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Review

Seed Storability in Forest Trees: Research Progress and Future Perspectives

by
Hao Cai
,
Jun Shao
and
Yongbao Shen
*
Collaborative Innovation Centre of Sustainable Forestry in Southern China, College of Forestry, Nanjing Forestry University, Nanjing 210037, China
*
Author to whom correspondence should be addressed.
Forests 2025, 16(3), 467; https://doi.org/10.3390/f16030467
Submission received: 26 January 2025 / Revised: 26 February 2025 / Accepted: 4 March 2025 / Published: 6 March 2025
(This article belongs to the Section Genetics and Molecular Biology)
Browse Figures
Figure 1
<p>Key factors influencing seed storability. This figure was summarized and modified according to Zhou et al. [<a href="#B7-forests-16-00467" class="html-bibr">7</a>] and Choudhary et al. [<a href="#B33-forests-16-00467" class="html-bibr">33</a>].</p> "> Figure 2
<p>Major reactions that generate and eliminate ROS occur during seed aging under humid conditions (RH above 60%). I, NADH dehydrogenase; II, succinate dehydrogenase; III, cytochrome bc<sub>1</sub> complex; VI, cytochrome c oxidase; V, ATP synthase. The figure visualization was created with <a href="http://BioRender.com" target="_blank">BioRender.com</a>.</p> "> Figure 3
<p>Lipid peroxidation during seed aging under humid conditions (RH above 60%). The figure visualization was created with <a href="http://BioRender.com" target="_blank">BioRender.com</a>.</p> "> Figure 4
<p>Conceptual diagram of molecular regulatory mechanisms to enhance seed storability. The figure visualization was created with <a href="http://BioRender.com" target="_blank">BioRender.com</a>.</p> "> Figure 5
<p>Endogenous hormonal pathways regulating seed storability.</p> "> Figure 6
<p>Past practices and future innovations in enhancing seed longevity and storability.</p> ">
Versions Notes

Abstract

:
The long-term storage of forest tree seeds holds critical significance for ecological restoration, forest resource conservation, and the sustainable development of forestry. In the context of plant biodiversity conservation, enhancing seed storability to achieve efficient utilization has garnered widespread attention. Seed storability, as a complex quantitative trait, is influenced by the combined effects of intrinsic seed characteristics and external environmental factors. The complexity of this issue presents significant challenges in maintaining seed longevity, particularly in the conservation of seeds from endangered species. This review discusses the essential factors affecting seed storability and the main causes of seed aging. It emphasizes the roles of molecular mechanisms, including raffinose family oligosaccharide (RFO), heat shock protein (HSP), late embryogenesis abundant (LEA) proteins, seed storage proteins (SSPs), and hormonal regulation, in modulating seed storability. Additionally, the evaluation criteria and methodologies for assessing seed storability are elaborated. The review highlights future research challenges, aiming to provide a comprehensive scientific foundation and practical guidance to improve seed storability. This will offer theoretical support for the sustainable management of forest resources.

1. Introduction

With the rapid growth of the global population and the overexploitation of natural resources by human activities, forest resources worldwide are under unprecedented pressure. Although reports indicate that over 80,000 plant species (more than 20%) are threatened with extinction [1], the issue of biodiversity loss remains easily overlooked. This not only threatens the stability of global ecosystems but also poses potential risks to the future supply of food and medicinal resources for humanity [2]. To address this critical challenge, the scientific community has undertaken various biodiversity conservation measures. Among these, seed storage has been widely recognized as an effective means of preserving plant genetic resources [3]. Currently, nearly 1750 seed banks exist globally, and they house about 6 million accessions. However, an analysis of seed bank data indicates that less than 10% of taxa in seed bank collections are globally threatened [4]. Therefore, in the context of plant biodiversity conservation, the scientific management of stored seeds to ensure their viability and germination capacity has become a hotspot in biological research.
Seeds, as the key carriers of plant reproduction, play a critical role in the conservation of forest genetic resources and the sustainable management of forestry [5]. Seed storability, also known as seed longevity or resistance to aging, refers to the ability of seeds to retain their viability and germination capacity during long-term storage [6]. It has been reported that seed storability varies significantly across species, primarily influenced by intrinsic and extrinsic factors [7]. Roberts (1973) classified seeds into orthodox and recalcitrant types based on their storage behavior [8]. Generally, orthodox seeds exhibit strong storage tolerance under low-temperature and low-humidity conditions, while recalcitrant seeds tend to age and lose viability more rapidly during storage [9]. However, this does not imply that orthodox seeds are inherently easy to store. In the case of limber pine (Pinus flexilis James), the timing of seed harvest is difficult to optimize [10]. If limber pine seeds are harvested before they acquire desiccation tolerance, they may lose seed viability even under low-temperature and low-humidity storage conditions. Additionally, researchers have studied the dual-fruit tree species Amphicarpaea edgeworthii Benth., which produces both aerial and subterranean seeds on the same plant. The aerial seeds are orthodox, whereas the subterranean seeds are sensitive to desiccation and quickly lose viability during dry storage [11]. This complexity presents challenges for maintaining seed longevity, especially in conserving endangered species. Therefore, understanding the factors influencing seed storability is critical for optimizing storage techniques and providing a theoretical foundation for forest breeding and cultivation.
Seed storability can be evaluated using natural aging (NA) methods, whereby seeds are preserved under conventional conditions, often at around 30% relative humidity (RH), and their germination rates are monitored over time to obtain accurate assessments [12]. However, waiting for seeds to age naturally under NA conditions is clearly impractical. To address this limitation, researchers employ experimental treatments to accelerate the aging process [13]. A common approach is to store seeds under conditions of high humidity or elevated temperature [14]. When seeds are stored at high temperatures with the ambient RH maintained at 100%, the method is known as accelerated aging (AA) [15]. Alternatively, if the seed moisture content (MC) is equilibrated (typically from 60 to 85% RH by adjusting the ambient humidity) prior to high-temperature storage, the method is referred to as the controlled deterioration test (CDT) [15]. With the appropriate modifications [16], both AA and the CDT are widely used to assess seed storage performance [17,18]. Since the physiological and biochemical properties of seeds undergo significant changes with variations in moisture content and temperature, AA/CD tests cannot accurately reflect the deterioration mechanisms under dry, low-temperature, and NA conditions [19,20,21]. To more precisely study seed storability, researchers have proposed the elevated partial pressure of oxygen (EPPO) method [13]. During EPPO storage, the absolute amount of O2 is increased by raising the pressure (up to 20 MPa), while the relative amount of O2 is the same as ambient air [22]. Notably, compared to the commonly used AA and CDT methods at higher moisture content, the EPPO method more realistically simulates the patterns of seed longevity.
It is well established that seed aging is an unavoidable process. Over time, seed viability gradually declines, accompanied by reduced germination rates and seedling vigor [23]. This process is not only reflected in changes to the seed’s physical structures but also involves physiological alterations, such as cellular metabolic activity, reactive oxygen species (ROS) accumulation, DNA damage, and loss of membrane integrity [24]. Notably, under long-term storage conditions, the degradation of stored substances such as lipids and proteins within seeds triggers oxidative stress, thereby accelerating seed aging [25]. Despite these challenges, seeds often contain a variety of protective compounds, including non-reducing sugars (sucrose and oligosaccharides), late embryogenesis abundant (LEA) proteins, and heat shock proteins (HSPs), which play active roles in delaying seed aging [26,27,28]. Advances in molecular biology have deepened scientists’ understanding of the molecular regulatory mechanisms underlying seed storability. Recent studies have revealed that seed storability is regulated by dormancy-associated genes, such as DOG1, ABI3, and GA3ox, whose high expression can significantly prolong seed longevity [29,30].
Although substantial progress has been made in understanding seed storability, it remains a complex trait involving intricate physiological, genetic, and molecular mechanisms. Therefore, exploring essential factors affecting forest tree seed storability, physiological changes during seed aging, and molecular regulatory mechanisms to improve seed storability is vital for enhancing seed storage characteristics, conserving biodiversity, and achieving sustainable development. By integrating and updating existing research findings, this review aims to enhance seed storability, thereby facilitating the conservation and development of forest germplasm resources to meet the evolving demands of forestry.

