment during 60 to 80 days after full bloom. The flowering rate as well as the expressions of flower-related genes, except for MdLEY (LEAFY), also increased, thereby promoting flowering. In addition, spraying with Spd significantly increased the contents of endogenous polyamines except for Spm in terminal buds by increasing the expressions of polyamine-associated genes. We hypothesize that the contribution of Spd to flowering is related to crosstalk among polyamines, hormone signals, and related gene expressions, which suggests that Spd participates in the apple floral induction process.

Exogenous spermidine enhances expression of Calvin cycle genes andphotosynthetic efficiency in sweet sorghum seedlings under salt stress

Salinity adversely affects plants resulting in disruption to plant growth and physiology. Previously, it has been shown that these negative effects can be alleviated by various exogenous polyamines. However, the role of spermidine (Spd) in conferring salinity tolerance in sorghum is not well documented. The effect of exogenous Spd on the responses of sweet sorghum (Sorghum bicolor L.) seedlings to salt stress (150 mM NaCl) was investigated by measuring photosynthetic carbon assimilation, Calvin cycle enzyme activities, and the the expression of respective genes. Application of 0.25 mM Spd alleviated the negative effects of salt stress on efficiency of photosystem II and CO2 assimilation and increased the activities of ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco) and aldolase. Salt stress significantly lowered the transcriptions of genes encoding Rubisco large subunit, Rubisco small subunit, 3-phosphoglyceric acid kinase, glyceraldehyde-3-phosphate dehydrogenase, triose-3-phosphate isomerase, fructose-1,6-bisphosphate aldolase, fructose-1,6-bisphosphate phosphatase, and sedoheptulose-1,7-bisphosphatase. However, transcriptions of genes encoding phosphoribokinase and Rubisco were up-regulated. The Spd application enhanced expressions of most of these genes. It appears Spd conferred salinity tolerance to sweet sorghum seedlings by enhancing photosynthetic efficiency through regulation of gene expressions and activities of key CO2 assimilation enzymes.

Polyamines are involved in GABA-regulated salinity-alkalinity stress tolerance in muskmelon

Salinity-alkalinity stress is a limiting environmental factor in muskmelon production. Treatment with γ-aminobutyric acid (GABA) protects crops against several abiotic stresses. Previous studies reported that GABA increased the content of polyamines (PAs) and improved the oxidative stress tolerance of muskmelon grown under salinity-alkalinity stress. However, it was unclear whether PAs were involved in GABA-regulated salinity-alkalinity stress tolerance. Here, we explored the roles of PAs in GABA-regulated Na+/K+ homeostasis and membrane lipid peroxidation. Our results showed that GABA pretreatment reduced the salinity-alkalinity stress-induced inhibition of muskmelon growth, significantly reduced the Na+/K+ ratio and enhanced the contents of free PAs, arginine, and methionine. GABA treatment upregulated the expression levels of PA-related genes under salinity-alkalinity stress. Application of exogenous spermidine reduced the salinity-alkalinity stress-induced inhibition of muskmelon growth, relative electrical conductivity, malondialdehyde content, and Na+/K+ ratio. By contrast, pretreatment of muskmelon plants with the Spd synthesis inhibitor dicyclohexylammonium sulfate (DCHA) eliminated GABA-remediated ion balance, membrane peroxidation injury, and plant growth under salinity-alkalinity stress conditions. In all, fed with GABA maintained the Na+/K+ balance via reducing Na+ content and increasing K+ uptake of muskmelon under salinity-alkalinity stress conditions, ultimately reduced membrane oxidation injury. And GABA provoked PAs levels involved in GABA-regulated Na+/K+ homeostasis and alleviation of membrane lipid peroxidation under salinity-alkalinity stress tolerance.

Spermidine Enhances Heat Tolerance of Rice Seeds by Modulating Endogenous Starch and Polyamine Metabolism.

Polyamines have been reported to be involved in grain filling and they might contribute to the construction of heat resistance of some cereals. In this study, the hybrid rice ‘YLY 689’ was used to explore the possible effects of exogenous spermidine (Spd) on seed quality under high temperature during the filling stage. Rice spikes were treated with Spd or its synthesis inhibitor cyclohexylamine (CHA) after pollination, and then the rice plants were transferred to 40 °C for 5-day heat treatment. The results showed that, compared with the control under high temperature, Spd pretreatment significantly improved the germination percentage, germination index, vigor index, seedling shoot height, and dry weight of seeds harvested at 35 days after pollination, while the CHA significantly decreased the seed germination and seedling growth. Meanwhile, Spd significantly increased the peroxidase (POD) activity and decreased the malondialdehyde (MDA) content in seeds. In addition, after spraying with Spd, the endogenous content of spermidine and spermine and the expression of their synthetic genes, spermidine synthase (SPDSYN) and spermine synthase (SPMS1 and SPMS2), significantly increased, whereas the accumulation of amylose and total starch and the expression of their related synthase genes, soluble starch synthase II-3 (SS II-3) and granules bound starch synthase I (GBSSI), also increased to some extent. The data suggests that exogenous Spd pretreatment could alleviate the negative impacts of high temperature stress on rice seed grain filling and improve the rice seed quality to some extent, which might be partly caused by up-regulating endogenous polyamines and starch metabolism.

