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Main conclusion Discrete categories of seed physiology can be explained through a unified concept of the structural and molecular mobility responses within cells to drying. Tolerance of desiccation is typically described by a threshold or low water content limit to survival. This convention provides fairly good distinction between orthodox and recalcitrant seeds, which show thresholds of less than about 0.07 and greater than about 0.2 g H₂₋O g DW⁻¹, respectively. Threshold water contents, however, are not direct measures of the intensity of water stress tolerated by seeds, not are they measures of cell response to water stress. More direct criteria, that accommodate both spatial and temporal effects of water loss, are required to explain variation of desiccation tolerance and longevity in seeds from diverse genetic backgrounds and growth conditions. This essay presents the argument that changes in cellular volume directly quantify primary responses to desiccating stress in a context that also links damage, as cellular constituents compress, and protection, as compressed molecules form stabilizing structure. During desiccation, fluid cytoplasm solidifies, and the newly formed spatial relationships among molecules determine whether and how long viability is maintained. The diversity of seed behaviors suggests complexity and opportunity to discover molecules and mechanisms that regulate survival and perception of time in cells that lack metabolic function.

Cryogenic technologies help to preserve plant biodiversity in seed banks, particularly in the tropics. Concerns about the rapid erosion of plant diversity have spawned a host of seed-banking initiatives ( 1 ). These repositories provide critical germ plasm needed to understand, maintain, and manage natural variation within and among species ( 2 ). However, numerous plant species and much of the humid tropics are underserved in these endeavors because of the perceived problem of seed recalcitrance ( 3 ). About 75 to 80% of angiosperm species ( 4 , 5 ) produce orthodox seeds that can survive drying and prolonged storage at −20°C. By contrast, 5 to 10% of angiosperm species produce recalcitrant seeds that do not survive desiccation ( 3 ) and are killed in the freezer when ice crystals form. How can their preservation be ensured?

The cytoplasm of anhydrobiotes (organisms that persist in the absence of water) solidifies during drying. Despite this stabilization, anhydrobiotes vary in how long they persist while dry. In this paper, we call upon concepts currently used to explain stability of amorphous solids (also known as glasses) in synthetic polymers, foods, and pharmaceuticals to the understand variation in longevity of biological systems. We use embryonic axes of pea (Pisum sativum) and soybean (Glycine max) seeds as test systems that have relatively long and short survival times, respectively, but similar geometries and water sorption behaviors. We used dynamic mechanical analysis to gain insights on structural and mobility properties that relate to stability of other organic systems with controlled composition. Changes of elastic and loss moduli, and the dissipation function, tan δ, in response to moisture and temperature were compared in pea and soybean axes containing less than 0.2 g H2O g-1 dry mass. The work shows high complexity of structure-regulated molecular mobility within dried seed matrices. As was previously observed for pea cotyledons, there were multiple relaxations of structural constraints to molecular movement, which demonstrate substantial localized, 'fast' motions within solidified cytoplasm. There was detected variation in the coordination among long-range slow, diffusive and short-range fast, vibrational motions in glasses of pea compared to soybean, which suggest higher constraints to motion in pea and greater 'fragility' in soybean. We are suggesting that differences in fragility contribute to variation of seed longevity. Indeed, fragility and coordination of short and long range motions are linked to stability and physical aging of synthetic polymers.

The lifespan or longevity of a seed is the time period over which it can remain viable. Seed longevity is a complex trait and varies greatly between species and even seed lots of the same species. Our scientific understanding of seed longevity has advanced from anecdotal ‘Thumb Rules,’ to empirically based models, biophysical explanations for why those models sometimes work or fail, and to the profound realisation that seeds are the model of the underexplored realm of biology when water is so limited that the cytoplasm solidifies. The environmental variables of moisture and temperature are essential factors that define survival or death, as well as the timescale to measure lifespan. There is an increasing understanding of how these factors induce cytoplasmic solidification and affect glassy properties. Cytoplasmic solidification slows down, but does not stop, the chemical reactions involved in ageing. Continued degradation of proteins, lipids and nucleic acids damage cell constituents and reduce the seed’s metabolic capacity, eventually impairing the ability to germinate. This review captures the evolution of knowledge on seed longevity over the past five decades in relation to seed ageing mechanisms, technology development, including tools to predict seed storage behaviour and non-invasive techniques for seed longevity assessment. It is concluded that seed storage biology is a complex science covering seed physiology, biophysics, biochemistry and multi-omic technologies, and simultaneous knowledge advancement in these areas is necessary to improve seed storage efficacy for crops and wild species biodiversity conservation.

