Effects of osmotic priming and ageing on onion seed germination

Onion seeds were osmotically primed in polyethylene glycol solution (342 g/kg water) either for 14 days before accelerated ageing at 40°C. 18% m.c. for 0, 24, 48, 72 or 96 h, or for 10, 14 or 17 days after ageing. Priming improved the rate of germination compared with non-primed seed. Priming before ageing delayed the loss of viability due to ageing, but priming after ageing had no effect on viability. Primed and dried onion seed was stored for 18 months at 10°C, 9% m.c. with no effect on viability; improvements in germination rate due to priming were maintained over the storage period. Conductivity measurements of seed leachates were not a consistently reliable indicator of germination performance.     

The effects of re-use of polyethylene glycol priming osmotica upon the development of microbial populations and germination of leeks and carrots

Three batches of leek seeds were osmotically primed successively in the same polyethylene glycol solution in a bubble column at a seed concentration of 100 g/litre for seven days at 15°C. Three batches of carrot seeds were similarly primed in a separate solution for six days at 15°C. The concentration of microorganisms in the solutions increased rapidly during priming of the first seed batch for both seed types, but increases during priming of the second and third batches were small. The seeds were the main source of the microorganisms; priming reduced the numbers of colonies of filamentous fungi and increased those of bacteria and yeasts. The priming treatments improved the percentage germination of the three seed batches of primed carrots and reduced the mean time to germination in both species and the mean time to emergence in compost. Percentage emergence was not affected by priming except in the third batch of primed carrot seed. The presence of large numbers of microorganisms in the priming solutions did not greatly affect seed performance when the same osmoticum was used three times with leeks and twice with carrots. Priming did not affect the number of abnormal seedlings.     

The Effects of Priming and Ageing on Resistance to Deterioration of Tomato Seeds

The influence of seed priming and ageing treatments on viabilityand rate of germination of tomato (Lycopersicon esculentum Mill.)seeds was examined under both long-term and controlled-deteriorationstorage conditions. Seeds of a single lot of tomato were eitherprimed or aged to increase or decrease the rate of germination(Argerich and Bradford, 1989). They were then stored at 6% moisturecontent (dry weight basis) at either 4 ?C or 30 ?C for 1 year.Both viability and germination rate were unaffected by eitherstorage temperature in control seeds, or by 4 ?C storage inprimed or aged seeds. At 30 ?C, however, viability and germinationrate of primed and aged seeds was markedly reduced after 6 monthsof storage. The temperature dependence of the germination rateand the spread of germination times within the population wasalso adversely affected by high temperature storage, particularlyfor primed seeds. Under controlled deterioration conditions(13.5% moisture content and 50 ?C), the rate of loss of viabilitywas greater for primed seed than for control or aged seeds.The relationship between seed viability and the mean germinationrate, however, was not influenced by the seed treatments. Thesedata are analysed in relation to current models of seed deteriorationduring storage and seed repair during priming. The results indicatethat enhancement of seed germination rates by priming treatmentssimultaneously lowers the resistance of seeds to deterioration.Primed tomato seeds must, therefore, be considered to be vigorousseeds with a reduced storage life. Key words: Tomato, controlled deterioration, seed germination rate, seed viability     

Effect of osmotic priming, rinsing and storage on the germination and emergence of carrot seed

Carrot seed, osmotically primed in polyethylene glycol solution (273 g/kg water) for 10 days at 15°C, was given 0–5 rinsing cycles, where each cycle comprised agitating the seed in 10 times its own bulk volume of distilled water for 20 to 30 s. Priming decreased the germination and emergence times whether the seeds were rinsed or not, and these effects were maintained even after storage for 450 days. The number of rinsing cycles had no effects, except that seed given five rinsing cycles deteriorated after 270 days storage.     