Effects of exogenous spermidine and spermine on antioxidant metabolism associated with cold-induced leaf senescence in Zoysiagrass (Zoysia japonica Steud.)

Zoysiagrass (Zoysia japonica Steud.) leaves lose their green colour as they undergo cold acclimation (CA) in late fall in many regions, and delaying leaf senescence can extend their green period. The objective of this study was to investigate whether exogenous spermidine (Spd) and spermine (Spm) can delay leaf senescence by enhancing reactive oxygen species (ROS)-scavenging antioxidant enzyme activity. Mature zoysiagrass (cv. Zenith) plants were treated with Spm (1 mM) or Spd (1 mM) or water (control). The treated plants were grown under normal conditions or subjected to cold acclimation CA for 35 days. CA treatment increased leaf electrolyte leakage (EL) and malondialdehyde (MDA) level, decreased chlorophyll (Chl) content, and induced leaf senescence. Foliar application of Spd and Spm reduced the decline in Chl content and reduced EL and MDA levels. CA treatment increased the ROS level and reduced antioxidant enzyme activity. Application of Spm and Spd reduced ROS accumulation and enhanced the activities of superoxide dismutase, peroxidase, catalase, and ascorbate peroxidase under CA conditions. The results of this study indicated that exogenous Spd and Spm at 1 mM can delay CA-induced leaf senescence through enhancement of ROS-scavenging antioxidant activity in zoysiagrass.

Tricarboxylic acid cycle activity suppresses acetylation of mitochondrial proteins during early embryonic development in Caenorhabditis elegans

The tricarboxylic acid (TCA) cycle (or citric acid cycle) is responsible for the complete oxidation of acetyl-CoA and formation of intermediates required for ATP production and other anabolic pathways, such as amino acid synthesis. Here, we uncovered an additional mechanism that may help explain the essential role of the TCA cycle in the early embryogenesis of Caenorhabditis elegans. We found that knockdown of citrate synthase (cts-1), the initial and rate-limiting enzyme of the TCA cycle, results in early embryonic arrest, but that this phenotype is not because of ATP and amino acid depletions. As a possible alternative mechanism explaining this developmental deficiency, we observed that cts-1 RNAi embryos had elevated levels of intracellular acetyl-CoA, the starting metabolite of the TCA cycle. Of note, we further discovered that these embryos exhibit hyperacetylation of mitochondrial proteins. We found that supplementation with acetylase-inhibiting polyamines, including spermidine and putrescine, counteracted the protein hyperacetylation and developmental arrest in the cts-1 RNAi embryos. Contrary to the hypothesis that spermidine acts as an acetyl sink for elevated acetyl-CoA, the levels of three forms of acetylspermidine, N1-acetylspermidine, N8-acetylspermidine, and N1,N8-diacetylspermidine, were not significantly increased in embryos treated with exogenous spermidine. Instead, we demonstrated that the mitochondrial deacetylase sirtuin 4 (encoded by the sir-2.2 gene) is required for spermidine's suppression of protein hyperacetylation and developmental arrest in the cts-1 RNAi embryos. Taken together, these results suggest the possibility that during early embryogenesis, acetyl-CoA consumption by the TCA cycle in C. elegans prevents protein hyperacetylation and thereby protects mitochondrial function.

