Possible Role of Cellular Polyamine Metabolism in Neuronal Apoptosis

Recent studies have shown that cellular levels of polyamines(PAs)are significantly altered in neurodegenerative diseases.Evidence from in vivo animal and in vitro cell experiments suggests that the cellular levels of various PAs may play important roles in the central nervous system through the regulation of oxidative stress,mitochondrial metabolism,cellular immunity,and ion channel functions.Dysfunction of PA metabolism related enzymes also contributes to neuronal injury and cognitive impairment in many neurodegenerative diseases.Therefore,in the current work,evidence was collected to determine the possible associations between cellular levels of PAs,and related enzymes and the development of several neurodegenerative diseases,which could provide a new idea for the treatment of neurodegenerative diseases in the future.

A PET probe targeting polyamine transport system for precise tumor diagnosis and therapy

Polyamine metabolism dysregulation is a hallmark of many cancers,offering a promising avenue for early tumor theranostics.This study presents the development of a nuclear probe derived from spermidine(SPM)for dual-purpose tumor PET imaging and internal radiation therapy.The probe,radiolabeled with either[68Ga]Ga for diagnostic applications or[177Lu]Lu for therapeutic use,was synthesized with exceptional purity,stability,and specific activity.Extensive testing involving 12 different tumor cell lines revealed remarkable specificity towards B16 melanoma cells,showcasing outstanding tumor localization and target-to-non-target ratio.Mechanistic investigations employing polyamines,non-labeled precursor,and polyamine transport system(PTS)inhibitor,consistently affirmed the probe?s targetability through recognition of the PTS.Notably,while previous reports indicated PTS upregulation in various tumor types for targeted therapy,this study observed no positive signals,highlighting a concentration-dependent discrepancy between targeting for therapy and diagnosis.Furthermore,when labeled with[177Lu],the probe demonstrated its therapeutic potential by effectively controlling tumor growth and extending mouse survival.Investigations into biodistribution,excretion,and biosafety in healthy humans laid a robust foundation for clinical translation.This study introduces a versatile SPM-based nuclear probe with applications in precise tumor theranostics,offering promising prospects for clinical implementation.

Effects of Salinity Stress on the Levels of Covalently and Noncovalently Conjugated Polyamines in Plasma Membrane and Tonoplast Isolated from Barley Seedlings

When the 7_d old barley ( Hordeum vulgare L.) seedlings were treated with different concentrations of NaCl for 3 d, the levels of the noncovalently conjugated polyamines (PAs) in the plasma membrane and tonoplast vesicles and the covalently conjugated PAs in the membrane proteins were promoted by NaCl of low concentrations and suppressed by NaCl of high concentrations. Among the noncovalently conjugated PAs in the vesicles, spermidine (Spd) level was the most abundant, while putrescine (Put) content was predominant among the covalently conjugated PAs, accounted for 40%-70%, 35%-60%, respectively. In addition, the TLC (thin_layer chromatography) profiles of the benzoylated PAs presented an unknown polyamine with Rf =0.92 (X 0.92 ), which conjugated covalently and noncovalently in root tonoplast and its content changed as well as Spd with NaCl treatment. The total PA contents in the roots were higher than that in the leaves, and the types and contents of covalently and noncovalently conjugated PAs in the tonoplast were higher than those in the plasma membrane. The results showed that the above two PAs associated with the membrane might be essential in salt adaption of cells and the maintenance of membrane function.

The Role of Polyamine Oxidative Degradation in the Process of Lettuce Seed Germination

[Objective]The aim of this study was to understand the role of polyamine oxidative degradation in the process of lettuce seed germination. [Method]After lettuce seed soaking treatments with aminoguanidine （AG,a specific inhibitor of polyamine oxidases）,seed germination rate,activities of polyamine oxidase （PAO） and diamine oxidase （DAO）,change of endogenous polyamine and H2O2 content were determined. [Result]Compared with the control,AG treatment strongly inhibited the seed germination,which also had an extremely significant difference in seed germination rate after incubation for 12 h. During the seed germination,activities of PAO and DAO significantly changed,while their activities firstly increased and then decreased,then DAO and PAO reached peaks at 24 h and 48 h respectively. AG treatment was strongly inhibitory for activities of DAO and PAO,whose activities even disappeared after incubation with AG for 24 h and 36 h. During the seed germination,endogenous Put reduced sharply in the first 24 h,then reduced slowly in 24-60 h,while Spd decreased slowly in the early stage of germination,and then sharply declined after 48 h. However,Spm content was low but slightly increased,and the total polyamine gradually decreased. AG treatment could significantly increase endogenous polyamines,especially Put and Spd contents. During the seed germination,H2O2 content gradually increased,and had a peak from 36 to 48 h,then kept a high level at last. AG treatment could significantly reduce H2O2 content. [Conclusion]During the seed germination,the changes of endogenous polyamine and H2O2 content correspond with the changes of PAO and DAO activities,which indicate that there is an active polyamine metabolism of oxidative degradation during the lettuce seed germination.