2. Key Factors Influencing Seed Storability

As previously discussed, the key factors influencing seed storability can be classified into two main categories (Figure 1). The first category is intrinsic to the seed itself. Studies have shown that genetic traits, seed coat structure, and internal components play crucial roles in determining seed storability [31]. For instance, variations in lipid content or the retention of chlorophyll within seeds can result in significant differences in seed storability [28,32]. The second category pertains to storage conditions, including RH, temperature, O2 concentration, microbial activity, and pest infestation [33]. The same seed variety may exhibit markedly different seed longevity under varying storage environments. Among these, storage temperature and humidity are particularly influential, with lower temperatures and reduced RH levels generally promoting extended seed longevity [34]. Recent research has also highlighted how changes in O2 concentration, microbial activity, and pest infestation can significantly impact seed survival rates [7]. Overall, seed storability is shaped by the intricate interplay of intrinsic and extrinsic factors. A deeper understanding and optimization of these factors can contribute to enhancing the seed storability and quality of stored seeds.

2.1. Intrinsic Factors of Seeds

Seed storability is a complex trait regulated by multiple genes. Studies have shown that due to genomic differences, seed longevity varies greatly among species. Some species inherently possess seeds with long seed longevity that can remain viable under adverse storage conditions, while others lose viability more quickly. For example, the seeds of the sacred lotus (Nelumbo nucifera Gaertn.) have demonstrated an extraordinary ability to germinate, even after 1300 years of storage [35]. In contrast, the seeds of species such as willow (Salix spp.) and elm (Ulmus pumila L.) showed much shorter seed longevity [36,37]. This variation can be attributed to differences in the biochemical parameters among species, such as phenolic compounds, flavonoids, and vitamin E, which are positively correlated with seed storability [24]. For instance, Arabidopsis thaliana L. mutants deficient in vitamin E lose viability faster than wild-type seeds under seed AA conditions (40 °C, RH 100%) [38].
However, genetic characteristics are not the sole determinants of seed storability. Seed coat structure also plays a critical role [39]. For seeds with hard seed coats, a complete seed coat acts as a natural physical and chemical barrier, protecting the internal structures from environmental factors such as pest infestation, microbial activity, and physical damage [6]. When the seed coat is damaged, atmospheric moisture and O2 can penetrate the seed, accelerating deterioration and reducing storage longevity. Additionally, inner seed coats rich in antioxidants such as anthocyanins, flavonoids, and vitamins can effectively eliminate ROS and mitigate oxidative stress, contributing to prolonged seed longevity [40]. Therefore, this type of seed is often stored with its protective outer layers intact. Seed maturity is also a critical factor influencing seed storability, particularly during the late stages of seed maturation [41]. At this stage, seeds undergo significant biochemical changes, including the degradation of chlorophyll and the accumulation of LEA proteins, which are essential for acquiring desiccation tolerance and longevity [30,42]. Recent findings indicate that seeds with deep dormancy exhibit higher viability after long-term storage, highlighting a significant positive correlation between seed dormancy and storability [43]. Furthermore, even within the same species, seeds grown in different geographical environments may exhibit varying levels of storability. For example, recent studies have found that seeds from tropical regions generally have longer longevity than those from temperate areas [44].

2.2. Extrinsic Environmental Factors

The external environment also significantly influences seed storability. Seeds from the same batch may exhibit different longevity under varying storage conditions. Among the environmental factors, RH and storage temperature are the most critical. RH directly affects seed MC, as water vapor in the surrounding environment tends to equilibrate with the seed’s MC. High RH leads to increased seed MC, which accelerates respiration, elevates the risk of microbial activity and pest infections, and shortens seed longevity [33]. To ensure effective long-term storage, researchers typically maintain RH below 60% and seed MC below 14% [45]. Storage temperature is another crucial factor. Harrington’s rule of thumb suggests that a 5 °C increase in storage temperature halves seed longevity [46]. Elevated storage temperatures accelerate the chemical reactions within seeds, particularly the oxidation reaction, leading to thermal-induced protein degradation and faster deterioration of storage substances, ultimately reducing seed longevity. Additionally, unstable storage temperatures exacerbate these reactions and cause RH fluctuations, further accelerating seed degradation [6]. When storage temperatures reach critical thresholds, the proteins and DNA within seeds may denature, and cell membrane permeability increases, resulting in loss of viability [47]. Therefore, stable storage temperatures and minimal RH fluctuations are essential in practical applications. The composition of storage atmosphere also impacts seed storability. High O2 concentrations should accelerate respiration and oxidation, reducing seed viability. In contrast, gases such as CO2, N2, and other inert gases have protective effects [13]. Additionally, microorganisms and pests, which can directly damage seed embryos or accelerate respiration, are critical threats to seed longevity.

