Seed storage at elevated partial pressure of oxygen, a fast method for analysing seed ageing under dry conditions

Background and Aims

Despite differences in physiology between dry and relative moist seeds, seed ageing tests most often use a temperature and seed moisture level that are higher than during dry storage used in commercial practice and gene banks. This study aimed to test whether seed ageing under dry conditions can be accelerated by storing under high-pressure oxygen.

Methods

Dry barley (Hordeum vulgare), cabbage (Brassica oleracea), lettuce (Lactuca sativa) and soybean (Glycine max) seeds were stored between 2 and 7 weeks in steel tanks under 18 MPa partial pressure of oxygen. Storage under high-pressure nitrogen gas or under ambient air pressure served as controls. The method was compared with storage at 45 °C after equilibration at 85 % relative humidity and long-term storage at the laboratory bench. Germination behaviour, seedling morphology and tocopherol levels were assessed.

Key Results

The ageing of the dry seeds was indeed accelerated by storing under high-pressure oxygen. The morphological ageing symptoms of the stored seeds resembled those observed after ageing under long-term dry storage conditions. Barley appeared more tolerant of this storage treatment compared with lettuce and soybean. Less-mature harvested cabbage seeds were more sensitive, as was the case for primed compared with non-primed lettuce seeds. Under high-pressure oxygen storage the tocopherol levels of dry seeds decreased, in a linear way with the decline in seed germination, but remained unchanged in seeds deteriorated during storage at 45 °C after equilibration at 85 % RH.

Conclusions

Seed storage under high-pressure oxygen offers a novel and relatively fast method to study the physiology and biochemistry of seed ageing at different seed moisture levels and temperatures, including those that are representative of the dry storage conditions as used in gene banks and commercial practice.     

Innovative Techniques for Large‐scale Seagrass Restoration Using Zostera marina (eelgrass) Seeds

The use of Zostera marina (eelgrass) seeds for seagrass restoration is increasingly recognized as an alternative to transplanting shoots as losses of seagrass habitat generate interest in large‐scale restoration. We explored new techniques for efficient large‐scale restoration of Z. marina using seeds by addressing the factors limiting seed collection, processing, survival, and distribution. We tested an existing mechanical harvesting system for expanding the scale of seed collections, and developed and evaluated two new experimental systems. A seeding technique using buoys holding reproductive shoots at restoration sites to eliminate seed storage was tested along with new techniques for reducing seed‐processing labor. A series of experiments evaluated storage conditions that maintain viability of seeds during summer storage for fall planting. Finally, a new mechanical seed‐planting technique appropriate for large scales was developed and tested. Mechanical harvesting was an effective approach for collecting seeds, and impacts on donor beds were low. Deploying seed‐bearing shoots in buoys produced fewer seedlings and required more effort than isolating, storing, and hand‐broadcasting seeds in the fall. We show that viable seeds can be separated from grass wrack based on seed fall velocity and that seed survival during storage can be high (92–95% survival over 3 months). Mechanical seed‐planting did not enhance seedling establishment at our sites, but may be a useful tool for evaluating restoration sites. Our work demonstrates the potential for expanding the scale of seed‐based Z. marina restoration but the limiting factor remains the low rate of initial seedling establishment from broadcast seeds.     

Comparing methods of weed seed exposure to predators

Several methods of seed exposure are used in seed predation studies, but how these methods influence the results remains poorly studied. In this article, two commonly used methods of seed exposure – seed cards and plasticine trays – were compared in the field and in the laboratory using three species of weed seeds. In the field, cards or trays with seeds were exposed either with or without cages to keep vertebrates out and either with or without impermeable roofs to provide protection from the weather. The overall seed retrieval from the control stands varied significantly between the methods of seed exposure, roof treatment and the species of seeds, and the scatter in the seed retrieval increased or decreased monotonically with the temperature or precipitation for some species of seeds. This indicates that the controls were more or less relevant depending on the weather conditions and species. The seed removal from cards varied between the species of seeds. The effect of exclosure cages indicated that invertebrates were the dominant seed predators of Capsella bursa‐pastoris, while in Poa annua and Stellaria media both vertebrates and invertebrates were important. Higher seed removal from plasticine trays compared to seed cards was found for all three species of seeds, and placing roofs over the seeds affected seed removal in C. bursa‐pastoris and P. annua. In the laboratory, seed consumption varied with the method of exposure in 10 out of 12 combinations of seed species and predator (two species of carabids and two species of isopods). The overall tendency was the opposite of the field observations: seed consumption was higher for seeds on filter paper and cards compared to seeds on tin trays. We conclude that seed cards are more convenient for short term studies in the field, while tin trays are useful in laboratory multi‐choice experiments. To measure the realistic consumption of invertebrate predators in the laboratory, filter paper seems to be the best option as it does not present an obstacle to predators eating the seed. Using roofs introduced further bias regarding the estimates of seed removal in the field and thus should be avoided.     

