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SEED PHYSIOLOGY, PRODUCTION & TECHNOLOGY

Effect of Priming Temperature on Storability of Primed sh-2 Sweet Corn Seed

Dep. of Agronomy, National Chung Hsing Univ., No. 250, Kuokuang Rd., Taichung, Taiwan 402, ROC

* Corresponding author (jmsung{at}nchu.edu.tw)

	ABSTRACT

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Priming offers a means to raise seed performance in many crop species, but the longevity of primed seed is generally decreased. The exact causes of the more rapid deterioration of primed seed are still not established. This study evaluated the effects of priming and storage temperatures on germination and antioxidative activities of sweet corn seed (Zea mays L.) carrying the shrunken-2 (sh-2) gene. Seed were solid-matrix primed in moistened vermiculite at 10, 15 or 20°C for 36 h, then air-dried to near original moisture level. Primed seed were stored at 25, 10, or -80°C for up to 12 mo. Solid-matrix priming improved germination, reduced lipid peroxidation, enhanced antioxidative activities, and increased seedling growth. Seed longevity was decreased when 20°C-primed seeds were stored at 25°C for 12 mo. Seed primed at 10 and 15°C had superior viability and vigor responses compared with nonprimed control seeds when they were stored at 25°C for 12 mo. Reduced storability of the 20°C-primed seeds was attributable to enhanced peroxidation and decreased antioxidative activities. Storage at 10 or -80°C extended the storability of matrix primed sh-2 seed for at least 12 mo. Enhanced antioxidative activity plays a role in maintaining the viability and vigor responses of solid matrix primed seed stored at cool (10°C) or subzero (-80°C) temperatures. Moreover, 10 or 15°C-primed sh-2 seed can retain viability for 12 mo, provided that the primed seed is stored at 10°C.

Abbreviations: AOS, activated oxygen species • APX, ascorbate peroxidase • ASC, ascorbate • CAT, catalase • DHA, dehydroascorbate • GR, glutathione reductase • GSH, reduced glutathione • GSSG, oxidized glutathione • MDA, malondialdehyde • MGT, mean germination time • SOD, superoxide dismutase

	INTRODUCTION

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SEED DETERIORATION is associated with loss of membrane integrity, changes in enzymatic activities, decline in protein and nucleic acid synthesis, and lesions in DNA (McDonald, 1999). These deteriorative changes have frequently been related to activated oxygen species (AOS)-induced oxidative injury (Hendry, 1993; Bernal-Lugo and Leopold, 1998; McDonald, 1999). Priming can reverse some of the aging-induced deteriorative events, and thus improve seed performance (Taylor et al., 1998). One crucial determinant of primed seed performance is postpriming storage environment. Long-term priming (10 d) of leek (Allium porrum L.) and onion (Allium cepa L.) seeds stored at 10°C retained viability after 1 yr (Drew et al., 1997). Carrot (Daucus carota L.) seed primed for 17 d lost some viability after 12 mo of 10°C storage (Dearman et al., 1987). Primed tomato (Lycopersicon lycopersicum L.) seed (7 d priming) stored at 10°C retained viability for 12 mo, but at 30°C viability was reduced (Alvarado and Bradford, 1988). A higher viability was reported for sweet pepper (Capsicum annuum L.) seeds primed for 4 d, following 3 yr of storage at 25°C (Thanos et al., 1989). In contrast, 1-d, short-term priming decreased the storage life of lettuce (Lactuca sativa L.) seed (Tarquis and Bradford, 1992). Tomato seed receiving 1 d of priming, however, had better storability (Van Pijlen et al., 1996). It appears that the effects of priming on seed longevity may be species specific, depending on the nature of the storage and priming conditions.

Van Pijlen et al. (1996) demonstrated that the adverse effects of priming during storage were caused by decreased DNA repair activity resulting from progression in the cell cycle. Increased lipid peroxidation, mediated by AOS attack during desiccation in the absence of protection mechanisms (e.g., superoxide dismutase and catalase) was also involved in reducing the longevity of primed seed (Bruggink et al., 1999).