3. Mechanisms of Seed Aging

After the seeds have reached maturity, they inevitably enter the aging and deterioration phase [48]. This process involves a series of complex physiological, biochemical, and metabolic regulatory mechanisms, and our current understanding of these mechanisms remains incomplete. To effectively preserve genetic resources and mitigate the significant economic losses caused by seed aging in forestry production, it is necessary to explore the mechanisms of seed aging and the series of important events that occur during the aging process in order to understand the underlying causes of seed aging. By summarizing previous research findings, we attribute the main causes of seed aging to the following five factors: ROS imbalance, lipid peroxidation, genetic material damage, protein damage, and cellular structural damage. Understanding the physiological changes during seed aging can help in the scientific development of strategies to protect seed vitality. This has a positive impact on enhancing seed resistance to aging and extending seed longevity.

3.1. ROS Imbalance

Seed aging largely follows the ‘free radical theory’, which suggests that the damage caused by the accumulation of free radicals is a potential mechanism for biological aging. During seed aging and deterioration, ROS are considered the main driving force [49]. ROS include reactive oxygen species such as superoxide anion (O2−), hydrogen peroxide (H2O2), and hydroxyl radicals (OH·), which are typically highly reactive [50]. During seed aging, although ROS are continuously produced, their levels are generally maintained below a certain threshold. This is largely due to a series of protective mechanisms present in seeds during storage, which effectively limit the generation and accumulation of ROS [51]. The protective mechanism at work depends on the water status of the seed [52]. Under humid conditions (RH above 60%), the antioxidant system in seeds works synergistically through enzymatic pathways (such as superoxide dismutase [SOD] and catalase [CAT]) and non-enzymatic pathways (such as ascorbic acid [AsA] and glutathione [GSH]) to scavenge ROS and maintain their dynamic balance (Figure 2) [53]. However, under dry conditions (RH below 30%), the cytoplasm of seeds enters a glassy state, significantly reducing molecular mobility, and the enzymatic antioxidant pathways are almost completely inhibited. At this point, the scavenging of ROS primarily relies on the slow action of non-enzymatic antioxidants (such as vitamin E, GSH and AsA) [54]. However, when seeds are stored under unsuitable conditions (i.e., high temperature, increased RH) or for extended periods, the balance between ROS production and removal is disrupted. This ROS imbalance leads to oxidative damage to macromolecules such as lipids, proteins, and DNA, particularly lipid peroxidation, which damages cell membrane structures, increases membrane permeability, causes cytoplasmic leakage, and impairs cell function [55]. During artificial seed aging (45 °C, RH 95%), it has been reported that the O2− content in oat (Avena sativa L.) seeds significantly increases, while the activities of CAT and SOD decrease significantly [56]. Additionally, when observing the storage effects of black poplar (Populus nigra L.) seeds under different storage temperatures, it was found that, at 3 °C, the levels of O2− and H2O2 increased during seed storage, which led to membrane damage [57]. The researcher posits that the aging of black poplar seeds stored at 3 °C is attributable to the fact that, although enzyme activity increases continuously during storage, the overall enzyme levels remain insufficient, resulting in a significant rise in redox potential, even under dry and low-temperature storage conditions. Clearly, this explanation warrants further scrutiny. The increase in enzyme activity may be due to the release of enzymes from damaged tissues triggered by oxidative stress resulting from ROS accumulation, and these released enzymes are likely not involved in scavenging ROS. Moreover, the release of these enzymes during storage may reduce the availability of reducing agents during seed imbibition, thereby impairing the seed’s ability to repair oxidative damage. This imbalance between ROS production and elimination ultimately leads to seed germination failure. In addition, excess ROS can also affect the function of mitochondria and other organelles, reducing energy metabolism efficiency and making it difficult for seeds to acquire sufficient energy during germination [58]. It is noteworthy that while ROS accelerate seed aging, their presence is not entirely harmful to seeds. Studies have shown that during the early stages of germination, ROS act as important signaling molecules that coordinate the synthesis and distribution of endogenous hormones, thus promoting rapid seed germination and enhancing seed vitality [59]. Importantly, while ROS generally need to be at low concentrations, they must be tightly regulated by the antioxidant system. When seeds age, an imbalance between ROS production and antioxidant clearance can lead to excessive ROS accumulation, causing oxidative stress, cell damage, and ultimately seed death.

3.2. Lipid Peroxidation

Membrane damage caused by lipid peroxidation is considered a common mechanism of seed deterioration [60]. During seed aging under dry/humid conditions, the increased concentration of ROS accelerates the breakdown of cell membranes, producing a variety of polyunsaturated fatty acids (PUFA) [61]. Under humid storage conditions (RH above 60%) or during the revival stage of dry seeds (when they absorb water), these PUFA can be further metabolized by essential enzymes such as phospholipase (PLA) and lipoxygenase (LOX) into more reactive fatty acids (ROO), including lipid hydroperoxides (LOOH) (Figure 3) [24]. It is noteworthy that these LOOH can be further degraded into several cytotoxic byproducts, such as 4-hydroxy-2-nonenal (4-HNE) and malondialdehyde (MDA), which disrupt the normal function of cellular organelles [62]. Among these compounds, MDA is the most well-known marker of lipid peroxidation. It has been reported that MDA affects protein oxidation, leading to protein modification, aggregation, and conformational changes, ultimately altering protein function [63]. When Lallemantia iberica Fisch. and C.A. Mey seeds were stored under adverse storage conditions (45 °C, 15% MC), the MDA concentration increased by 15% compared to the beginning of storage, suggesting that membrane damage caused by lipid peroxidation may be related to the deterioration of Lallemantia iberica seeds [64]. Similar findings have been observed in the aging studies of Metasequoia glyptostroboides Hu and W.C. Cheng, soybeans (Glycine ma (L.) Merr.), and European beech (Fagus sylvatica L.) seeds [65,66,67]. Additionally, excessive lipid peroxidation generates free radicals that cause membrane disruption and damage biomacromolecules, further accelerating seed deterioration [55]. Recent seed humid storage studies indicate that volatile compounds (aldehydes or ketones) generated during lipid peroxidation may lead to microbial infections in seeds and provide nutrients for pests, exacerbating seed damage [7].