The Effects of Priming and Accelerated Ageing upon the Nucleic Acid Content of Leek Seeds and their Embryos

Using seed priming and accelerated ageing techniques, a singlelot of leek (Allium porrum) seeds was manipulated to producefour lots of seeds with different germination performance. Changesin content of the major nucleic acid species in whole seedsand embryos of two of these lots (primed and unprimed), weredetermined over the early stages of germination. The major effectof priming was an increased level of RNA species in the seedsand embryos, and this difference was maintained during germination.Comparison of nucleic acid levels in the dry seeds of thesetwo lots and two others (aged and aged then primed) indicatedthat there was no correlation with germination performance.Similar comparisons of the nucleic acid levels in the embryosof seeds imbibed for 1 d showed only a limited correlation betweenrRNA levels and germination performance. Analysis of these datasuggests that accelerated ageing has an adverse effect uponendosperm cells, which results in the degradation of their nucleicacids during priming. Furthermore, the viability of these agedseeds also falls during priming. The data also indicate thatratios of rRNA to DNA correlate with germination performanceof the four lots of seeds studied. It is proposed that sucha relationship is indicative of the efficiency of a primingtreatment, and may be useful in comparisons of naturally varyingseed lots. Key words: Leek, seed, germination, priming, nucleic acids     

Effects of matriconditioning on onion seed germination, seedling emergence and associated physical and metabolic events

The effect of matriconditioning, the physiological presowing seed technique, using Micro-Cel E on Allium cepa L. cv. Czerniakowska seed quality was studied. Several ratios of seeds, carrier, water and time of priming were tested. The most effective treatment for improving onion seed germination at most tested temperatures was priming to a ratio of 2 g seed:1 g Micro-Cel:3 g water for 5 days in light at 15 °C. Matriconditioning greatly improved the germination and emergence percentage, seedling fresh and dry weight and reduced electrolyte leakage compared to that of untreated seeds; this beneficial effect was especially evident at suboptimal temperatures. Matriconditioning improved the germinability of aged seeds, the effect being more pronounced in the more aged seeds. No significant differences in ethylene production by primed and non-primed seeds were observed in the absence of its precursor, 1-aminocyclopropane-1-carboxylic acid (ACC), but its presence during imbibition caused an increase in ethylene production; an enhanced activity of in vivo ACC oxidase in Allium cepa matriconditioned seeds in comparison to untreated seeds, indicates that the endogenous level of ACC is a limiting factor of ethylene production. Likewise, the activity of ACC oxidase isolated from matriconditioned seeds was higher than that from untreated seeds. Higher endo--mannanase and total dehydrogenase activities were observed in primed air-dried seeds in comparing to non-primed seeds.     

Biochemical Changes During Osmopriming of Leek Seeds

Osmotic priming treatments reduced both the mean time to germinationand the spread of germination for two leek seed-lots of highviability but differing vigour. In addition the differencesin germination performance between these two seed-lots was abolishedby the priming treatments. In the unprimed seed-lots, differencesin germination performance were reflected in differences inrates of protein biosynthesis in leek embryo tissue during germination.Osmopriming treatments abolished these differences upon subsequentgermination of osmotically primed seed and furthermore inducedhigh levels of protein biosynthesis in embryo tissue. DNA synthesiswas detectable in leek embryos during the priming period inthe absence of any cell division and was followed by a five-foldincrease in the rate of DNA synthesis in embryo tissue upongermination following priming at which time the rates of DNAsynthesis in these leek embryos was significantly greater thanthat found at any time over the first 4 d of germination inembryos of unprimed leek seeds. The increases in rates of bothprotein and DNA synthesis observed upon germination of primedseed occurred only after a 6–12 h lag period during whichtime there is little increase in these rates above those foundat the end of priming Analysis of nucleotide and nucleotide sugar levels in leek embryosboth during and after priming showed that only traces of GTPand CTP and low levels of ATP and UTP were present in embryosduring priming. After a 6 h lag period following the end ofpriming these levels increased sharply, probably via de novosynthesis. A similar pattern was found for UDP glucose levelsduring priming and subsequent germination. These results indicatethat there is considerable biochemical activity during primingand that the significant benefits in germination performanceof primed leek seeds is accompanied by marked increases in protein,DNA and nucleotide biosynthesis after a lag period of 6–12h following the end of the priming period Allium porrum, leek, seed, osmopriming, germination, protein synthesis, nucleic acids, nucleotides, nucleotide sugars     