Relationship Between Polyamine Metabolism in Roots and Salt Tolerance of Barley Seedlings

Plant growth rate (GR), contents of free polyamines (fPAs) and bound polyamines (bPAs) and activities of some key enzymes involved in polyamine (PA) metabolism in the roots of two barley (Hordeum valgare L.) cultivars differing in salt sensitivity were investigated with 0-300 mmol/L NaCl treatments. With 0-200 mmol/L NaCl treatments, activities of arginine decarboxylase (ADC) and transglutaminase (TGase) and PA oxidase (PAO) in the roots of barley seedlings all increased, while TGase and PAO activities decreased slightly at 300 mmol/L NaCl. As a result, free Put (fPut) content increased continuously with increasing concentrations of NaCl, while levels of free Spd (fSpd) and an unknown PA (fPAx) and bPAs (bPut, bSpd and bPAx), as well as (fSpd + fPAx)/fPut ratio rose at 50-200 mmol/L NaCl and reduced at 300 mmol/L NaCl. However, no significant change in the tetra-amine spermine (Spin) content was observed. Statistical analysis showed that GR was very significantly positively correlated with (fSpd + fPAx)/fPut ratios and the contents of bPAs, whereas a significant inverse correlation existed between GR and the ratios of fPA contents to bPA levels. These results showed that, under salt stress, the balance between fSpd, fPAx and fPut levels and an equipoise between fPA and bPA contents in roots were important to salt tolerance of barley seedlings.

Effects of Polyamine on Leaf Physiological and Biochemical Indexes of Maize Seedlings under Osmotic Stress

[Objective]The research aimed to probe into the mechanism of polyamine enhancing the tolerance of plants to drought stress and provide theoretical basis for application of polyamine in process of maize drought resistance. [Method]With PEG-6000 simulating natural drought,the change in content of soluble protein and relative water content were investigated in seedling leaves of two maize cultivars,Nongda 108 and Yedan 13 under osmotic stress with exogenous Spd treatment. [Result]On the 7th day,leaf relative water content and the content of soluble protein decreased more significantly in leaves of Yedan 13 （drought-sensitive） than in Nongda 108 （drought-tolerant）. Exogenous Spd treatment not only obviously inhibited the decrease of leaf relative water content,but also increased the content of soluble protein. [Conclusion]Exogenous Spd treatment could enhance the tolerance of maize seedlings to osmotic stress,via improving the content of soluble protein in seedling leaves.

Mechanism of the Effect of Polyamines on the Activity of Tonoplasts of Barley Roots Under Salt Stress

The seeds of barley Hordeum vulgare L. cv. Jian 4) were soaked with 0.1 mmol/L putrescine (Put) and 0.5 mmol/L spermidine (Spd), and then the seedlings were treated with 200 mmol/L NaCl. The growth rate (GR), dry matter accumulation, distribution of ions, the amount of polyamines (PAs) bound to tonoplast proteins as well as lipid composition and the activity of tonoplast vesicles isolated from roots were investigated. The results showed that soaking with Put or Spd could retard salt injury, promote GR and dry matter accumulation, and increase K+/Na+ in the roots. Compared with NaCl_treated plants, phospholipid content in root tonoplast rose by soaking with Put and Spd, while the level of galactose in lipids was decreased. Moreover, the ratio in noncovalently conjugated PA contents of (Spd+PAx (an unknown PA)) to (Put+Dap (diaminopropane)), and the total contents of covalently and noncovalently conjugated PAs were all increased. Statistical analysis indicated that the ratio of (Spd+PAx) to (Put+Dap) was significantly and positively correlated with the activities of membrane associated enzymes H+_ATPase and H+_PPase.

