Effects of polyamines on the early development of cystocarps,spore release,survival,and germination rate of Gracilariopsis lemaneiformis

Three polyamines(PAs)(spermidine(Spd),spermine(Spm),and putrescine(Put))were used as growth regulators to the marine macroalgae Gracilariopsis lemaneiformis and to female gametophyte and reproductive tissues(cystocarps were constructed),which led to early maturation of cystocarps and spore release.Laboratory cultivation revealed that exogenous PA application accelerated the development of cystocarps,and the mean sporeling growth rate was 20%–50%/d during the initial week of cultivation.The spore count(SC),survival rate(SR),and germination rate(GR)of G.lemaneiformis showed different responses to PA(10-6 mol/L and 10-3 mol/L)treatments under different temperatures(18,26,and 34℃),light intensities(30,60,and 90μmol photons/(m^(2)·s)),salinities(25,30,and 35),and nutrient levels(25꞉2.5,50꞉5.0,and 75꞉7.5μmol/L NH4Cl꞉K2HPO4),and the optimal conditions were 26℃,light intensity of 60μmol/(m^(2)·s),salinity of 35,and nutrient level of 50꞉5.0μmol/L NH4Cl꞉K2HPO4.Under the optimum conditions,the SC per cystocarp increased by 82.38%,and the spore SR and GR increased by 33.07%and 46.44%,respectively.Each experiment lasted for 7 days,with the highest SC on Day 4 and plateauing on Day 7.The results indicate that PAs could promote the early maturation,spore release,survival,and germination of G.lemaneiformis under laboratory conditions.

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.

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.

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.

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.

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.

Protective Effect of Hydrogen Peroxide against Paraquat Toxicity in Young Pea Plants: Possible Role of Endogenous Polyamines

Young pea plants grown as a water culture were sprayed with hydrogen peroxide and 24 h later were treated with the herbicide paraquat. The effects of paraquat and H2O2 treatment were investigated on the endogenous concentrations of free proline, and free and conjugated putrescine, spermidine and spermine. Application of PQ increased the free proline and Put, and decreased conjugated and bound polyamines. The results clearly demonstrate the oxidative damages provoked by PQ application. Single treatment with H2O2 provoked considerable decrease in the free fraction of the three PAs studied 2 h after light exposition, but caused an increase in their respective conjugated fractions. Data suggest that PA changes after H2O2 treatment are due to conversion from free to conjugated form. Three hours later free PAs in H2O2 treated-plants reached control levels;however the conjugated remained higher than the control. No alterations were detected in proline concentration after H2O2 treatment. Pretreatment with Н2О2 activated some component of the plant protection mechanisms by causing an alteration in free/conjugated PAs ratio in plants subsequently subjected to PQ treatment. Data concerning PA and proline concentrations in plants treated with both compounds supposed that H2O2 shows a protective role against PQ and improves the plant tolerance to the oxidative stress generated by paraquat.

Assessment of Polyamines and Trehalose in Wheat Microspores Culture for Embryogenesis and Green Regenerated Plants

Most aspects of microspore culture protocol have the capacity to cause stress to microspores, hence, less stressful treatments might be required to avoid deleterious effects. In stressed plants, polyamines and trehalose can act as compatible solutes or osmoprotectants by stabilizing proteins and biological membranes. To improve green plant regeneration in wheat microspore culture, this study assessed the effects of polyamines (putrecine, spermidine, spermine) and trehalose on androgenic response namely embryogenesis, green plant regeneration and ploidy of green plants regenerated in three spring wheat genotypes. Microspores of the genotypes produced significant numbers of embryos and green plants among polyamine treatments but trehalose had no effect (P ≤ 0.05). Polyamine treatments for 30 min generally produced more green plants per 100 microspores than the 60 min treatments in all three genotypes. At least three out of twelve polyamine treatments in each genotype improved the production of double haploid plants and seed setting in regenerants. Wheat genotype, concentration and duration of polyamine treatment had significant impact on embryogenesis and regeneration of green plants in this study. The study also showed that polyamines could be used to accelerate cultivar development in wheat breeding.