Responses of Vegetable Seeds to Controlled Hydration

Leek, onion and carrot seeds were imbibed in water and in solutionsof polyethylene glycol (PEG) 6000 over the range –0.5to –4.0 MPa osmotic potential, for periods up to 28 dat 15 C. Seeds started to germinate after 7 and 14 d at –0.5MPa and –1.0 MPa PEG, respectively, but in the lattercase, germination did not exceed 5%. No germination occurredin solutions of lower (more negative) osmotic potential. Seedmoisture content increased with osmotic potential in all threespecies and the relationships were unaffected by the durationof imbibition in solutions of –1.0 to –4.0 MPa,though leek seeds had higher moisture contents than the otherspecies for any given osmotic potential. Linear relationships between response to priming (differencebetween mean germination times of primed and untreated seeds)and seed moisture content were obtained for each species, positiveresponses being obtained above 30–35% seed moisture contentwith optima at 46, 44.5 and 44% seed moisture contents in leek,onion and carrot, respectively. Priming had no effect on embryovolume or cell number per embryo in leek and onion. Carrot embryovolume increased by 43% and cell number per embryo doubled inprimed compared with untreated seeds, whereas seeds imbibedin water showed only a slight increase in cell number per embryoat the stage when radicles were beginning to penetrate the seedcoat. Allium cepa L. cv. Rijnsburger Robusta, onion, Allium porrum L. cv. Winterreuzen, leek, Daucus carota L. cv. Nantaise, carrot, germination, priming, polyethylene glycol, seed moisture, cell number, embryo volume     

Comparative evaluation of hydro-, chemo- , and hormonal-priming methods for imparting salt and PEG stress tolerance in Indian mustard (<Emphasis Type="Italic">Brassica</Emphasis><Emphasis Type="Italic">juncea</Emphasis> L.)

The present study was aimed to evaluate the effect of different seed priming methods to enhance the sodium chloride (NaCl) and polyethylene glycol-8000 (PEG-8000) stress tolerance in Indian mustard (Brassica juncea L.). Seeds subjected to different priming treatments such as water (hydro-priming), calcium chloride (CaCl₂) (chemo-priming), and abscisic acid (ABA) (hormonal-priming) showed increased rate of germination as compared to non-primed seeds. The primed and non-primed seeds were grown for 15 days and then the seedlings were independently subjected to iso-osmotic salt (150 mM NaCl) or PEG-8000 (20%) stress. The different biochemical responses were studied 10 days after treatment. Under NaCl and PEG stress, the dry weight and total chlorophyll content were higher in primed sets as compared to non-primed treatment which was also evident by the phenotype of the seedlings. In general, the higher activities of superoxide dismutase and glutathione reductase resulted in lower oxidative damage, in terms of malondialdehyde content, under NaCl and PEG stress in hydro-primed set as compared to non-primed, ABA-, and CaCl₂-primed treatments. Besides, the level of total phenolics and accumulation of osmolytes such as free proline, glycine betaine, and total soluble sugars was also lower in hydro-primed set as compared to other primed and non-primed treatments. The study thus suggests the use of hydro-priming as a simple and cost-effective strategy to alleviate the NaCl and PEG induced stress in B. juncea.     

Prehydration and Priming Treatments that Advance Germination also Increase the Rate of Deterioration of Lettuce Seeds

The influence of prehydration in water or priming in –1.5 MPa polyethylene glycol 8000 solution for various periods,followed by redrying, on germination rate and longevity of lettuce(Lactuca sativa L.) seeds (achenes) was determined during controlleddeterioration at 10% moisture content (fresh weight basis) and40°C. Short prehydration treatments (up to 1 h) had littleeffect on either germination rate or longevity, but significantlyimproved root growth rates. Increasing durations of prehydrationor priming reduced the mean time to germination by up to 61%relative to untreated seeds, but also reduced mean seed longevityby as much as 84% Prehydration and priming altered the relationshipsbetween germination rate and viability and between normal andabnormal seedlings during ageing. Prehydration in abscisic acidor at a temperature inhibitory to germination did not preventthe reduction in longevity under controlled deterioration conditions.While prehydration or priming treatments effectively acceleratesubsequent germination rates of lettuce seeds, the redried seedsare nonetheless highly susceptible to deterioration in storage. Key words: Lettuce, Lactuca sativa L., seed priming, seed deterioration, germination rate     

Osmoconditioning as a Means of Counteracting the Ageing of Pepper Seeds during High-temperature Storage