Effects of Polyamines on Plants under Salt Stress

Salt stress is one of the worldwide abiotic stresses resulting in growth re- duction, chlorosis, wilting, and plant death. These exhibitions might result from men- tal toxicity and osmotic stress induced by salt. The two aspects of stress would af- fect vital metabolic pathways, reactive oxygen species scavenging system, lipid per- oxidation and photosynthetic apparatus. Thus, exploring ways to improve crop pro- ductivity or alleviate harmful effects under salt is one of the major areas of concern. Polyamines are aliphatic nitrogen organic cations which are implicated in a wide range of plant physiological processes such as morphogenesis, flower differentiation and initiation, they also play a role in biotic or abiotic stress responses. At the physiological level, polyamines modify the activities of many enzymes included in salt stress response and can bond to photosynthetic apparatus, thus changing the photosynthetic efficiency. At molecular level, polyamines can modify expressions of the polyamine-related genes directly or indirectly. Significant researches had been done to understand the effects of polyamines on plant salt resistance, but several questions still need to be answered. The present review is focused specifically on the effects of polyamines on physiological and molecular changes in plants under salt stress.

Effect of Polyamine Priming on Chilling Tolerance of Lolium perenne during Seed Imbibition

[Objective] This study aimed to investigate the effect of potyamine priming on physiological and biochemical variations of Lolium perenne embryos and seed germination. [Method] With annual Lolium perenne （Diamond T and Grazing-8000） as experimental materials, after priming with 0.5 mmol/L putrescine （Put）, spermidine （Spd） and spermine （Spm） for 24 h and chilling imbibition at 5 ℃ for 12, 24, 36 and 48 h, the effect of Put, Spd and Spm priming on chilling tolerance and germination ability of annual Lolium perenne seeds during imbibition was investigated. [Result] Put, Spd and Spm priming improved the activities of peroxidase （POD）, catalase （CAT）, superoxide dismutase （SOD） and ascorbate peroxidase （APX） and content of soluble protein content under low temperature stress, significantly in-creased the germination rate, and shortened the average germination duration. After chilling imbibition for 48 h, compared with the control, the average germination rate of annual Lolium perenne seeds was improved by 15.5% and 12.0% after Put, Spd and Spm priming, and the average germination duration was shortened by 1.21 and 1.14 d. During seed imbibition, the chilling tolerance of Grazing-8000 was stronger than that of Diamond T. Overall, Put, Spd and Spm treatment could increase the chilling tolerance of annual Lolium perenne seeds during imbibition, and improve the germination ability of seeds under low temperature stress. [Conclusion] Results of this study provided theoretical basis for the application of seed priming technology in the production of annual ryegrass.

两种预氧化法与氯胺消毒联用对polyamine形成NDMA的影响

预氧化处理与有机高分子絮凝剂联用会使含氮消毒副产物亚硝基二甲胺(NDMA)质量浓度上升,典型水处理工艺条件下NDMA的质量浓度为60～100 ng/L,为了解不同因素的影响及其反应机理,研究次氯酸钠氯化预氧化及高锰酸钾预氧化与聚环氧氯丙烷二甲胺(polyamine)在不同条件下联用并用氯胺消毒后NDMA的生成量.结果表明:随着预氧化剂投加量、pH、polyamine投加量及Br-质量浓度的增加,NDMA的生成量也相应增加;低质量浓度NH_4^+会导致NDMA生成量增加17. 5%左右,但随质量浓度增加,NDMA生成量逐渐减少至初始水平;随着NO_2^-质量浓度增加,NDMA生成量显著减少,当NO_2^-的质量浓度达到10 mg/L时,NDMA的生成量为最初的15%左右;氯化预氧化过程符合UDMH机理,而高锰酸钾预氧化过程包含UDMH机理及羟基自由基机理.

Effect of polyamine on seed germination of wheat under drought stress is related to changes in hormones and carbohydrates