3.3. Damage to Genetic Material

During the long-term storage of seeds, an increased frequency of genetic mutations occurs in aging seeds due to accelerated nucleic acid degradation and the reduced synthesis inhibition [68]. ROS, such as O2− and H2O2, interact with DNA molecules, leading to base oxidation, DNA double-strand breaks (DSBs), and other forms of DNA damage [69,70]. DSBs are considered the most severe type of nucleic acid damage, causing the loss of large chromosomal regions and limiting seed germination. In addition, ROS can react with DNA bases and the phosphate backbone, resulting in oxidative base modifications. One of the most common modifications is 8-oxo-7,8-dihydroguanosine (8-oxo-G), which involves hydroxylation at the C-8 position of guanine [71]. 8-oxo-G is a potentially mutagenic compound that can mis-pair with adenine (A) residues, leading to single-strand DNA breaks [72]. In fact, seeds possess robust damage repair systems that minimize disruptions to biomolecules and cellular structures. DSBs repair primarily occurs through homologous recombination and non-homologous end joining [73]. Studies have confirmed that DNA damaged during the early stages of seed aging can be repaired [74]. In addition to causing DNA damage, ROS accumulation may also trigger RNA degradation [75]. RNA’s single-stranded structure makes it more susceptible to oxidation than DNA, particularly functional RNA molecules such as mRNA and rRNA [76]. When mature, dry seeds stored under suboptimal conditions are placed in germination chambers, the long-lived mRNA within the seeds is translated into proteins immediately after imbibition [77]. The abundance and integrity of these proteins are positively correlated with seed storage longevity. However, due to the structural instability of RNA, it degrades rapidly during aging.

3.4. Protein Damage

Protein carbonylation is one of the primary causes of seed aging during dry/humid storage, significantly impacting the metabolic stability and functional integrity of seed cells [78]. During seed aging, the ROS can induce the irreversible oxidative modifications of proteins, resulting in a marked increase in carbonylation levels [79]. Studies have shown that ROS induces protein carbonylation through two major pathways as follows: the first involves direct metal-catalyzed oxidation, inserting carbonyl groups into the side chains of amino acids such as arginine, lysine, and proline; the second involves reactions between lipid peroxidation intermediates (MDA and 4-HNE) and proteins, leading to carbonyl modifications [80,81]. These modifications disrupt protein structure and function, forming a molecular basis for the decline in seed vigor. Accelerated seed aging experiments have further validated the critical role of carbonylation in seed aging. For instance, Arabidopsis seeds stored under high-temperature (40 °C) and high-humidity (RH 85%) conditions showed significantly increased protein carbonylation levels with aging time, correlating positively with ROS accumulation [21]. Similar results have been observed in rice (Oryza sativa L.), elm, and European beech seeds during NA [82,83,84]. The accumulation of carbonylated proteins directly impairs key cellular metabolic functions. For example, the catalytic activities of essential enzymes such as malate dehydrogenase in the tricarboxylic acid (TCA) cycle, phosphoglycerate mutase in glycolysis, and ATP synthase subunit α in ATP synthesis are significantly reduced [21]. Additionally, the aggregation of carbonylated proteins further hinders normal cellular metabolic pathways, exacerbating energy crises and leading to seed death [79,85,86]. Protein carbonylation is not confined to the cytosol; it is also widely distributed across subcellular organelles such as mitochondria and peroxisomes. Particularly in humid conditions, those subcellular organelles typically serve as the primary sites for ROS generation. Mitochondrial proteins in complexes I and III of the electron transport chain (ETC) are highly sensitive to carbonylation. Their oxidative modifications directly impede electron transport and accelerate the energy depletion of seed cells [82,87]. This metabolic collapse is closely associated with a reduction in seed storage longevity. However, during the germination of Arabidopsis seeds, many proteins undergo carbonylation modifications, yet these seeds still develop into vigorous seedlings [88]. This phenomenon suggests that protein carbonylation is not purely a harmful process. In fact, with prolonged seed storage, protein carbonylation has been positively correlated with increased ROS levels and has also been shown to alleviate seed dormancy in sunflower (Helianthus annuus L.) and Arabidopsis [89,90]. For example, mutations in the NADPH oxidase genes AtRbohB and AtRbohD in Arabidopsis significantly reduce protein carbonylation levels, leading to prolonged seed dormancy [91]. These findings suggest that there is a balance between the beneficial effects of protein carbonylation in breaking seed dormancy and its detrimental impact on seed storability. However, further studies are needed to elucidate how this balance is achieved.

3.5. Damage to Cellular Ultrastructure

Seed aging causes significant and irreversible changes to the cellular ultrastructure during dry/humid storage [92]. Studies have shown that aged seeds exhibit withered embryos, with plastids undergoing significant contraction and various organelles, such as the endoplasmic reticulum, mitochondria, and lysosomes, swelling [59]. Mitochondria, as the center of energy synthesis and metabolism in cells, are often the first organelles to be damaged during seed aging [79]. In non-aged seeds, mitochondria typically possess an intact double-membrane structure with well-developed cristae and are embedded in a dense cytoplasm [39]. However, as seeds age, the morphology and function of mitochondria undergo significant changes, including swelling, cristae fragmentation, and irregular inner membrane structures [93]. In an artificial aging study of elm seeds (the seeds were aged at 37 °C above water for 5 d), researchers found that mitochondria in non-aged seeds appeared as punctate structures uniformly distributed within the cytoplasm. As aging progressed, the mitochondria gradually lost their functionality and developed abnormal morphologies, such as worm-like, enlarged, or diffuse shapes. When the seeds completely lost viability, the mitochondria aggregated in a diffuse form near the cell membrane [39]. Besides mitochondria, other organelles are also affected by seed aging. The Golgi apparatus and endoplasmic reticulum may rupture or disappear, the thylakoid membranes of chloroplasts become irregular, and lipid bodies break down into larger or irregular lipid droplets [24]. These damages to cellular ultrastructure directly impair the metabolic functions, significantly accelerating the aging process of seeds.