Sweet pepper seeds were osmotically conditioned in 0.4 M mannitolsolution for 4 d (at 25 °C, in darkness) before or afterstorage at 35 °C for up to six months, and their germinationand viability was compared with that of untreated seeds storedunder the same conditions. Seeds that had been osmoconditionedprior to storage retained a high rate of germination and germinatedto a high final percentage (from 80 to 50 per cent) at both15 and 25 °C throughout the storage period. By contrast,both the rate and total level of germination of untreated pepperseeds declined rapidly at both germination temperatures, andby three months of storage the total level of seed viabilitywas already less than 10 per cent. Seeds that were first storedat 35 °C, and then osmoconditioned just prior to germination,showed a decline in germinability which when tested at 25 °Cwas the same as for untreated seeds, while tested at 15 °Coccurred at a slightly slower rate than for untreated seeds. It is evident that osmoconditioning prior to storage, in additionto the acceleration of germination, resulted in a dramatic delayof the ageing rate, thus increasing considerably the longevityof seeds. On the other hand, osmoconditioning after storagedid not seem to have any significant effect on seed viability,though it enhanced the germination rate. Capsicum annuum, sweet pepper, seed, germination, osmoconditioning, priming, storage, viability, ageing, longevity     

Aspirin priming circumvents the salinity-induced effects on wheat emergence and seedling growth by regulating starch metabolism and antioxidant enzyme activities

Though seed priming has been emerged as an effective and pragmatic approach, efforts are being made to discover and optimize the new priming agents which are cheaper and easily accessible to the farmers. Here, we established two independent experiments, to ascertain the role of aspirin priming in salinity tolerance of wheat. In the first experiment, various concentrations of aspirin (125, 250, 375, and 500 ppm) were examined for emergence and seedling growth of wheat. A non-primed control, hydropriming, and hydrogen peroxide priming treatments were also maintained for comparison. Among the different treatments, seeds primed with 125 and 250 ppm aspirin depicted better emergence, vigorous seedling growth, and higher starch metabolism. Therefore, these treatments were further used in the second experiment under salinity stress (10 dS m^?1). Salinity stress caused delayed and erratic emergence hampered the shoot and root growth, chlorophyll contents, and enhanced the lipid peroxidation and phenolics content in wheat seedlings. However, wheat seed priming particularly with aspirin effectively alleviated the negative effects of salinity on most of the observed parameters. Aspirin priming also significantly enhanced the activities of antioxidant enzymes (catalase, peroxidase, and superoxide dismutase), and reduced oxidative stress in wheat seedlings. Vigorous growth and greater salinity tolerance of wheat seedlings derived from aspirin primed seeds were related with better starch metabolism, strong antioxidative defense system, and lower lipid peroxidation.     

‘On-farm’ seed priming with zinc in chickpea and wheat in Pakistan

A series of on-station trials was implemented between 2002 and 2006 to assess the response of wheat (Triticum aestivum L.) and chickpea (Cicer arietinum) to zinc (Zn) added by soaking seeds (priming) in solutions of ZnSO₄ before sowing. Wheat seed was primed for 10 h in 0.3% Zn and chickpea for 6 h in 0.05% Zn. Seed treatments increased the seed concentration in wheat from 27 to 470 mg/kg and in chickpea from 49 to 780 mg/kg. Priming wheat seeds with 0.3% Zn significantly increased the mean shoot dry mass, Zn concentration and Zn uptake of 15-day-old seedlings relative to non-primed controls and seeds primed with water alone. Using 0.4% Zn further increased shoot Zn concentration but depressed shoot dry mass to the level of the non-primed control. In seven trials, mean grain yield of wheat was significantly increased from 2.28 to 2.42 t/ha (6%) by priming with water alone and to 2.61 t/ha (14%) by priming with 0.3% Zn. Mean grain yield of chickpea in seven trials was increased significantly from 1.39 to 1.65 t/ha (19%) by priming seeds with 0.05% Zn. The effect of priming chickpea seeds with water was intermediate (1.49 t/ha) and not statistically separable from the non-primed and zinc-primed treatments. Increased grain yield due to priming in both crops was associated with increases in total biomass but there was no significant effect of priming on harvest index. In addition to increasing yield, priming seeds with Zn also significantly increased grain zinc concentration, by 12% in wheat (mean of three trials) and by 29% in chickpea (one trial) and the total amount of Zn taken up by the grain (by 27% in wheat and by 130% in chickpea). Using ZnSO₄ to prime seeds was very cost-effective, with net benefit-to-cost ratios of 75 for wheat and 780 for chickpea. An erratum to this article can be found at     