Aged soybean seeds show widespread cleavage of long mRNA transcripts with no particular relationship to transcript function, consistent with extant hypotheses of aging mechanisms. Abstract Seeds exist in the vulnerable state of being unable to repair the chemical degradation all organisms suffer, which slowly ages seeds and eventually results in death. Proposed seed aging mechanisms involve all classes of biological molecules, and degradation of total RNA has been detected contemporaneously with viability loss in dry-stored seeds. To identify changes specific to mRNA, we examined the soybean (Glycine max) seed transcriptome, using new, whole-molecule sequencing technology. We detected strong evidence of transcript fragmentation in 23-year-old, compared with 2-year-old, seeds. Transcripts were broken non-specifically, and greater fragmentation occurred in longer transcripts, consistent with the proposed mechanism of molecular fission by free radical attack at random bases. Seeds died despite high integrity of short transcripts, indicating that functions encoded by short transcripts are not sufficient to maintain viability. This study provides an approach to probe the asymptomatic phase of seed aging, namely by quantifying transcript degradation as a function of storage time.

Dry seeds do not show obvious signs of life and so testing for viability, health and life expectancy can be challenging. Usually testing seed quality involves adding water and measuring metabolic capacity or growth potential by vital staining or germination assays. Importantly, most laboratory seed tests are intended to assay immediate viability, while most genebanks need tests that predict seed performance in the distant future. All currently available assays require considerable a priori knowledge of germination conditions and seeds large enough to dissect. Germination conditions are often unknown for seeds produced from wild species and are an important criterion for seed testing. Using standardized methods (i.e., commercially available kits) we tested the feasibility of adapting a new seed quality assay that measures RNA integrity and is promising for cultivated species, to seeds from wild species. Most of the 100 wild species we include are rare or endangered and in need of preservation through genebanking. To determine the feasibility of measuring RNA integrity in seeds from wild populations, we compared the quality of RNA extracted from seeds that were recently harvested to those of the same species that have been genebanked for 16 to 41 years, with various seed traits examined for interference with RNA extraction and characterization. We demonstrate reliable characterization of RNA quality across a diverse group of plants, despite variation in germination requirements, seed morphology or composition. RIN (RNA Integrity Number) values were usually high across all samples and variables, attesting to both the quality of newly collected material as well as its maintenance under genebanking conditions. This study conclusively demonstrates the feasibility of reliably extracting and characterizing RNA from dry seeds collected from wild populations, regardless of a variety of seed traits and morphologies. Relationships between RNA quality and seed age and viability require further exploration.

Worldwide, oak species are threatened with extinction due to habitat loss, pathogens, and changing fire regimes. Ex situ conservation through tissue culture may protect the remaining genetic diversity of Quercus dumosa, or the coastal sage scrub oak, from further loss. We designed three basal salt formulations based on the mineral composition of shoot tips and first leaves from mature Q. dumosa and explored carbohydrate source, stress-mitigating compounds, and plant growth regulator concentrations to develop a method of cultivating many Q. dumosa culture lines in vitro. All three novel basal salt formulations led to decreased necrosis compared with commercial basal salt formulas WPM, MS, and DKW. Substitution of 30 g L−1 sucrose with glucose and adding 250 mg L−1 ascorbic acid, 5.2 mg L−1 SNP sodium nitroprusside, and 103 mg L−1 y-aminobutyric acid improved culture health overall. In an experiment involving 115 culture lines, 0.66 mg L−1 6-benzylaminopurine produced the highest average shoots per explant, but 0.33 mg L−1 produced the greatest proportion of shoots 2 cm or greater. Incubation for 24 h in 20 mg L−1 indole-3-butyric acid led to the most rooting. These methods show promise for the ex situ conservation of many genotypes of endangered Q. dumosa.