4. Molecular Regulatory Mechanisms to Enhance Seed Storability

Seed aging is an unavoidable physiological change during long-term storage, with profound influence on germplasm conservation and forestry production. Therefore, understanding and regulating the genes and mechanisms associated with seed storability is critical for optimizing forest tree seed storage techniques. Studies have shown that seeds have evolved complex physiological and molecular mechanisms to mitigate the adverse effects of aging in response to changing environmental conditions [94]. As previously mentioned, orthodox seeds enhance their storability by actively reducing MC during maturation to improve desiccation tolerance. However, recalcitrant seeds are highly sensitive to desiccation and cannot withstand dehydration damage without losing viability, which significantly limits their storage potential [95]. Additionally, the dynamic changes in the composition and content of soluble sugars in seeds influence their physiological state and storage tolerance. For example, the accumulation of sucrose and RFO not only stabilizes cell membranes during seed dehydration but also enhances storability by preventing lipid peroxidation [27]. HSPs play an essential protective role under environmental stress during seed maturation. The expression of HSP genes, including those of the small HSP family, is induced during seed development. These proteins maintain physiological activity and stability by facilitating proper protein folding, preventing aggregation, and repairing damaged proteins, thus improving seed storability [96]. Similarly, LEA proteins, which accumulate during the late stages of embryogenesis, are essential factors in seeds’ responses to dehydration stress. LEA proteins enhance desiccation tolerance and extend seed longevity by stabilizing cellular structures and protecting macromolecules under extreme dehydration conditions [28]. SSP also plays a vital role in seed storability [97]. Concurrently, genes involved in hormone regulation are central in modulating seed storability. Abscisic acid (ABA), a crucial hormone in seed maturation and dormancy, promotes the expression of dormancy-related genes such as ABI3 and DOG1 through its signaling pathways, regulating dormancy depth and resistance to aging. These genes influence seeds’ stress resistance and survival capabilities by modulating ABA synthesis and signaling [59]. In summary, seed storability is governed by a complex network of genetic and physiological mechanisms. Genes related to HSP, LEA proteins, and hormone regulation synergistically contribute to the development and maintenance of seed storability (Figure 4). Understanding these mechanisms and their interactions will provide essential theoretical and practical guidance for developing more efficient forest tree seed storage technologies.

4.1. The Role of Sugar Metabolism in Enhancing Seed Storability

In plants, sugars serve as essential regulators of nearly all fundamental processes throughout the seed life cycle, including maturation, germination, and aging [98]. During seed maturation, the accumulation of soluble non-reducing sugars helps reduce MC loss, facilitating the transition of the cytoplasm from a liquid to a glassy state, which minimizes metabolic activity during seed storage [59]. High temperature and humidity hinder the cytoplasm’s transition to a glassy state, whereas low temperature and low moisture promote its formation [99]. During seed desiccation, oligosaccharides such as sucrose and RFO replace water in the cells, promoting the formation of the glassy state [100]. In this state, RFOs help prevent vesicle fusion, maintain the liquid crystalline phase of cell membranes, and effectively protect the membrane from water-induced damage [101]. Sucrose has been shown to preserve the integrity of cellular membranes, while RFOs enhance the protective effects of sucrose by reducing lipid crystallization [102]. The accumulation of RFO has been reported to significantly improve the storage longevity of seeds from the Brazilian native tree Erythrina speciosa Andrews. This is primarily because RFOs, as reserve energy sources, gradually accumulate during seed development and are redistributed from vacuoles to the cytosol during the later stages of seed maturation. This redistribution helps Erythrina speciosa seeds maintain the liquid crystalline phase of cell membranes during dehydration in seed maturation [103]. Similarly, in a study on the long-term seed storage of European beech seeds at −10 °C, researchers found that seeds with a low sucrose-to-RFO ratio typically exhibited higher germination ability. The accumulation of RFO members such as stachyose also contributed to maintaining seed viability during seed storage [104]. These findings support the notion that non-reducing sugars, particularly from the raffinose family, may fluctuate during seed storage. However, an increase in RFO content contributes to ROS and lipid peroxidation, thereby enhancing seed vigor and longevity. Galactinol synthase (GolS) is a crucial enzyme in RFO biosynthesis [105]. GolS significantly influences carbon allocation between sucrose and RFOs and facilitates RFO synthesis by converting myo-inositol and UDP-galactose into galactinol [26]. In black poplar, an endangered tree species, GDSL proteins enhance seed storage performance under low-temperature and desiccation conditions by regulating fatty acid metabolism and membrane stability [106]. Intriguingly, when researchers knocked out GolS genes (GolS1 and GolS2) in Arabidopsis, seeds of the GolS1 mutants and GolS1 GolS2 double mutants exhibited low concentrations of galactinol and a sharp decline in seed vigor [107]. This further highlights the close association between RFO metabolic genes and seed storability. Suppressing the expression of GolS1 and GolS2 in the endosperm negatively regulates seed longevity by reducing galactinol concentrations. Although these studies reveal the critical role of sugar metabolism in enhancing seed storability, most of the research has focused on model plants. Studies on the seed storability of forest tree species remain relatively limited, especially for endangered trees in the wild.

4.2. The Role of HSP in Enhancing Seed Storability

HSP, a class of molecular chaperones, plays a crucial role in plant responses to abiotic stress and seed storage processes [108]. By regulating the expression of downstream genes, HSP contributes to maintaining seed viability and enhancing storage tolerance through the following mechanisms: (i) promoting the proper folding of newly synthesized proteins; (ii) preventing oxidative damage to proteins; and (iii) repairing damaged proteins [59]. These functions make HSPs essential regulators of seed storability. It has been reported that HSP expression is typically regulated by heat shock factors (HSFs), among which HSFA9 is a key transcription factor (TF) [109]. In transgenic tobacco (Nicotiana tabacum L.) subjected to CDT (50 °C for 5 h), the overexpression of the sunflower heat shock factor HaHSFA9 significantly increased the accumulation of HSP, thereby extending seed longevity [110]. Although this study primarily simulated stress tolerance through heat treatment rather than directly evaluating seed storability, it highlighted the potential mechanism by which HSFA9 enhances seed stress resistance through the regulation of the HSP network. However, other studies have shown that HSFA9’s role in seed longevity under optimal storage conditions is limited. For instance, in Medicago truncatula Gaertn., HSFA9 mutant seeds exhibited similar longevity to wild-type seeds under optimal storage conditions but aged significantly faster under high-humidity or high-temperature environments [30]. This indicates that HSFA9 plays a critical role in regulating seed aging processes, particularly under adverse storage conditions. Additionally, HSFA9 regulates the expression of other related TFs, including HSP70, HSFB2A, HSFA2, and molecular chaperones such as ROF1 and BAG6. These proteins collectively contribute to antioxidant defense and protein stability, synergistically enhancing seed vigor and storage tolerance. OsHSP18.2 is another key HSP associated with seed longevity [111]. In Arabidopsis, during artificial aging experiments (45 °C, RH 100%), the overexpression of OsHSP18.2 reduces ROS accumulation, protecting other proteins from oxidative damage and significantly improving seed vigor and storability. The expression of this protein increases markedly after artificial aging, further underscoring its critical role in maintaining seed viability [111]. Moreover, the expression of HSP acts in coordination with other regulatory factors. ABI3, an important regulatory factor in the ABA signaling pathway, indirectly enhances HSP expression by activating the promoter of HSFA9, thereby improving seed storage tolerance [112]. In Chinese cork oak (Quercus variabilis Blume), ABI3 indirectly induces seed desiccation tolerance by regulating the expression of HSPA9, which increases HSP content, thereby improving seed storage longevity [113]. In summary, HSPs enhance seed storability and longevity through antioxidant defense, the maintenance of protein homeostasis, and collaboration with other chaperone proteins. These mechanisms highlight the significance of HSPs as key regulators in extending seed storage life.