Yield-related agronomic traits evaluation for hybrid wheat and relations of ethylene and polyamines biosynthesis to filling at the mid-grain filling stage

Because of the yield increase of 3.5-15%compared to conventional wheat,hybrid wheat is considered to be one of the main ways to greatly improve the wheat yield in the future.In this study,we performed a principal component analysis(PCA)on two-line hybrids wheat and their parents using the grain weight(GW),the length of spike(LS),the kernel number of spike(KSN),and spike number(SPN)as variables.The results showed that the variables could be classifed into three main factors,the weight factor(factor 1),the quantity factor 1(factor 2)and the quantity factor 2(factor 3),which accounted for 37.1,22.6 and 18.5%,respectively of the total variance in different agronomic variables,suggesting that the GW is an important indicator for evaluating hybrid combinations,and the grain weight of restorer line(RGW)could be used as a reference for parents selection.The hybrid combination with a higher score in factor 1 direction and larger mid-parent heterosis(MPH)of the GW and its parents were used to carry out the analysis of grain fling,1-aminocylopropane-1-carboxylicacid(ACC)and polyamine synthesis related genes.The results suggested that the GW of superior grain was significantly higher than that of inferior grains in BS1453xJS1(H)and its parents.Both grain types showed a weight of H between BS 1453(M)and JS1(R),and a larger MPH,which may be caused by their differences in the active fling stage and the grain fling rate.The ADP-glucose pyrophosphorylase(AGPase),granule bound starch synthase I(GBSS/),starch synthaseⅡ(SSS),and starch branching enzyme-Ⅰ(SBE-1)expression levels of H were intermediated between M and R,which might be closely related to MPH formation of the GW.Compared with R and H,the GW of M at maturity was the lowest.The expression levels of spermidine synthase 2(Spd2),ornithine decarboxylase(ODC)and S adenosylmethionine decarboxylase(SAMDC)had significantly positive correlations with the grain flig rate(1=0.77,0.51,0.59"),suggesting their major roles in the grain flling and heterosis formation.These provide a theoretical basis for improving the GW of photo-thermo-sensitive male sterile lines(PTSMSL)by changing the endogenous polyamine synthesis in commercial applications.

Interactive Effects of Drought Stress and Phytohormones or Polyamines on Growth and Yield of Two M(<i>Zea maize</i>L) Genotypes

Two maize genotypes (Nefertiti and Bashaier) were picked up from nine maize genotypes during the early vegetative growth (25 days) to be cultivated in open field upon the crop yield under the different drought stress levels (90,70,50,30) or under the interaction effect of drought stress and phytohormones or polyamines. According to the data of growth criteria, the maize genotype Nefertiti was found to be the most drought sensitive genotype, while the genotype Bashaier was found to be the most drought resistant genotype. Additionally while the photosynthetic pigments remained more or less unchanged in genotype Bashaier, their biosynthesis destroyed earlier in the drought sensitive genotype (Nefertiti). Also while the genotype Bashaier absorbed and accumulated a sufficient amount of mono and divalent cations (K+, Ca++ and Mg++), the genotype Nefertiti did not. Accordingly while the genotype Bashaier gave a crop yield up to 50% field capacity, the genotype Nefertiti gave a crop yield only up to 70% field capacity and failed to give a crop yield beyond this level. The interaction effect of drought stress and phytohormones and polyamines improved the all above characteristics. Interestingly each of these activators considerably improved the production of crop yield only in genotype Bashaier specially polyamines they produced more than 60% field capacity and at the level of 30% field capacity (the level which did not give crop yield in this genotype). However, phytohormones in generally did not make an important effect on the crop yield in genotype Nefertiti although they improved the dry matter production during the vegetative stages. Such situation seemed to be complicated and borne many questions to be studied in the future.

Effects of Polyamines Synthesis Inhibitors Sprayed after Heading on Contents and Components of Polyamines in Rice Grains at the Ripening Stage

With a higher polyamine rice （indica variety Jinlingxiang） as experimental material, the changes in contents and components of polyamines （PAs） in rice grains after harvesting were investigated under the treatments by spraying the panicles and flag leaves of rice with PAs synthesis inhibitors, D-Arg （0.5, 1.0, 1.5 and 2.0 mmol/L）, methylglyoxal bis-guanylhydrazone （MGBG） （0.1, 0.5, 1.0 and 1.5 mmol/L） and cyclohexylamine （CHA） （1.0, 5.0, 10.0, 15.0 and 20.0 mmol/L）, at the heading, flowering and grain filling stages. The inhibition of D-Arg on the contents of free and bound PAs and its total amounts in rice grains was significant, especially treated at the flowering and grain filling stages. Treatments with MGBG and CHA affected mainly the components of PAs, and their effects on the total amounts of PAs were not significant except MGBG at the flowering stage and CHA at both heading and flowering stages.

Polyamines and polyamine-metabolizing enzymes in schizophrenia: Current knowledge and concepts of therapy

Polyamines play preeminent roles in a variety of cellular functions in the central nervous system and other organs.A large body of evidence suggests that the polyamine pathway is prominently involved in the etiology and pathology of schizophrenia.Alterations in the expression and activity of polyamine metabolizing enzymes,as well as changes in the levels of the individual polyamines,their precursors and derivatives,have been measured in schizophrenia and animal models of the disease.Additionally,neuroleptic treatment has been shown to influence polyamine concentrations in brain and blood of individuals with schizophrenia.Thus,the polyamine system may appear to be a promising target for neuropharmacological treatment of schizophrenia.However,for a number of practical reasons there is currently only limited hope for a polyamine-based schizophrenia therapy.