The environment during priming also influences the seed response to priming. Bradford (1986) reported that priming at 15°C resulted in enhanced seed performance for beet (Beta vulgaris L.), carrot, celery (Apium graveolens L.), lettuce, onion, corn, and soybean [Glycine max (L.) Merr.]. Ozbingol et al. (1998) found that the optimum temperature (27–28°C) for priming was the same as the optimum temperature for germination of tomato seed. Haigh et al. (1986) concluded that temperatures during priming (15, 20, or 25°C) had little effect on subsequent emergence responses of onion seeds. These findings suggest that different temperatures must be evaluated for each crop species to determine which provides the best priming result.

Accumulated evidence shows that 20°C-primed sh-2 seed deteriorated more rapidly than nonprimed sh-2 seed when they were stored at 10 or 25°C for 12 mo (Chang and Sung, 1998). Since it is known that subzero storage temperature preserves seed better do than cool (10°C) and room (25°C) temperatures (Roos, 1989), it is reasonable to hypothesize that the longevity of primed seed stored at -80°C may be greater. Moreover, longevity may vary among the seed primed and stored at different temperatures. Thus, to test these hypotheses, we investigated the germination, seedling growth, and lipid peroxidation of sh-2 sweet corn seed primed at 10, 15, or 20°C stored at 25, 10, and -80°C for 12 mo. The changes in activity of several peroxide-scavenging enzymes in relation to priming, storage duration, and storage temperature are also measured. Collectively, these data provide more insights about how priming and storage temperatures affect sh-2 sweet corn seed longevity.

	MATERIALS AND METHODS

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Seed Materials
A commercially produced seed lot of sh-2 sweet corn hybrid cultivar Honey 236 (92 g kg^-1 on fresh weight basis) was obtained from a local vendor. For priming treatment, 135 replicates each of 150 g of seed were mixed with 300 g of vermiculite No. 3 to which 375 mL of distilled water were added (Sung and Chang, 1993), sealed in a plastic bag, mixed to provide uniform seed-substrate contact, and incubated at 10, 15, or 20°C for 36 h to test temperature effects. Chiu (2000) reported that, under 10, 15 or 20°C priming temperature, 36 h of hydration duration was optimum for the tested seed lot. The partially hydrated seeds were then separated from the vermiculite, and air-dried at 25°C for 48 h to near original moisture level. After drying, all the seeds were sealed in aluminum foil bags coated with polyethylene and stored at 25, 10, or -80°C (45 bags per storage treatment) for up to 12 mo.

Germination Test
Seed germination and moisture content were determined at 0, 3, 6, 9, or 12 mo storage. Laboratory germination tests were conducted in three replicates of 50 seeds. Seed was planted 1.5 cm deep in a plastic tray filled with 100 g No. 3 vermiculite and watered with 200 mL of distilled water. Seed was incubated in controlled chambers with 12-h photoperiod and 300 W m^-2 light intensity at 25°C and watered as necessary. Germination was defined as the point coleoptiles were visible above the vermiculite surface and counted daily for 10 d for mean germination time (MGT) calculation (Chang and Sung, 1998). Seed moisture contents (fresh weight basis) were determined by sampling three replicates (20 seeds per sample) and weighing seed before and after heating in a forced-air oven at 103°C for 24 h. Seedling dry weights were determined 14 d after sowing. Leaching tests were conducted in three 10-seeds replicates that were soaked in 20 mL of deionized water at 25°C for 24 h. Leachates were measured with a conductivity meter (Suntex, model 17A, Taipei, Taiwan; cell constant 1.217).