4.3. The Role of LEA Proteins in Enhancing Seed Storability

During the late stages of seed maturation, seed cells undergo a significant dehydration process that activates a group of genes encoding hydrophilic proteins known as LEA proteins [28]. LEA proteins primarily accumulate during the late embryogenesis stage of seed maturation and combine with sugar molecules to form a glassy cytoplasm, which protects cellular components and maintains seed viability [114]. This protective mechanism is critical for seed desiccation tolerance and long-term storage. LEA proteins possess a unique intrinsically disordered structure that can transition into a stable α-helix under desiccation conditions [115]. This structural transformation forms a protective barrier that prevents irreversible damage to cell membranes, proteins, and other biological macromolecules [116]. In Arabidopsis, the overexpression of the LEA gene AtEM6 significantly enhances seed desiccation tolerance and viability, confirming the positive role of LEA proteins in seed longevity [117]. Moreover, LEA2 proteins, also known as dehydrins (DHNs), which belong to Group 2 LEA proteins, play a particularly prominent role in seed longevity regulation. Proteomic analyses indicate that DHN accumulation decreases progressively with an increase in artificial aging (40 °C, RH 85%) time, closely correlating with the loss of germination capacity in Arabidopsis seeds [21]. Researchers have validated this hypothesis through RNA interference (RNAi) targeting the LEA14 gene, showing that the decreased expression of LEA14 and its related genes, such as XERO and RAB18, significantly shortens seed longevity. This finding suggests that DHNs play an active role in extending seed storability [118]. BP65 (seed biotinylated protein of 65 kDa of apparent molecular mass), a Group 3 LEA protein identified in pea (Pisum sativum L.), predominantly accumulates during the transition of embryonic cells from seed maturation to dormancy, a phase where metabolic activity slows down [119,120]. This protein reportedly regulates intracellular biochemical reactions by binding free biotin, an essential cofactor of carboxylases. Since carboxylation reactions may trigger unnecessary energy consumption during metabolic activity in the embryo development stage, reduced SBP65 mRNA levels and biotinylated protein accumulation in the viviparous mutant (vip-1) of pea significantly compromise seed metabolic stability, desiccation tolerance, and longevity [120]. Additionally, other LEA proteins in Medicago truncatula have also been found to play critical roles in seed longevity regulation [121]. In Astronium fraxinifolium Schott, a threatened tree species from Brazil, genes encoding LEA proteins were among the most upregulated DEGs in seeds compared to aged seeds, highlighting their importance in seed lifespan regulation [122]. Although these studies have elucidated the crucial roles of LEA proteins in seed storability, their precise mechanisms remain to be further explored. Future research should focus on the functional diversity and interaction networks of LEA proteins in different plant species. This could pave the way for molecular breeding strategies targeting improved storage tolerance in forest tree seeds, providing novel methods and theoretical foundations for the conservation and utilization of forest genetic resources.

4.4. The Role of SSP in Enhancing Seed Storability

Seed storage proteins (SSPs) are critical nutritional and functional components in seeds [97]. These proteins are primarily categorized into three major classes: globulins, albumins, and prolamins. Among them, globulins are the most common SSPs, widely distributed in legumes and grasses [123]. For instance, the 11S and 7S globulins in soybean accumulate significantly during seed maturation and play an active role in maintaining cellular homeostasis and enhancing seed storability [28]. During the AA (30 °C, RH 75%) process of poplar seeds, the abundance of 11S globulin SSPs increases significantly in seeds with low vigor [124]. Under dry storage conditions, globulins exhibit critical oxidative regulatory functions, particularly the 12S globulin. In non-aged dry seeds, the α-subunit of 12S globulin undergoes preferential carbonylation, whereas the β-subunit is fully carbonylated after seed aging [21,84,88]. This selective carbonylation suggests that the high oxidative sensitivity of the α-subunit may promote protein mobilization during seed germination through the 20S proteasome-mediated protein degradation pathway, thereby enhancing seed vigor [125]. Additionally, mutations in the CRU3 gene in Arabidopsis, which encodes 12S globulin, lead to a significant reduction in seed vigor. This finding further underscores the critical role of SSPs in delaying seed aging and maintaining longevity [126]. Furthermore, due to their high abundance and affinity for oxidation, SSPs are considered effective ROS scavenging systems that protect the essential cellular components required for embryo survival [126]. During seed dry storage, this ROS-buffering function is closely associated with the molecular chaperone activity of SSP [59]. By mitigating oxidative damage induced by ROS, these proteins not only enhance the long-term storability of seeds but also contribute positively to seed germination and seedling growth [127]. In summary, SSPs are not only key nutritional reserves during seed development but also support long-term storability and germination capacity through multiple mechanisms. However, further research is needed to fully elucidate the specific molecular mechanisms of SSPs in seed aging and environmental adaptation.