Branched Polyamines Functionalized with Proposed Reaction Pathways Based on ¹H-NMR, Atomic Absorption and IR Spectroscopies

Three novel branched polyamines N,N,N’,N’-tetrakis-[3((pyridine-2-methyl)-amine) propyl]-1,4- butanediamine (1), N,N,N’,N’-tetrakis-[N-((2-methylpyridine)ethyl)propanamide]ethylenediamine (2) and N,N,N’,N’-tetrakis-[3((2-hidroxibenziliden)-amine)propyl]-1,4-butanediamine (3), were synthesized starting from 2-pyridinecarboxaldeyde with DAB-Am-4 for 1, PAMAM G0 for 2 and from salicylaldehyde with DAB-Am-4 for 3. The pathway reactions have been proposed by 1H-NMR, IR and Atomic Absorption Spectroscopy. The optimal reaction time was set by IR spectroscopy following aldehyde? peak modification. 1 and 2 were obtained as both hydrochlorides and as free amines and 3 only as free imine. These polyamines were characterized by UV-Vis, IR, 1H-NMR and 13C-NMR and Mass Spectrometry.

Role of Polyamines in Ozone Exposed Ischemic-Reperfused Hearts

The effect of chronic ozone exposure to ischemia reperfusion (I/R) injury in isolated perfused rat hearts was previously demonstrated. The present study tested our hypothesis that chronic ozone exposure led to attenuation of polyamines in the heart, which may limit sensitivity to I/R. Sprague Dawley rats were continuously exposed for 8 hrs/day for 28 days to filtered air or 0.8 ppm ozone. Isolated hearts were previously subjected to 0.5 hour of global ischemia followed by 1 hour of reperfusion after which global polyamine content was examined between the two groups. Spermidine production was significantly increased in the experimental group compared to control group (of I/R hearts). These results suggest that ozone-induced sensitivity to chronic I/R injury activates myocardial polyamine stress response characterized by increased enzymatic activities and accumulation of spermidine. Collectively, these results suggest that I/R possibly disturbs polyamine metabolism, and increased oxidative stress and concomitant reduced myocardial cell viability.

Polyamines in developing rice grains and their relations with grain filling

Many studies have reported that polyamines have an effect on a number of growth and developmental process in plants, such as cell division, embryogenesis, senescense, and flowering. However, there are few reports on the physiological significance in of polyamines grain filling of rice. We studied the level of polyamines in the developing grains and their relations with grain filling and grain weight, using 8 rice varieties including hybrid combinations. It was showed that the content of Put (puterscine), Spd (spermidine), and Spm (spermine), especially Spd and Spm, was

Changes of Protective Enzymes and Accumulation of Polyamines Induced by Boron Deficiency in Roots of Rape Seedlings

The effects of boron deficiency on the membrane permeability, the lipid peroxidation of membrane, the activities of the protective enzymes and the accumulation of polyamines in the roots of rape ( Brassica napus L. cv Zhongyou 821) plants were examined using solution culture experiment. Compared to the 20 mmol B m -3 treatment, boron deficient treatment (2 mmol B m -3 ) decreased root dry weight and increased the rate of solute leakage and malondialdehyde (MDA) concentration in the roots of rape. Similar patterns of change were observed in the level of lipid peroxidation (MDA concentration) and the rate of solute leakage under boron deficiency. Results suggested that a significant alteration of membrane composition had occurred under B deficiency. The concentration of putrescine and the ratio of putrescine to spermidine + spermine in the roots of rape increased significantly, and the activities of superoxide dismutase and catalase decreased in the roots of rape during B deficiency. But peroxidase activity in root of B deficient plant was higher than that of control plant.

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.

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.

FUNCTIONS OF POLYAMINES IN ROSIN SIZING UNDER NEUTRAL PAPERMAKING CONDITIONS

Functions of the polyamines in neutral rosin sizing were investigated using X-ray photoelectron spectroscopy (XPS) and laser diffraction particle analysis. The polyamine with a higher charge density and a smaller unite size could retain more rosin and cover larger fiber surfaces. The XPS spectra demonstrated that polyallylamine (PAAm) and polyvinylamine (PVAm) could react with a rosin size to form a -OC-N-CO- structure, but polydimethylamino ethyl methacrylate (PDMAEMA) could not. The formation of this structure may be a key step for effective sizing.