The relative contributions of seed and pollen dispersal to gene flow and genetic diversity in seedlings of a tropical palm

Seed and pollen dispersal shape patterns of gene flow and genetic diversity in plants. Pollen is generally thought to travel longer distances than seeds, but seeds determine the ultimate location of gametes. Resolving how interactions between these two dispersal processes shape microevolutionary processes is a long‐standing research priority. We unambiguously isolated the separate and combined contributions of these two dispersal processes in seedlings of the animal‐dispersed palm Oenocarpus bataua to address two questions. First, what is the spatial extent of pollen versus seed movement in a system characterized by long‐distance seed dispersal? Second, how does seed dispersal mediate seedling genetic diversity? Despite evidence of frequent long‐distance seed dispersal, we found that pollen moves much further than seeds. Nonetheless, seed dispersal ultimately mediates genetic diversity and fine‐scale spatial genetic structure. Compared to undispersed seedlings, seedlings dispersed by vertebrates were characterized by higher female gametic and diploid seedling diversity and weaker fine‐scale spatial genetic structure for female gametes, male gametes and diploid seedlings. Interestingly, the diversity of maternal seed sources at seed deposition sites (N _em) was associated with higher effective number of pollen sources (N _ep), higher effective number of parents (N _e) and weaker spatial genetic structure, whereas seed dispersal distance had little impact on these or other parameters we measured. These findings highlight the importance maternal seed source diversity (N _em) at frugivore seed deposition sites in driving emergent patterns of fine‐scale genetic diversity and structure.     

Interactions between a seed‐eating neotropical rodent,the Azara's agouti (Dasyprocta azarae), and the Brazilian ‘pine’ Araucaria angustifolia

We studied the seed predation and scatter‐hoarding behaviour of Azara's agoutis Dasyprocta azarae (Rodentia: Dasyproctidae) in relation to the seeds of the Brazilian ‘pine’, Araucaria angustifolia (Araucariaceae), the rodent's main winter food source. We compared seed‐removal rates, seed‐caching rates, cache distances and recovery rates between a summer period of food abundance (with a low demand for A. angustifolia seeds and no such seeds naturally available) and a winter period of food scarcity (with a high demand for A. angustifolia seeds). We investigated whether the relative seed value affected the rodent's seed‐handling behaviour. We predicted that during the high seed‐demand period (winter): (1) cache distances would be greater; (2) fewer seeds would be stored; (3) more seeds would be recovered and the seed‐recovery time would be lower. In support of our first two predictions, the caching distances were greater in winter (mean ± SE = 15.67 ± 5.11 m) than in summer (9.40 ± 1.59 m), and agoutis hoarded >9 times more seeds in summer (55) than in winter (6). Our third prediction was not supported, and the proportion of unrecovered caches and buried seed recovery times did not differ between winter (mean ± SE = 3.00 ± 0.00 days, n = 5 seeds) and summer (11.05 ± 3.68 days, n = 20 seeds). The high resource density (during summer) rather than the density of A. angustifolia seeds likely influenced seed fate. Agoutis acted mainly as predators, leaving few intact seeds, caching a low proportion of handled seeds (? 8%) and rapidly consuming the caches. Agoutis may cache seeds to keep them safe from competitors on a short‐term basis rather than maintaining medium‐ or long‐term reserves for use during food‐scarcity periods.     