Drought is a multifaceted stress condition that inhibits crop growth. Seed germination is one of the critical and sensitive stages of plants, and its process is inhibited or even entirely prevented by drought. Polyamines （PAs） are closely associ- ated with plant resistance to drought stress and seed germination. However, little is known about the effect of PA on the seed germination of wheat under drought stress. This study investigated the involvement of PAs in regulating wheat seed germination under drought stress. Six wheat genotypes differing in drought resistance were used, and endogenous PA levels were measured during seed germination under different water treatments. In addition, external PAwas used for seed soaking and the variation of hormones, total soluble sugar and starch were measured during the seed germination under different water treatments. These results indicated that the free spermidine （Spd） accumulation in seeds during the seed germination period favored wheat seed germination under drought stress; however, the free putrescine （Put） accumulation in seeds during the seed germination period may work against wheat seed germination under drought stress. In addition, seed soaking in Spd and spermine （Spm） significantly relieved the inhibition of seed germination by drought stress; howev- er, soaking seeds in Put had no significant effect on seed germination under drought. External Spd and Spm significantly increased the endogenous indole-3-acetic acid （IAA）, zeatin （Z）＋zeatin riboside （ZR）, abscisic acid （ABA）, and gibberellins （GA） contents in seeds and accelerated the seed starch degradation and increased the concentration of soluble sugars in seeds during seed germination. This may promote wheat seed germination under drought stress. In conclusion, free Spd and Put are key factors for regulating wheat seed germination under drought stress and the effects of Spd and Put on seed germination under drought notably related to hormones and starch metabolism.

Relationship between ATPase activity and conjugated polyamines in mito-chondrial membrane from wheat seedling roots under osmotic stress

The effects of osmotic stress on the ATPase activity, the contents of —SH group and conjugated polyamines in mitochondrial membrane from wheat seedling [Triticum aestivum L. cv. Yumai No.18(drought-tolerant) and cv. Yumai No.9(drought-sensitive)] roots were investigated. The results showed that ATPase activity and —SH group content decreased with polyethylene glycol(PEG) 6000(-0.55 MPa) treatment for 7 d, in concert with the decrease of the ratio of noncovalently conjugated spermidine(NCC-Spd)/noncovalently conjugated putrescine(NCC-Put) and increase of the covalently conjugated putrescine(CC-Put). Osmotic stress injury to Yangmai No.9 seedlings was alleviated greatly with 1 mmol/L exogenous spermidine(Spd), in concert with marked increases of the ratio of NCC-Spd/NCC-Put, —SH group contents and ATPase activity in mitochondrial membrane. Under osmotic stress, the concomitant treatment of Yumai No.18 seedlings with methylglyoxyl bis(guanylhydrazone) (MGBG), an inhibitor of S-adenosyl methionine decarboxylase(SAMDC), and phenanthrolin (o-Phen), an inhibitor of transglutaminase(TGase), caused a significant decrease of the ratio of NCC-Spd / NCC-Put, CC-Put contents, respectively, in concert with the marked decreases of ATPase activity, —SH group content and its tolerance to osmotic stress. All the results above suggested that osmotic stress tolerance of wheat seedlings was associated with the ATPase activity, the contents of —SH group, NCC-Spd and CC-Put in mitochondrial membrane.

Regulatory Function of Polyamine Oxidase-Generated Hydrogen Peroxide in Ethylene-Induced Stomatal Closure in Arabidopsis thaliana

Hydrogen peroxide （H2O2） is an important signaling molecule in ethylene-induced stomatal closure in Arabidopsis thaliana. Early studies on the sources of H2O2 mainly focused on NADPH oxidases and cell-wall peroxidases. Here, we report the involvement of polyamine oxidases （PAOs） in ethylene-induced H2O2 production in guard cells. In Arabidopsis epidermal peels, application of PAO inhibitors caused the failure of ethylene to induce H2O2 production and stomatal closure. Results of quantitative RT-PCR analysis and pharmacological experiments showed that AtPAO2 and AtPAO4 transcripts and activities of PAOs were both induced by ethylene. In transgenic Arabidopsis plants over-expressing AtPAO2 and AtPAO4, stomatal movement was more sensitive to ethylene treatment and H2O2 production was also significantly induced. The increased H2O2 production in the transgenic lines compared to the wild-type plants suggests that AtPAO2 and AtPAO4 probably are involved in ethylene-induced H2O2 production. Several factors which induce stomatal closure such as dehydration and high salinity all enhanced the expression of AtPAO2 and AtPAO4 to different degrees. Moreover, GFP- AtPAOs fusion protein localized in the nucleus, cytoplasm, and cell wall of the guard cells. Therefore, our results strongly indicated that PAO is a source of H2O2 generation in Arabidopsis guard cells and plays crucial roles in stomatal movement.