Hydro and halo priming: influenced germination responses in wheat Var-HUW-468 under heavy metal stress

The beneficial effect of seed priming in improving critical growth stages like seed germination and early growth phases has been accepted by Plant Physiologists for many important field crops. In the present investigation, studies were made to see the effect of heavy metal stress imposed during germination using solution of HgCl₂ in four different concentrations (0.0, 0.50, 0.75 and 1.00 mM) in Petri dishes on primed and non-primed seeds of wheat. Priming has been done with distilled water (hydro), Mg(NO₃)₂ and Ca(NO₃)₂ (halo) salts. Different germination parameters, such as germination percentage, radicle and plumule lengths, seedling emergence, soluble and insoluble sugar contents and activity of α-amylase in endosperm were studied at different study periods. Primed seeds increased all the germination parameters except insoluble sugar content in respect to non-primed control in the absence of HgCl₂. However, the use of primed seeds has shown to overcome the inhibitory effects of heavy metal stress imposed in the form of HgCl₂ solution during the period of germination. Hence, the work concludes the mitigating effects of priming under heavy metal stress.     

The use of an ELISA to quantitate the extent of 11S globulin mobilization in untreated and primed sugar beet seed lots

Seed priming (controlled imbibition) is a widely used technique for improving crop establishment, because it allows a reduction of the time to radicle emergence following seed imbibition and synchronization of individual seeds within seed lots with respect to germination timing. The major problem encountered in seed priming is the control of seed imbibition to a level permitting pre-germinative processes to proceed but that blocks radicle emergence. If not, the consequence of drying back the seeds to initial moisture content for storage purposes could be a total loss of the treated batch. This is because, as long as radicle growth has not begun, seeds may be re-dried without any permanent deleterious effects upon subsequent germination or growth. Recently, we reported the discovery of a molecular marker of sugar beet seed priming, corresponding to the basic B-subunit of the seed storage protein 11S globulin. An ELISA based upon this molecular marker has been used to analyse how different sugar beet seed lots respond to a priming treatment. The results demonstrate that this ELISA allows us to readily distinguish between the primed seeds and the corresponding untreated seeds.     

A comparison of two large-scale seed priming techniques

Leek seed lots of high (91%) and low (82%) viability were primed in aerated polyethylene glycol (PEG) solutions in Bubble-columns and by a non-osmotic priming technique in both 1987 and 1988 and the seeds were then sown in the field. Both methods of priming established a similar seed moisture content during treatment and, in the laboratory, produced seeds with more rapid and uniform germination than untreated seeds, with a greater advantage for Drum compared with Bubble-column priming in PEG. In the field, both priming techniques gave seedling emergence responses similar to those from priming in PEG by the laboratory-scale technique on filter paper. Both large-scale priming methods gave earlier and more uniform emergence than untreated seeds and gave similar or slightly higher levels of seedling emergence, except on one sowing occasion when seeds were stored before sowing followed by sowing into a drying seedbed.     

The Effects of Priming and Ageing on Seed Vigour in Tomato

A comparison was made of the effects of seed priming or ageingtreatments on the performance of tomato (Lycopersicon esculentumMill. cv. UC204C) seeds according to a number of indices ofseed vigour. A single lot of tomato seeds was primed in 120mol m^–3 K2HPO4 + 150 mol m^–3 KNO₃ for 5 d at 20?C, or aged at 13% moisture content (dry weight basis) and 50?C for 6 d. Germination percentage (>98%) was unaffectedby priming and reduced to 85% by ageing. X-ray photographs andlongitudinal sections revealed the formation of free space surroundingthe embryo in dry primed seeds, which was not evident in controlor aged seeds. Priming increased the rate of germination atall temperatures above the base temperature (T_b), while ageingdecreased it. T_b was unaffected by priming and only slightlyincreased by ageing. The variation in individual times to germinationwas approximately doubled in both primed and aged seed comparedto the control, based upon the slopes of probit germinationpercentage versus log thermal time curves. Root growth aftergermination tests and seedling growth in both greenhouse andfield tests were not influenced by either priming or ageing.The conductivity test was found to be unreliable as a vigourtest for tomato seeds. The results identify several indiceswhich can be used to quantify seed vigour in tomato. They alsoillustrate that seed priming can enhance seed performance accordingto some criteria, while having no effect or decreasing qualityaccording to other criteria. Seed vigour can apparently be separatedinto various components which can be independently influencedby seed enhancement treatments. Key words: Tomato, seed germination rate, seed priming, seed vigour     