4.5. The Role of Endogenous Hormones in Enhancing Seed Storability

Endogenous hormones play a crucial role in maintaining seed storability by regulating cellular metabolism, antioxidant capacity, and gene expression, thereby influencing the aging process of seeds (Figure 5). ABA is one of the key endogenous hormones, with various genes in the ABA signaling pathway significantly affecting seed storability. For example, in the woody plant Calocedrus macrolepis Kurz, which is an endangered species, the seed storability was improved by ABA accumulation [128]. In Arabidopsis, the ABA content in the CYP707A2 mutant is approximately six times higher than that in the wild type, which significantly extends seed longevity and vigor [129]. Additionally, the overexpression of the OsDET1 gene increases ABA accumulation, markedly enhancing seed dormancy and storage tolerance in rice seeds [130]. This suggests that enhancing seed dormancy can effectively improve storage tolerance. The DOG1 gene, a key regulator of ABA accumulation, plays a vital role in establishing seed dormancy during development. Arabidopsis seeds with DOG1 mutations exhibit significant declines in vigor and storability, further indicating that DOG1 may extend seed longevity by promoting dormancy [131]. However, the relationship between seed dormancy and longevity may be complex. An analysis of inbred populations revealed that the DOG1-Cape Verde Islands allele both shortens seed longevity and increases dormancy [132]. This suggests that, under natural conditions, dormancy and longevity may be regulated by independent mechanisms, which could vary between different germplasms.
It is widely recognized that chlorophyll retention in mature seeds is often considered a negative factor affecting their storability and is closely related to low storage tolerance [133]. Notably, ABA regulates chlorophyll degradation through its signaling pathway during seed maturation, ensuring high seed vigor and storage durability during storage [134]. For instance, during seed maturation, ABA promotes chlorophyll degradation by activating key genes ABI3 and ABA4, which ensures high seed vigor and storability in mature seeds [135]. Interestingly, during chlorophyll degradation, ABA also induces the expression of genes encoding chlorophyll b reductase, such as the NYC1 gene in Arabidopsis. Mutants of NYC1 show increased chlorophyll retention, leading to a rapid decline in seed vigor during storage [136]. The loss of ABI3 suppresses NYC1 expression, resulting in green seeds even after drying. This further confirms that ABA positively affects seed vigor and longevity by regulating chlorophyll degradation. Recent studies have revealed that transcription factors ABI4 and ABI5 in the ABA signaling pathway play negative regulatory roles in primary seed dormancy. In legume species such as Medicago truncatula, seeds of the Mtabi4 mutant and the double mutant Mtabi4/Mtabi5 exhibit chlorophyll retention and reduced longevity. Notably, Mtabi4 mutants developing in darkness show delayed chlorophyll breakdown and impaired photosystem II degradation [137]. This indicates that ABI4 not only facilitates chlorophyll degradation but also prolongs seed storability by suppressing photosynthesis. In studies on recalcitrant seeds of Chinese cork oak, the downregulation of ABI5 alters the expression of the downstream related genes, resulting in increased sensitivity of seeds to desiccation, which negatively impacts seed longevity [113].
Gibberellins (GAs), as members of the tetracyclic diterpenoid compound family, play critical roles in plant development and reproduction [134]. Previous studies have revealed that increased GA levels positively contribute to seed longevity during seed storage [138]. For example, in Arabidopsis, the simultaneous knockdown of the transcription factors ATHB25, ATHB22, and ATHB31 significantly shortens seed longevity. The core component of the GA signaling pathway, DELLA proteins, acts as a negative regulator of GA signaling and plays a crucial role in regulating seed dormancy and germination [139]. It has been found that the new tomato (Solanum lycopersicum L.) mutant seeds lacking DELLA activity exhibit reduced longevity during dry seed storage [140]. In Arabidopsis, seeds of DELLA mutants, characterized by sustained GA signaling activity, show enhanced tolerance to aging [141]. This further emphasized the positive role of GA in seed longevity. The central regulatory gene GA3ox, a pivotal component of GA metabolism, is essential for GA biosynthesis and promotes seed germination when highly expressed [142]. TF ATHB25 positively regulates GA3ox2, enhancing the synthesis of GA-related metabolites and thereby improving seed storability in Arabidopsis [141]. However, while GA regulation plays a vital role in breaking dormancy and initiating germination, its function in extending seed storability is complex and remains controversial. On one hand, GA promotes embryo growth and breaks dormancy, accelerating germination, but this characteristic may have adverse effects during prolonged seed storage [143]. This could be attributed to the overactivation of GA signaling, which accelerates seed aging. Moreover, GA may weaken the antioxidant system, leading to the accumulation of ROS and increased cellular damage. Therefore, excessive GA production or enhanced signaling under storage conditions could shorten seed longevity. In studying GA’s role in regulating seed longevity, the intricate signaling network involving GA and other hormones such as ABA must be considered. Current research indicates that GA and ABA jointly regulate seed longevity and dormancy through downstream genes such as DOG1, ABI3, and RGL2 [144,145,146]. The differential expression of these genes leads to diverse interactions between GA and ABA in various species. Finally, a study on Arabidopsis mutants deficient in GA synthesis and signaling suggests that GAs’ role in longevity remains inconclusive [135]. Therefore, the relationship between GA signaling and seed longevity warrants further investigation.
Scientists have cloned and studied numerous genes associated with seed dormancy, some of which show potential for enhancing seed storability. However, leveraging dormancy enhancement to improve seed storability faces many challenges in practical applications. Increased seed dormancy often leads to reduced germination rates, uneven seedling emergence, and poor seedling establishment. These issues not only reduce nursery production efficiency but may also delay forestry planting schedules. Particularly in ecological restoration projects, delays could result in significant economic losses. Therefore, identifying and utilizing genes that enhance seed storability without compromising germination is a critical goal in forest genetic improvement. Although dormancy-related genes demonstrate the potential to improve storability under laboratory conditions, their practical benefits remain unverified in production. Balancing the trade-off between enhanced dormancy and seed storability presents an urgent challenge. Future research should focus on identifying and functionally validating genes capable of significantly improving seed storability without affecting seed dormancy, providing practical solutions for the conservation and utilization of forest germplasm resources. Furthermore, advances in molecular biology enable scientists to precisely regulate the expression levels of key genes through gene editing and molecular marker technologies, facilitating the synergistic improvement of seed dormancy and storability. For instance, regulating dormancy-associated genes such as DOG1, ABI3, and GA3ox could simultaneously enhance seed longevity and seedling vigor. Future research directions should prioritize identifying genes that improve seed storability independently of seed dormancy enhancement, accompanied by extensive validation and functional optimization. This approach will help scientists better understand the relationship between seed dormancy and storability, bringing greater economic and ecological benefits to forestry production and environmental conservation.