Determinations of Peroxidative Products
Malondialdehyde (MDA) and total peroxide were determined on three replicates of 5 seeds. The seeds were hand-ground in a mortar and pestle with 4 mL 50 g L^-1 trichloroacetic acid at 4°C and then centrifuged at 14 000 g for 20 min. The supernatants were used for MDA (Heath and Packer, 1968) and total peroxide (Sagisaka, 1976) determinations. For Maillard reaction product (as indicated by browning intensity) determinations, three replicates of 5 seeds were hand-ground with 4 mL of deionized water at 4°C in motar and pestle and then centrifuged at 14 000 g for 20 min. The supernatants were used for browning intensity measurements (Yen and Lai, 1987)

Determinations of Enzymes Activities
For enzyme activity measurements, three replicates of 5 seeds were hand-ground at 4°C in a mortar and pestle with 4 mL of 0.1 M potassium phosphate buffer (pH 7.0), followed by centrifugation at 20 000 g for 20 min. The supernatants were used for determination of enzyme activity. Superoxide dismutase (SOD; EC. 1.15.1.1) activity was assayed according to the method of Stewart and Bewley (1980). Catalase (CAT; EC. 1.11.1.6) activity was assayed according to the method of Kato and Shimizu (1987). Glutathione reductase (GR; EC 1.6.4.2) activity was determined according to the method of Foster and Hess (1980). Ascorbate peroxidase (APX; EC 1.11.1.11) and dehydroascorbate reductase (DHAR; EC 1.8.5.1) activities were determined by the methods of Nakano and Asada (1981). The SOD activity was expressed as the unit of enzyme activity needed to inhibit the reaction by half per g fresh weight per min. The activities of CAT, DHAR, GR, and APX were expressed per gram fresh weight, and one unit represented 1 µmol of substrate undergoing reaction per minute. Portions of enzyme extract were used for determination of soluble protein (Sung and Chang, 1993).

Determinations of Glutathione and Ascorbate Contents
Glutathione was determined on three replicates of 5 seeds. The seeds were hand-ground in a cold mortar and pestle with 4 mL ice-cold 50 g L^-1 sulfosalicyclic acid and centrifuged at 20 000 g for 20 min. The supernatants were used for total glutathione, reduced glutathione (GSH), and oxidized glutathione (GSSG) determinations (Smith, 1985). For ascorbate and dehydroascorbate determinations, a modification of the method of Law et al. (1983) was used. Three replicates of 5 seeds were homogenized in a cold mortar and pestle with 4 mL ice-cold 50 g L^-1 trichloracetic acid solution and centrifuged at 16 000 g for 20 min, and 5 µL of supernatant was used for total ascorbate and ascorbate (ASC) determinations. Dehydroascorbate (DHA) content was deduced as the difference between total ascorbate and ascorbate contents.

Statistics
The experimental design for emergence test was a randomized complete block design with three replicates. Percentage data were arcsin-transformed before analysis. All data were subjected to an analysis of variance, and a LSD was calculated when a significant (P < 0.05) F ratio occurred for treatment effects.

	RESULTS

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Germination and Seedling Dry Weight
Unstored nonprimed seeds used in the study had 88% germination and 4.1 d MGT (Fig. 1A) . No changes in seed moisture content occurred during storage (Fig. 1G). Nonprimed seeds stored at 25°C for up to 6 mo showed no evident decline in germination percentage, but marked reductions in germination percentage and extensions in MGT (Fig. 1D) were observed for nonprimed seeds stored for 9 and 12 mo. Germination percentage and MGT of nonprimed seeds were decreased slightly by 12 mo of 10°C storage (Fig. 1B and E). Electrolyte leakage increased and seedling dry weight decreased for sh-2 seeds stored at 10°C (Fig. 2C) , whereas germination percentage and seedling dry weight of nonprimed seeds stored at -80°C for 12 mo were unchanged (Fig. 1C and 2C).

View larger version (36K): [in this window] [in a new window]   Fig. 1. Germination, mean germination time (MGT) and seed moisture level of sh-2 sweet corn seed primed under different temperatures and nonprimed control stored at 25, 10, and -80°C for up to 12 mo. Vertical bars represent the mean and SE of three replications. LSD (P < 0.05) for germination, MGT and seed moisture level, across all the treatments, are 5.99, 0.45, and 0.04 respectively.