Physiological and enzymatic activity of pepper seeds (Capsicum annuum) during priming

Pepper ( Capsicum annuum L. cv. Keystone Resistance Giant 3) seeds were monitored during priming to determine if seed treatments which accelerate the rate of germination could be correlated with specific physiological changes within the seeds. Pepper seeds primed with −0.90 and −1.35 MPa NaCl solutions at 23°C for 18 days did not completely equilibrate with the osmotic potential of the priming solution. Seed respiratory rates indicated that priming extends the lag phase of germination following imbibition. Soluble protein levels increased 115% in primed seeds, and the uptake and incorporation of [¹⁴C(U)] labelled amino acids into the acid insoluble fraction increased throughout the priming treatments. Alcohol dehydrogenase (EC 1.1.1.1, anaerobic metabolism), glucose-6-phosphate dehydrogenase (EC 1.1.1.49) and 6-phosphogluconate dehydrogenase (EC 1.1.1.44, pentose phosphate pathway) activities remained stable throughout the priming treatment, but were higher after 6 days. than the water-imbibed controls. Aldolase (EC 4.2.1.1. glycolysis) and isocitrate lyase (EC 4.1.3.1, glyoxylate cycle) activities increased with imbibition and were 61 and 56% (respectively) higher in primed seeds as compared to the water-imbibed controls after 12 days. Treatment with the −0.90 MPa NaCl solution was more effective than the −1.35 MPa solution in improving the rate of germination, yet there were no significant differences between the protein concentrations or enzyme activities of the two priming treatments. However, the incorporation of labelled amino acids into pepper seeds was significantly higher in the −0.90 MPa priming treatment.     

Priming of seeds with NaCl induces physiological changes in tomato plants grown under salt stress

The effects of seed priming with 6 M NaCl solution have been investigated with respect to growth and physiological responses of tomato plants ( Lycopersicon esculentum Mill. cv. Pera) exposed to 70 and 140 m M NaCl nutrient solutions from 11 to 60 days after sowing. Tomato seedlings from primed seeds emerged earlier than from non-primed seeds. At 70 m M , a lower shoot and root dry weight reduction was found in plants from primed seeds at the different harvests (30, 45 and 60 days after sowing), while at 140 m M the positive effect of seed priming was only shown in roots. Significant changes in Na⁺ and CI⁻ accumulation with seed priming were only found in roots at 60 days after sowing, with ion accumulation in roots being higher in plants grown at 70 and 140 m M from primed seeds. In leaves of salt-treated plants, significant increases in sugars and organic acids with seed priming were found from 30 days after sowing, and these increases were higher at longer treatment times. In roots, however, only the organic acids tended to increase in plants from primed seeds, although they increased less than in leaves, especially at 60 days after sowing. These results support the hypothesis that priming of seeds with NaCl induces physiological changes in the plants, changes which are shown more clearly at advanced growth stages.     

Wet-dry cycling extends seed persistence by re-instating antioxidant capacity

Seeds in the field experience wet-dry cycling that is akin to the well-studied commercial process of seed priming in which seeds are hydrated and then re-dried to standardise their germination characteristics. To investigate whether the persistence (defined as in situ longevity) and antioxidant capacity of seeds are influenced by wet-dry cycling, seeds of the global agronomic weed Avena sterilis ssp. ludoviciana were subjected to (1) controlled ageing at 60% relative humidity and 53.5°C for 31 days, (2) controlled ageing then priming, or (3) ageing in the field in three soils for 21 months. Changes in seed viability (total germination), mean germination time, seedling vigour (mean seedling length), and the concentrations of the glutathione (GSH) / glutathione disulphide (GSSG) redox couple were recorded over time. As controlled-aged seeds lost viability, GSH levels declined and the relative proportion of GSSG contributing to total glutathione increased, indicative of a failing antioxidant capacity. Subjecting seeds that were aged under controlled conditions to a wet-dry cycle (to ?1 MPa) prevented viability loss and increased GSH levels. Field-aged seeds that underwent numerous wet-dry cycles due to natural rainfall maintained high viability and high GSH levels. Thus wet-dry cycles in the field may enhance seed longevity and persistence coincident with re-synthesis of protective compounds such as GSH.