Relationship Between Polyamines Metabolism and Cell Death in Roots of Malus hupehensis Rehd. Under Cadmium Stress

The free putrescine （Put） content, the hydrogen peroxide （H202） content and the polyamine oxidase （PAO） activity in roots of Malus hupehensis Rehd. var. pinyiensis Jiang （PYTC） were significantly increased, and reached its peak at 1, 2 and 6 h, respectively, under cadmium treatment. The free spermine （Spin） and spermidine （Spd） contents were dramatically decreased, and reached the minimum value at 4-6 h, then remained relatively stable. The change in total free polyammes （PAs） content was consistent with that of free Put. The number of root dead cells was gradually increased after treatment for 24 h, and the typical characteristics of programmed cell death （PCD） were displayed at 48 h. Throughout the Cd treatment process, changes in PAs metabolism appeared to be prior to cell death increase, and the H2O2 content was always maintained at a high level. These results indicated that polyamines could initiate cell death by generating H2O2 in roots of Malus hupehensis Rehd. under CdSO4 stress.

Melatonin treatment induces chilling tolerance by regulating the contents of polyamine,γ-aminobutyric acid,and proline in cucumber fruit

The mechanism of melatonin(MT)induced chilling tolerance in harvested cucumber fruit was investigated at commercial maturity.In this study,cucumber fruits were treated with 100μmol L^(-1) MT at 4℃ and 90% relative humidity for 15 d of storage.In comparison with the control,cucumber treatment with MT resulted in reduced chilling injury(CI),decreased electrolyte leakage and enhanced firmness.The fruits treated with MT showed higher chlorophyll contents in storage conditions with suppressed chlorophyllase enzyme activity.MT treatment increased arginine decarboxylase(ADC)and ornithine decarboxylase(ODC)enzyme activities.Moreover,enhanced expression of the Cucumis sativus ADC(CsADC)and C.sativus ODC(CsODC)genes resulted in the accumulation of polyamine contents.Similarly,proline levels exhibited higher levels among treated fruits.Meanwhile,the proline synthesizing enzymes △1-pyrroline-5-carboxylate syntheses(P5CS)and ornithine aminotransferase(OAT)were significantly increased,while a catabolic enzyme of proline dehydrogenase(PDH)activity was inhibited by treatment.In addition,MT induced expression of C.sativus OAT(CsOAT)and C.sativus P5CS(CsP5CS)genes.Cucumber fruits treated with MT also exhibited higher γ-aminobutyric acid(GABA)content by enhanced GABA transaminase(GABA-T)and glutamate decarboxylase(GAD)enzyme activities and a higher C.sativus GAD(CsGAD)gene expression.To sum up,the results show that MT treatment enhanced chilling tolerance,which was associated with the regulation of polyamines,as well as proline and γ-aminobutyric acid.

Catalytic hydrolysis of phosphate diester (BNPP) and plasmid DNA by mononuclear macrocyclic polyamine metal complexes

The activities of the catalytic hydrolysis of phosphate diester （BNPP） [bis（p-nitrophenyl）phosphate diester] and plasmid DNA （pUC 18） by mononuclear macrocyclic polyamine metal complexes have been investigated in this paper. The results showed that the highest activity in hydrolysis of BNPP was obtained with le--Zn（II） complex （composed of lipophilic group） as catalyst. The hydrolysis rate enhancement is up to 3.64 × 10^4 fold. These metal complexes could effectively promote the cleavage of plasmid DNA （pUC18） at physiological conditions.

Relationship between food polyamines and gross domestic product in association with longevity in Asian countries

The relationship between gross domestic pro- duct (GDP) and dietary profile, with a focus on polyamine intake, was investigated in 35 Asian countries. Data on food supply, GDP, and health condition were collected from databases of the United Nations, the International Monetary Fund, and the World Health Organization, respectively. The amount of polyamine intake from food was estimated using polyamine concentrations listed in published sources. The amounts of putrescine, spermidine, spermine, and total polyamines in 1000 kcal of food in Asian countries were 39.07 ± 17.98, 33.74 ± 14.35, 14.05 ± 6.60 and 86.85 ± 33.96 μmol/1000 kcal, respectively. Putrescine, spermidine, and spermine constituted 44, 39, and 17% of total polyamine, respectively. Vegetables contributed the largest amount of both putrescine and spermidine and the second largest amount of spermine (45.5%, 62.2% and 27.2% of total putrescine, spermidine, and spermine, respectively). Meat was the richest source of spermine and contributed the greatest amount (50%) of this polyamine. We showed a significant positive association between GDP and the amount of polyamine per 1000 kcal of food. Lifespan was associated with both GDP and the amount of polyamine per energy quotient of food. As several basic research studies have shown that polyamines help prolong longevity, it follows that polyamines may have a role in determining the lifespan of humans.