Exogenous spermidine elevating cadmium tolerance in Salix matsudana involves cadmium detoxification and antioxidant defense

In this study, exogenous spermidine role on Salix matsudana tolerance to cadmium was evaluated. Spermidine and cadmium presented antagonistic effects on the biomass, copper and zinc concentrations in S. matsudana. cadmium mainly distributed in the cell wall of subcellular fraction; 46.97%–60.43% of cadmium existed in a sodium chloride-extracted form. Cadmium contents in roots, leaves, and twigs ranged from 2002.67 to 3961.00, 111.59 to 229.72, and 102.56 to 221.27 mg/kg, respectively. Spermidine application elevated cadmium concentrations in the roots, cuttings, and cell wall and the ratio of deionized water-extracted cadmium, but decreased cadmium levels in the twigs and leaves and the fractions of cadmium extracted by ethanol and sodium chloride, respectively. Putrescine and malondialdehyde were important indicators of cadmium-induced oxidative damage. Exogenous spermidine alleviated the accumulation of superoxide anion, hydrogen peroxide, malondialdehyde via promoting the levels of spermidine, soluble protein, superoxide dismutase, reductive ascorbate, glutathione reductase, and glutathione peroxidase in S. matsudana leaves under the corresponding cadmium stress. The results indicated that S. matsudana was a candidate for cadmium rhizoremediation and extraction in leaves; the spermidine application enhanced the cadmium tolerance of S. matsudana through promoting cadmium accumulation in roots, cell wall, and less bioactive chemical forms and the antioxidative ability.

Correction to: Effects of Exogenous Spermidine and Elevated CO2 on Physiological and Biochemical Changes in Tomato Plants Under Iso-osmotic Salt Stress

The original version of this article unfortunately contained errors in two authors' names. The given and family names of the authors were incorrectly swapped as Zhang Yi and Shi Yu instead the authors' names should be Yi Zhang and Yu Shi.

Spermidine-Regulated Biosynthesis of Heat-Stable Antifungal Factor (HSAF) in Lysobacter enzymogenes OH11.

Heat-Stable Antifungal Factor (HSAF) and its analogs are antifungal natural products produced by the biocontrol agent Lysobacter enzymogenes. The production of HSAF is greatly influenced by environmental stimuli and nutrients, but the underlying molecular mechanism is mostly unclear. Here, we found that HSAF production in L. enzymogenes OH11 is strictly controlled by spermidine, which is the most prevalent triamine in bacteria. When added into OH11 cultures, spermidine regulated the production of HSAF and analogs in a concentration-dependent manner. To verify the role of spermidine, we deleted LeSDC and LeADC genes, encoding S-adenosylmethionine decarboxylase and arginine decarboxylase, respectively, that are the key enzymes for spermidine biosynthesis. Both deletion mutants produced barely detectable spermidine and HSAF including its analogs, whereas the antifungals production was restored by exogenous spermidine. The results showed that the OH11 cells must maintain a proper spermidine homeostasis for the antifungals production. Indeed, the expression level of the key HSAF biosynthetic genes was significantly impaired in LeSDC and LeADC mutants, and exogenous spermidine restored the gene expression level in the mutants. Ornithine is a key substrate for HSAF biosynthesis, and OH11 genome contains arg1 and arg2 genes, encoding arginases that convert arginine to ornithine. While the expression of arg1 and arg2 was affected slightly upon mutation of LeSDC and LeADC, exogenous spermidine significantly increased the arginase gene expression in LeSDC and LeADC mutants. Together, the data revealed a previously unrecognized mechanism, in which spermidine controls antibiotic production through controlling both the biosynthetic genes and the substrate-production genes.

Exogenous Spermidine Promoted Ca2+ Absorption in Lettuce Roots and Reduced the Incidence of Tipburn

Exogenous Spermidine Promoted Ca2+ Absorption in Lettuce Roots and Reduced the Incidence of Tipburn

Effects of Exogenous Spermidine and Elevated CO2 on Physiological and Biochemical Changes in Tomato Plants Under Iso-osmotic Salt Stress

The original version of this article unfortunately contained errors in two authors' names. The given and family names of the authors were incorrectly swapped as Zhang Yi and Shi Yu instead the authors' names should be Yi Zhang and Yu Shi.

Leaf gas exchange and grain yield of common bean exposed to spermidine under water stress

Three prevalent aliphatic polyamines (PAs) include putrescine, spermidine, and spermine; they are low-molecular-mass polycations involved in many physiological processes in plants, especially, under stressful conditions. In this experiment, three bean (Phaseolus vulgaris L.) genotypes were subjected to well-watered conditions and two moderate and severe water-stressed conditions with and without spermidine foliar application. Water stress reduced leaf relative water content (RWC), chlorophyll contents, stomatal conductance (gs), intercellular CO2 concentration (Ci), transpiration rate, maximal quantum yield of PSII (Fv/Fm), net photosynthetic rate (PN), and finally grain yield of bean plants. However, spermidine application elevated RWC, gs, Ci, Fv/Fm, and PN, which caused an increase in the grain yield and harvest index of bean plants under water stress. Overall, exogenous spermidine could be utilized to alleviate water stress through protection of photosynthetic pigments, increase of proline and carotenoid contents, and reduction of malondialdehyde content.