View larger version (42K): [in this window] [in a new window]   Fig. 2. Seedling dry weight, seed leakage and soluble protein content of sh-2 sweet corn seed primed under different temperatures and nonprimed control stored at 25, 10, and -80°C for up to 12 mo. Vertical bars represent the mean and SE of three replications. LSD (P < 0.05) for seedling dry weight, seed leakage, and soluble protein content, across all the treatments, are 1.85, 1.23, and 0.05, respectively.

Lipid Peroxidation and Protein Browning
Unstored control seeds had measurable levels of MDA, total peroxides and browning products (Fig. 3A) . The seed stored at 25°C for 6 to 12 mo accumulated more MDA, total peroxide, and browning products (Fig. 3A, D and G). Nonprimed seed stored at 10 and -80°C for 6, 9, and 12 mo accumulated less MDA, total peroxides, and browning products than those stored at 25°C (Fig. 3).

View larger version (42K): [in this window] [in a new window]   Fig. 3. Malondialdehyde, total peroxide and browning intensity of sh-2 sweet corn seed primed under different temperatures and nonprimed control stored at 25, 10, and -80°C for up to 12 mo. Vertical bars represent the mean and SE of three replications. LSD (P < 0.05) for malondialdehyde, total peroxide, and browning intensity, across all the treatments, are 0.05, 2.56, and 0.05, respectively.

Antioxidative Systems
The activities of antioxidative enzymes in both 25 and 10°C-stored nonprimed seeds decreased during 12 mo storage (Fig. 4 and 5) . Under -80°C condition, only SOD, APX, and GR showed a decrease in enzyme activity during 12 mo storage. Primed seed generally exhibited greater activities of antioxidative enzyme (except dehydroascorbate reductase, DHAR) than nonprimed control seed before storage. Under 25 and 10°C storage conditions, the activities of antioxidative enzymes decreased with increasing storage duration, with the rates of decline for 10 and 15°C-primed seed lower than for 20°C-primed seed. At -80°C storage, only slight decreases in SOD, APX, and GR activities occurred with increasing storage duration for primed seeds (Fig. 4 and 5). Marked decreases in soluble protein content occurred for the nonprimed and primed seed during storage (Fig. 2); the decreases were less with 10 and -80°C storage (Fig. 2H and I) compared with 25°C-stored seeds (Fig. 2G).

View larger version (44K): [in this window] [in a new window]   Fig. 4. Activities of superoxide dismutase, catalase, and ascorbate peroxidase of sh-2 sweet corn seed primed under different temperatures and nonprimed control stored at 25, 10, and -80°C for up to 12 mo. Vertical bars represent the mean and SE of three replications. LSD (P < 0.05) for superoxide dismutase, catalase, and ascorbate peroxidase activities, across all the treatments, are 0.39, 1.98, and 3.54, respectively.

View larger version (31K): [in this window] [in a new window]   Fig. 5. Activities of glutathione reductase and dehydroascorbate reductase of sh-2 sweet corn seed primed under different temperatures and nonprimed control stored at 25, 10, and -80°C for up to 12 mo. Vertical bars represent the mean and SE of three replications. LSD (P < 0.05) for glutathione reductase and dehydroascorbate reductase, across all the treatments, are 7.32 and 0.61, respectively.

View larger version (28K): [in this window] [in a new window]   Fig. 6. Reduced glutathione (GSH) and oxidized glutathione (GSSG) of sh-2 sweet corn seed primed under different temperatures and nonprimed control stored at 25, 10, and -80°C for up to 12 mo. Vertical bars represent the mean and SE of three replications. LSD (P < 0.05) for GSH and GSSG, across all the treatments, are 16.57 and 0.93, respectively.

View larger version (28K): [in this window] [in a new window]   Fig. 7. Ascorbate (ASC) and dehydroascorbate (DHA) of sh-2 sweet corn seed primed under different temperatures and nonprimed control stored at 25, 10, and -80°C for up to 12 mo. Vertical bars represent the mean and SE of three replications. LSD (P < 0.05) for ASC and DHA, across all the treatments, are 2.53 and 1.33, respectively.