Effect of diethylstilbestrol on polyamine metabolism in hamster epididymis

<abstract>Aim: To investigate the effect of diethylstilbestrol (DES), one of the most potent endocrine disruptors, on the metabolism of polyamines in hamster epididymis. Methods: Male golden hamsters of 7-week-old were kept under a light and dark cycle of 14 h and 10 h for 1 week to stimulate maximally the gonadal function. DES was injected subcutaneously at doses of 0.01 mg·kg-1·day-1, 0.1 mg·kg-1·day-1 and 1 mg·kg-1·day-1 for one week. Results: DES treatment caused a significant decrease in the weight of epididymis. The activity of epididymal ornithine decar boxylase (ODC) increased 1 day after DES treatment, kept at a high level for 4 days and then decreased to nearly normal level at day 7. The activity of spermidine/spermine N1-acetyltransferase (SSAT) also increased transiently after DES treatment. The contents of putrescine, spermidine, spermine and N1-acetylspermidine were increased 1 day -4 days after DES treatment and restored to normal at day 7. All these changes showed a marked difference between the caput and the cauda. Conclusion: The polyamine biosynthesis in the hamster epididymis can be affected by DES, a xenoestrogen. DES may probably affect polyamine metabolism in the epididymis by regulating the rate-limiting enzymes involved in the polyamine biosynthesis.

Endogenous Levels of Polyamines under Water-Deficit Stress during Cotton’s Reproductive Development

Polyamines, putrescine (PUT), spermidine (SPD) and spermine (SPM) are implicated in plants’ responses under conditions of abiotic stress. Previous research in other crops has indicated that polyamines and changes in their concentrations are associated with drought tolerance under conditions of water-deficit stress;however, no information exists on cotton (Gossypium hirsutum L.). Growth chamber experiments were conducted with two cotton cultivars differing in drought tolerance, ST5288B2F (drought-sensitive) and Siokra L23 (drought-tolerant) in order to investigate the distribution of free polyamines, the effect of water-deficit stress on the polyamine metabolism of cotton reproductive units and their subtending leaves as well as the possible relationship between polyamines and drought tolerance in cotton. Our results showed that cotton ovaries contained significantly higher levels of total free polyamines compared to their subtending leaves under both control and water stress conditions. Water-deficit stress significantly increased PUT concentrations in ST5288B2F, while SPM levels significantly decreased in Siokra L23. The results indicated that water-deficit stress significantly affected cotton polyamine metabolism in reproductive structures and their subtending leaves;however, no clear relationship between drought-tolerance and changes in polyamine accumulation was established. Further research is needed to elucidate the mechanism according to which water-deficit stress affects polyamine metabolism.

Polyamines mediate the effect of post-anthesis soil drying on starch granule size distribution in wheat kernels

Polyamines(PAs) are important endogenous plant growth regulators responding to environmental stress and mediating many physiological processes including grain filling in cereals.This study investigated whether PAs mediate the effect of post-anthesis soil drying on starch granule size distribution,starch content,and weight of superior and inferior kernels of wheat(Triticum aestivum L.).Two wheat cultivars were grown in pots.Three treatments,well-watered(WW),moderate soil drying(MD) and severe soil drying(SD),were imposed from 9 days post-anthesis until maturity.PA levels in kernels and small,medium and large granules were measured.The results showed that superior kernels had much higher free spermidine(Spd) and free spermine(Spm) concentrations,larger volumes of medium starch granules,and smaller-sized large granules than did inferior kernels under all the treatments.Compared to WW,MD significantly increased the concentrations of free Spd and free Spm,activities of soluble starch synthase and granule-bound starch synthase,volume of medium granules,and starch content and kernel weight of inferior kernels,and decreased the size of large granules.SD produced the opposite effect.Application of Spd or Spm to spikes produced effects similar to those of MD,and application of an inhibitor of Spd and Spm synthesis produced effects similar to those of SD.These results suggest that PAs mediate the effect of post-anthesis soil drying on starch biosynthesis in wheat kernels by regulating key enzymes in starch synthesis and that elevated PA levels under MD increase the volume of medium granules and kernel weight of inferior kernels.