Seed dynamics of resprouting shrubs in grassy woodlands: Seed rain,predators and seed loss constrain recruitment potential

Abstract Measuring the fate of seeds between seed production and seedling establishment is critical in understanding mechanisms of recruitment limitation of plants. We examined seed fates to better understand the recruitment dynamics of four resprouting shrubs from two families (Fabaceae and Epacridaceae) in temperate grassy woodlands. We tested whether: (i) pre‐dispersal seed predation affected seed rain; (ii) post‐dispersal seed predation limited seed bank accumulation; (iii) the size of the seed bank was related to seed size; and (iv) viable seeds accumulated in the soil after seed rain. There was a distinct difference in seed production per plant between plant families with the legumes producing significantly more seeds per individual than the epacrids. Seed viability ranged from 43% to 81% and all viable had seed or fruit coat dormancy broken by heat or scarification. Pre‐dispersal predation by Lepidopteran larvae removed a large proportion of seed from the legume seed rain but not the epacrids. Four species of ants (Notoncus ectatomoides, Pheidole sp., Rhytidoponera tasmaniensis and Iridomyrmex purpureus) were major post‐dispersal seed removers. Overall, a greater percentage of Hardenbergia (38%) and Pultenaea (59%) seeds were removed than the fleshy fruits of Lissanthe (14%) or Melichrus (0%). Seed bank sizes were small (<15 seeds m^?2) relative to the seed rain and no significant accumulation of seed in the soil was detected. Lack of accumulation was attributed to seed predation as seed decay was considered unlikely and no seed germination was observed in our study sites. Our study suggests that seed predation is a key factor contributing to seed‐limited recruitment in grassy woodland shrubs by reducing the number of seeds stored in the soil.     

Antisense suppression of LOX3 gene expression in rice endosperm enhances seed longevity

Lipid peroxidation plays a major role in seed longevity and viability. In rice grains, lipid peroxidation is catalyzed by the enzyme lipoxygenase 3 (LOX3). Previous reports showed that grain from the rice variety DawDam in which the LOX3 gene was deleted had less stale flavour after grain storage than normal rice. The molecular mechanism by which LOX3 expression is regulated during endosperm development remains unclear. In this study, we expressed a LOX3 antisense construct in transgenic rice (Oryza sativa L.) plants to down‐regulate LOX3 expression in rice endosperm. The transgenic plants exhibited a marked decrease in LOX mRNA levels, normal phenotypes and a normal life cycle. We showed that LOX3 activity and its ability to produce 9‐hydroperoxyoctadecadienoic acid (9‐HPOD) from linoleic acid were significantly lower in transgenic seeds than in wild‐type seeds by measuring the ultraviolet absorption of 9‐HPOD at 234 nm and by high‐performance liquid chromatography. The suppression of LOX3 expression in rice endosperm increased grain storability. The germination rate of TS‐91 (antisense LOX3 transgenic line) was much higher than the WT (29% higher after artificial ageing for 21 days, and 40% higher after natural ageing for 12 months). To our knowledge, this is the first report to demonstrate that decreased LOX3 expression can preserve rice grain quality during storage with no impact on grain yield, suggesting potential applications in agricultural production.     

植冠与土壤种子库储存种子的萌发特性及策略

为明确黄土丘陵沟壑区植物种子库如何调控种子萌发来提高个体适合度,选择研究区7种具有种子库的主要物种为研究对象,以刚成熟和室内储存种子为对照,比较植冠宿存(5个宿存期)和土壤埋藏(5a埋藏期)对植物种子萌发特性的影响,探讨植冠种子库与土壤种子库储存下的种子萌发策略。结果表明:7种植物种子经过不同种子库储存后萌发特性表现出明显的种间差异,黄刺玫(Rosa xanthina)和水栒子(Cotoneaster multiflorus)种子萌发力表现为植冠宿存不变型、土壤储存增强型,土壤储存明显提高水栒子种子萌发速率;达乌里胡枝子(Lespedeza davurica)和狼牙刺(Sophora davidii)种子萌发力表现为植冠宿存增强型、土壤储存减弱型,种子萌发历时表现为植冠宿存延长型,土壤种子库储存还可加快达乌里胡枝子萌发速率、缩短萌发历时;茭蒿(Artemisia giralaii)和铁杆蒿(Artemisia gmelinii),种子萌发率随植冠宿存时间先升高后降低,随土壤储存时间先降低后升高,土壤储存可推迟其萌发,铁杆蒿种子萌发速率在植冠与土壤储存后均加快;紫丁香(Syringa oblata)种子萌发率随植冠宿存先升高后降低,土壤储存明显加快其种子启动萌发与速率。在黄土丘陵沟壑区,植物种子经过植冠或土壤种子库储存,或增加、加快、提早萌发充分利用有利条件提高占据性,或减少、减缓、推迟萌发分摊不利条件的风险;而且该区植物植冠与土壤储存后种子萌发特性间的关系,体现各自适应环境与应对干扰的分工与协作策略,主要表现为:单一主导型和相辅相成型。     