	DISCUSSION

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Many environmental factors are known to influence the success of priming, priming temperature being critical (McDonald, 1999). In this study, we found a rapid decline in viability for 20°C-primed sh-2 seed during 25°C storage. Rates of germination decline were less pronounced when the seed was primed at 15 or 10°C (Fig. 1). Reduced electrolyte leakage and improved seedling growth confirmed the superior quality of 10 and 15°C-primed seed (Fig. 2). In addition, both MDA and total peroxides were lower in 15 and 10°C-primed seed than in 20°C-primed seed (Fig. 3), suggesting that lipid peroxidation also was decreaseing. Enhanced antioxidative activities are probable reasons for the reduced lipid peroxidation in 10 and 15°C-primed seed (Fig. 4 and 5). The higher levels of soluble and lower levels of browning observed in 10 and 15°C-primed seed compared with 20°C-primed seed support this argument (Fig. 2). Thus, decreased lipid peroxidation, under the enhanced protection by antioxidative mechanisms, might extend the longevity of 10 and 15°C-primed seed stored at 25°C.

Decreased longevity of 20°C-primed seed stored at 25°C might also result from a combination of hydration and temperature effects. The final water gain for 20°C primed-seed, at the end of hydration, was higher than 10 and 15°C-primed seed (Chiu, 2000). Under higher seed moisture conditions, 20°C-primed seed might pass the threshold of desiccation tolerance (i.e., irresistible cell division), thus injuring membranes in the seed beyond the point of recovery, even though they dehydrated faster than 10 or 15°C-hydrated seeds during redrying period (Chiu, 2000).

During storage, all seed undergoes deterioration, with the rate dependent on storage temperature and crop species. Successful long-term storage of primed seed generally requires drying the seeds below 100 g kg^-1 moisture and storing them at low temperature conditions (e.g., 10°C) (Dearman et al., 1986, 1987). Chang and Sung (1998) reported that storage at low temperature (<10°C) was required to maintain quality of primed sh-2 seeds, provided that the seed moisture was lower than 90 g kg^-1. In the present study, solid-matrix primed seed with 100 g kg^-1 moisture level stored at 10°C had much higher viability, vigor, and antioxidative activity than nonprimed seed (Fig. 1). In contrast, storage at 25°C for 12 mo reduced germination percentage and seedling dry weight from solid-matrix primed seed (Fig. 2). Thus, to maintain high quality in solid matrix primed sh-2 seed for extended storage period, the seed moisture should be kept at the level below 100 g kg^-1 and the seed should be stored at cool temperature (10°C). Primed sh-2 seed may also be stored satisfactorily at -80°C conditions for at least 12 mo. This would appear not to be commercially practical, however, except for germplasm conservation of inherently short-lived species.

In conclusion, our results confirm that 3 mo storage at 25°C of 20°C-primed sh-2 sweet corn seed reduced seed quality and inhibited antioxidative activity. The quality of 10 or 15°C-primed seed could be maintained for 6 mo even when they were stored at 25°C. However, 10 or -80°C storage extended the storability of solid-matrix primed sh-2 seeds to 12 mo. The improved antioxidative activity seems to play a role in maintaining the viability and vigor of solid-matrix primed seed stored at low temperature.

	ACKNOWLEDGMENTS

 
This work was supported by the National Science Council of ROC (NSC87-2313-B-005-056).

Received for publication July 31, 2001.

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This Article Abstract Figures Only Full Text (PDF) Free Alert me when this article is cited Alert me if a correction is posted Services Similar articles in this journal Similar articles in Web of Science Alert me to new issues of the journal Download to citation manager Citing Articles Citing Articles via HighWire Citing Articles via Web of Science (18) Citing Articles via Google Scholar Google Scholar Articles by Chiu, K. Y. Articles by Sung, J. M. Search for Related Content PubMed Articles by Chiu, K. Y. Articles by Sung, J. M. Agricola Articles by Chiu, K. Y. Articles by Sung, J. M.