Seeds of different species are believed to have characteristic shelf lives, although data confirming this are scarce, and a mechanistic understanding of why this should be remains elusive. We have quantified storage performance of c. 42,000 seed accessions, representing 276 species, within the USDA National Plant Germplasm System (NPGS) collection, as well as a smaller experiment of 207 cultivars from 42 species. Accessions from the NPGS collection were harvested between 1934 and 1975, and had relatively high initial germination percentages that decreased at a variable rate during storage at both 5 and –18°C. Germination time courses, which represent the average performance of the species, were fitted to Avrami kinetics, to calculate the time at which germination characteristically declined to 50% (P50). These P50 values correlated with other longevity surveys reported in the literature for seeds stored under controlled conditions, but there was no correlation among these studies and seed persistence observed in the classic buried seed experiment by Duvel. Some plant families had characteristically short-lived (e.g. Apiaceae and Brassicaceae) or long-lived (e.g. Malvaceae and Chenopodiaceae) seeds. Also, seeds from species that originated from particular localities had characteristically short (e.g. Europe) or long (e.g. South Asia and Australia) shelf lives. However, there appeared to be no correlation between longevity and dry matter reserves, soluble carbohydrates and parameters relating to soil persistence or resource allocation. Although data from this survey support the hypothesis that some species tend to survive longer than others in a genebank environment, there is little information on the attributes of the seed that affect its storage performance.

Premise of the study: Fern spores are unicellular and haploid, making them a potential model system to study factors that regulate lifespan and mechanisms of aging. Aging rates of nongreen spores were measured to compare longevity characteristics among diverse fern species and test for orthodox response to storage temperature and moisture. Methods: Aging of spores from 10 fern species was quantified by changes in germination and growth parameters. Storage temperature ranged from ambient room to -196°C (liquid nitrogen); spores were dried to ambient relative humidity (RH) or using silica gel. Key results: Survival of spores varied under ambient storage conditions, with one species dying within a year and two species having greater than 50% survival after 3 years. Few changes in germination or growth were observed in spores stored at either -80°C or -196°C over the same 3-yr study period. Spores stored at -25°C aged anomalously quickly, especially those dried to ambient RH or subjected to repeated freeze-thaw cycles. Conclusions: Spore longevity is comparable to orthodox seed longevity under ambient storage conditions, with wide variation among species and shelflife extended by drying or cooling. However, faster aging during freezer storage may indicate a similar syndrome of damage experienced by seeds categorized as \"intermediate\". The damage is avoided by storage at -80°C or liquid nitrogen temperatures, making cryoconservation an effective and broadly applicable tool to extend fern spore longevity. The study demonstrates that spore banks are a feasible approach for ex situ conservation of this important plant group.

More than a billion people worldwide may be selenium (Se) deficient, and supplementation with Se-rich Brazil nuts may be a good strategy to prevent deficiency. Since different forms of Se have different nutritional value, and Se is toxic at elevated levels, careful seed characterization is important. Variation in Se concentration and correlations of this element with other nutrients were found in two batches of commercially available nuts. Selenium tissue localization and speciation were further determined. Mean Se levels were between 28 and 49 mg kg−1, with up to 8-fold seed-to-seed variation (n = 13) within batches. Brazil nut Se was mainly in organic form. While present throughout the seed, Se was most concentrated in a ring 1 to 2 mm below the surface. While healthy, Brazil nuts should be consumed in moderation. Consumption of one seed (5 g) from a high-Se area meets its recommended daily allowance; the recommended serving size of 30 g may exceed the allowable daily intake (400 μg) or even its toxicity threshold (1200 μg). Based on these findings, the recommended serving size may be re-evaluated, consumers should be warned not to exceed the serving size and the seed may be sold as part of mixed nuts, to avoid excess Se intake.