The ecophysiology of seed persistence: a mechanistic view of the journey to germination or demise

Seed persistence is the survival of seeds in the environment once they have reached maturity. Seed persistence allows a species, population or genotype to survive long after the death of parent plants, thus distributing genetic diversity through time. The ability to predict seed persistence accurately is critical to inform long‐term weed management and flora rehabilitation programs, as well as to allow a greater understanding of plant community dynamics. Indeed, each of the 420000 seed‐bearing plant species has a unique set of seed characteristics that determine its propensity to develop a persistent soil seed bank. The duration of seed persistence varies among species and populations, and depends on the physical and physiological characteristics of seeds and how they are affected by the biotic and abiotic environment. An integrated understanding of the ecophysiological mechanisms of seed persistence is essential if we are to improve our ability to predict how long seeds can survive in soils, both now and under future climatic conditions. In this review we present an holistic overview of the seed, species, climate, soil, and other site factors that contribute mechanistically to seed persistence, incorporating physiological, biochemical and ecological perspectives. We focus on current knowledge of the seed and species traits that influence seed longevity under ex situ controlled storage conditions, and explore how this inherent longevity is moderated by changeable biotic and abiotic conditions in situ, both before and after seeds are dispersed. We argue that the persistence of a given seed population in any environment depends on its resistance to exiting the seed bank via germination or death, and on its exposure to environmental conditions that are conducive to those fates. By synthesising knowledge of how the environment affects seeds to determine when and how they leave the soil seed bank into a resistance–exposure model, we provide a new framework for developing experimental and modelling approaches to predict how long seeds will persist in a range of environments.     

Seed traits and bird species influence the dispersal parameters of wetland plants

相似文献   

Heteromorphic seeds of wheat wild relatives show germination niche differentiation

• Crop wild relatives are fundamental genetic resources for crop improvement. Wheat wild relatives often produce heteromorphic seeds that differ in morphological and physiological traits. Several Aegilops and Triticum species possess, within the same spikelet, a dimorphic seed pair, with one seed being larger than the other. A comprehensive analysis is needed to understand which traits are involved in seed dimorphism and if these aspects of variation in dimorphic pairs are functionally related.
• To this end, dispersal units of Triticum urartu and five Aegilops species were X‐rayed and the different seed morphs weighed. Germination tests were carried out on seeds, both dehulled and left in their dispersal units. Controlled ageing tests were performed to detect differences in seed longevity among seed morphs, and the antioxidant profile was assessed in terms of antioxidant compounds equipment and expression of selected antioxidant genes. We used PCA to group seed morphs sharing similar patterns of germination traits, longevity estimates and antioxidant profile.
• Different seed morphs differed significantly in terms of mass, final germination, germination timing, longevity estimates and antioxidant profile in most of the tested species. Small seeds germinated slower, had lower germination when left in their dispersal units, a higher antioxidant potential and were longer‐lived than large seeds. The antioxidant gene expression varied between morphs, with different patterns across species but not clearly reflecting the phenotypic observations.
• The results highlight different trait trade‐offs in dimorphic seeds of Aegilops and T. urartu, affecting their germination phenology and longevity, thereby resulting in recruitment niche differentiation.

     

Effect of water content and temperature on seed longevity of seven Brassicaceae species after 5 years of storage

Maximising seed longevity is crucial for genetic resource preservation and longevity of orthodox seeds is determined by environmental conditions (water content and temperature). The effect of water content (down to 0.01 g·H₂O·g^?1) on seed viability was studied at different temperatures for a 5‐year storage period in taxonomically related species. Seeds of seven Brassicaceae species (Brassica repanda, Eruca vesicaria, Malcolmia littorea, Moricandia arvensis, Rorippa nasturtium‐aquaticum, Sinapis alba, Sisymbrium runcinatum) were stored at 48 environments comprising a combination of eight water contents, from 0.21 to 0.01 g·H₂O·g^?1 DW and six temperatures (45, 35, 20, 5, ?25, ?170 °C). Survival curves were modelled and P50 calculated for those conditions where germination was reduced over the 5‐year assay period. Critical water content for storage of seeds of six species at 45 °C ranged from 0.02 to 0.03 g·H₂O·g^?1. The effect of extreme desiccation at 45 °C showed variability among species: three species showed damaging effects of drying below the critical water content, while for three species it was neither detrimental nor beneficial to seed longevity. Lipid content could be related to longevity, depending on the storage conditions. A variable seed longevity response to water content among taxonomically related species was found. The relative position of some of the species as long‐ or short‐lived at 45 °C varied depending on the humidity at which storage behaviour was evaluated. Therefore, predictions of survival under desiccated conditions based on results obtained at high humidity might be problematic for some species.     

Mammal Abundances and Seed Traits Control the Seed Dispersal and Predation Roles of Terrestrial Mammals in a Costa Rican Forest

In Neotropical forests, mammals act as seed dispersers and predators. To prevent seed predation and promote dispersal, seeds exhibit physical or chemical defenses. Collared peccaries (Pecari tajacu) cannot eat some hard seeds, but can digest chemically defended seeds. Central American agoutis (Dasyprocta punctata) gnaw through hard‐walled seeds, but cannot consume chemically defended seeds. The objectives of this study were to determine relative peccary and agouti abundances within a lowland forest in Costa Rica and to assess how these two mammals affect the survival of large seeds that have no defenses (Iriartea deltoidea, Socratea exorrhiza), physical defenses (Astrocaryum alatum, Dipteryx panamensis), or chemical defenses (Mucuna holtonii) against seed predators. Mammal abundances were determined over 3 yrs from open‐access motion‐detecting camera trap photos. Using semi‐permeable mammal exclosures and thread‐marked seeds, predation and dispersal by mammals for each seed species were quantified. Abundances of peccaries were up to six times higher than those of agoutis over 3 yrs, but neither peccary nor agouti abundances differed across years. Seeds of A. alatum were predominantly dispersed by peccaries, which did not eat A. alatum seeds, whereas non‐defended and chemically defended seeds suffered high levels of predation, mostly by peccaries. Agoutis did not eat M. holtonii seeds. Peccaries and agoutis did not differ in the distances they dispersed seeds. This study shows that seed fates are contingent upon many factors such as seed defenses, frugivore–granivore abundances, and seed‐handling capabilities. Mammal–seed interactions are complex; the outcomes of these interactions depend on the inherent characteristics of seeds and their potential dispersers.     

Dispersal mode constrains allocation of carbon and mineral nutrients in seeds of forest and savanna trees

• Species vary in seed size and content of stored reserves, which can be related to dispersal strategies and type of habitat in which they are found. We compare seed carbon and nutrient reserves of anemochorous and zoochorous trees from the Cerrado of central Brazil.
• We measured seed dry mass, lipids, non‐structural carbohydrates (starch and total soluble sugars), carbon and mineral nutrients in ten forest and 13 savanna species, each classified as having wind‐ or animal‐dispersed seeds. We used phylogenetically independent contrasts to test for correlations among these traits.
• Seeds of anemochorous species were lighter, with higher concentrations of C, N, P, Ca and Mg. Lipids were the dominant carbon reserve for most anemochorous species, underpinning the importance of allocation to compact carbon reserves. Starch, lipids or soluble sugars were the major carbon reserve in zoochorous seeds. Savanna and forest species did not differ in seed mass or in total carbon reserves. However, seeds of forest species had higher concentrations of starch than seeds of savanna species. Lipid and starch negatively correlated across species, suggesting a trade‐off between starch and lipids as major seed carbon reserves. Calcium was positively correlated with Mn and B, while Mg was positively correlated with C, N, P, K, S, Zn and B. Potassium, S and Cl were positively correlated, while P was positively correlated with Mg and Zn.
• Dispersal mode rather than vegetation type constrained seed mass and seed storage allocation patterns in forest and savanna trees. We provide evidence that similar mechanisms are involved in seed storage of carbon and mineral nutrients across species.

     

An Effective Method for Collecting and Storing Seeds from Zostera marina (Eelgrass) in the Yellow Sea,China

The coast of the Yellow Sea in China, like many other temperate coastal zones, has been experiencing a dramatic decline in the abundance of seagrass. Intensive efforts have been made to restore seagrass communities along the coast to restore the function of the coastal ecosystem. Transplanting adult Zostera marina shoots is labor‐intensive, time‐consuming, expensive, and detrimental to donor beds; thus, restoring seagrass communities through the use of seeds is highly valued in current, large‐scale restoration trials. In this study, an effective method for collecting, processing, and storing Z. marina seeds was developed. From 2009 to 2013, respectively, 122,000, 421,000, 364,000, 1,041,000, and 1,091,000 seeds were successfully collected. Two‐way analysis of variance (ANOVA) showed the interaction between salinity and temperature significantly affected the cumulative germination rate (CGR) (p < 0.01) during the storage period and the viability (p < 0.01) of seeds after storage. The germination rate after storage was significantly affected by salinity and temperature (p < 0.01). The highest viability (89.8 ± 1.0%) and germination rate (75.6 ± 4.5%) were found among seeds stored at 4°C and a salinity of 44.5 psu for 7 months. The cost for planting 1 ha of sea bottom with Z. marina seeds ranged from $2,613 to $80,900 depending on the seeding density and seed loss during storage. The average cost per Z. marina seed in this study was $0.00586.     

Enhanced seed viability and lipid compositional changes during natural ageing by suppressing phospholipase Dα in soybean seed

Changes in phospholipid composition and consequent loss of membrane integrity are correlated with loss of seed viability. Furthermore, phospholipid compositional changes affect the composition of the triacylglycerols (TAG), i.e. the storage lipids. Phospholipase D (PLD) catalyses the hydrolysis of phospholipids to phosphatidic acid, and PLDα is an abundant PLD isoform. Although wild‐type (WT) seeds stored for 33 months were non‐viable, 30%–50% of PLDα‐knockdown (PLD‐KD) soybean seeds stored for 33 months germinated. WT and PLD‐KD seeds increased in lysophospholipid levels and in TAG fatty acid unsaturation during ageing, but the levels of lysophospholipids increased more in WT than in PLD‐KD seeds. The loss of viability of WT seeds was correlated with alterations in ultrastructure, including detachment of the plasma membrane from the cell wall complex and disorganization of oil bodies. The data demonstrate that, during natural ageing, PLDα affects the soybean phospholipid profile and the TAG profile. Suppression of PLD activity in soybean seed has potential for improving seed quality during long‐term storage.     

Artificial ageing of Vicia faba L. seeds caused by prolongation of G-1 phase

Broad bean (Vicia faba L. “Inovec”) seeds were artificially aged by means of storage at 30 %, resp. 25 % water content at 25 °C for 7-days to study the consequences on germination, root length and frequency of chromosomal aberrations. Under these conditions, significant changes in all parameters were observed. An increase of frequency of chromosomal aberrations in ana-telophase cells was confirmed by evaluation of c-metaphase cells. Synergic effect of artificial seed ageing was studied on different harvests of old seeds. Possible principles of this effect on cell level are discussed.     

Deterioration of western redcedar (Thuja plicata Donn ex D. Don) seeds: protein oxidation and in vivo NMR monitoring of storage oils

Deterioration of conifer seeds during prolonged storage hasa negative impact on reforestation and gene conservation efforts.Western redcedar (Thuja plicata Donn ex D. Don) is a speciesof tremendous value to the forest industry. The seeds of thisspecies are particularly prone to viability losses during long-termstorage. Reliable tools to assess losses in seed viability duringstorage and their underlying causes, as well as the developmentof methods to prevent storage-related deterioration of seedsare needed by the forest industry. In this work, various imagingmethods and biochemical analyses were applied to study deteriorationof western redcedar seeds. Seedlots that exhibited poor germinationperformance, i.e. those that had experienced the greatest lossesof viability during prolonged storage, exhibited greater abundanceof oxidized proteins, detected by protein oxidation assays,and more pronounced changes in their in vivo ¹³C NMR spectra,most likely due to storage oil oxidation. The proportion ofoxidized proteins also increased when seeds were subjected toaccelerated ageing treatments. Detection of oxidized oils andproteins may constitute a reliable and useful tool for the forestindustry. Key words: Conifer seeds, in vivo NMR spectroscopy, MRI, oil peroxidation, protein carbonylation, seed deterioration, seed storage, storage lipids, western redcedar Received 23 May 2007; Revised 28 November 2007 Accepted 17 December 2007