5. Seed Storability Assessment Indicators and Methods

5.1. Traditional Indicators

As mentioned above, seed storability is typically assessed using NA or AA methods [147]. Consequently, it is widely used in both research and practical applications [13]. Notably, whether using NA or AA, traditional indicators for evaluating seed vigor typically include the germination rate, TTC (triphenyl tetrazolium chloride) staining rate, and relative electrical conductivity (Figure 6) [38]. Recent studies have also proposed using the time required for the germination rate to decline to 50% of its initial rate (P50) as a dynamic evaluation of seed vigor [148,149]. It is worth noting that during long-term storage, seed aging is often accompanied by damage to the biological membrane system, manifesting as the accumulation of large amounts of free fatty acids and ROS. These characteristics provide a basis for developing seed storability assessment methods. For instance, lipoxygenase activity, fatty acid metabolism analysis, and ROS-related enzyme activity assays have been widely applied to evaluate the degree of seed aging [150]. Moreover, studies have shown that changes in seed components—such as the physicochemical properties of fatty acid content—are more sensitive to variations in seed longevity than traditional vigor indicators [151]. It is noteworthy that, as seeds age, the gradual impairment of mitochondrial function drives a shift from aerobic respiration to anaerobic respiration, leading to the production of ethanol [152]. Consequently, the accumulation of ethanol has been widely used as a key indicator for assessing the degree of seed aging and seed quality.

5.2. Emerging Technologies

With the rapid advancement of molecular biology and omics technologies, tools such as metabolomics, proteomics, and transcriptomics have been widely applied to investigate seed quality and longevity (Figure 6). To date, numerous key factors influencing seed storability have been identified in various plant species (Table 1). For instance, a metabolomics analysis revealed significant differences in metabolite profiles among rice varieties under AA conditions (45 °C, RH 60%). Results showed that seeds with longer longevity had markedly higher levels of flavonoids (e.g., quercetin-3-arabinoside and kaempferol), amino acids (e.g., cysteine derivatives), and sugars (e.g., glucose) [153]. In recent studies, researchers using the EPPO method have discovered that the functional Rc gene can regulate the synthesis of proanthocyanidins and their accumulation in fruit peel, thereby enhancing resistance to oxidative damage [52]. These findings are consistent with the role of phenolic compounds, such as flavonoids, in enhancing seed longevity under dry storage conditions, as demonstrated by their ability to scavenge ROS and protect cellular structures. Therefore, these metabolites play critical roles in extending seed longevity and could serve as potential biomarkers for predicting seed storability. Similar observations have been validated in Sinojackia xylocarpa Hu seeds [154], further confirming that changes in metabolite levels are an important means of assessing seed storability. In addition, transcriptomics studies have successfully identified genes closely associated with seed vigor. For example, in maize (Zea mays L.), TFs such as ZmActdf, ZmUBQ, and ZmGAPDH were found to be significant biomarkers affecting seed vigor [155]. These genes play essential roles in seed aging and storability. It is noteworthy that protein damage accumulation during seed aging is one of the key factors affecting seed storability. The accumulation of isoaspartate (isoAsp) accelerates aging [156]. However, regulating genes that repair damaged proteins, such as the PIMT gene in rice, significantly improves seed vigor. Studies have demonstrated that the overexpression of OsPIMT1 in transgenic seeds reduced isoAsp content and enhanced embryo vigor [157]. This suggests that the PIMT gene has potential as an important marker for evaluating seed vigor. Moreover, tandem mass tag (TMT) labeling technology has been employed to analyze the effects of artificial aging (45 °C; 50% RH) on protein expression in wheat (Triticum aestivum L.) seeds. The results indicated that proteins such as oleate synthase, hemoglobin-1, non-specific lipid transfer proteins, and lectins were significantly correlated with seed viability [158]. These findings further suggest that differential protein expression could serve as markers for seed aging under artificial aging. Despite the significant progress in revealing the determinants of seed storability through omics tools, the application of genetic engineering to enhance the seed storability of forest trees remains underexplored.
Currently, artificial intelligence (AI) technologies provide new tools for seed storability research. Phenomics approaches based on imaging and sensors have been developed to efficiently evaluate seed quality and longevity [159]. For instance, these technologies have excelled in predicting seed longevity in Brassica napus L., leveraging high-throughput seed phenotyping to precisely monitor seed germination rates and seedling survival curves, thereby improving assessment efficiency and accuracy [160]. Additionally, non-destructive detection methods such as oxygen-sensing technology, near-infrared spectroscopy, hyperspectral imaging, chlorophyll fluorescence imaging, and X-ray micro-computed tomography (micro-CT, μCT) have also been widely applied for seed vigor evaluation [161,162,163,164]. However, the detection precision of these techniques needs further optimization to meet the demands of complex research and practical applications.
Table 1. Key factors influencing seed storability.
Table 1. Key factors influencing seed storability.
NameMethodsTechnologyMain FindingReferences
Rice
(Oryza sativa L.)		AAMetabolomics analysisSeeds with longer longevity had markedly higher levels of flavonoids (quercetin-3-arabinoside and kaempferol), amino acids (cysteine derivatives), and sugars (glucose).[153]
Pedunculate oak (Quercus robur L.)NAMetabolomics analysisThe significant increase in the abundance of phenolic compounds, amino acids, phosphorylated monosaccharides, and carbohydrates.[165]
Maize
(Zea mays L.)		AATranscriptomics
analysis		TFs (ZmActdf, ZmUBQ, and ZmGAPDH) play essential roles in seed aging and storability.[155]
Metasequoia glyptostroboides Hu and W.C.ChengAATranscriptomics
analysis		Forty hub genes, such as Rboh, ACO, HSF, and CML play important roles in the antioxidant network. [66]
Astronium fraxinifolium SchottAATranscriptomics
analysis		Differentially expressed genes from photosystems, GRPs, and ubiquitin-conjugating enzymes play roles in the response to seed aging.[122]
Wheat
(Triticum aestivum L.)		AAProteomics
analysis		Proteins such as oleate synthase, hemoglobin-1, non-specific lipid transfer proteins, and lectins were significantly correlated with seed viability.[158]
Cariniana legalis (Martius) O. KuntzeNAProteomics
analysis		The decrease in the accumulation of proteasome subunit α-type and phosphoglucose isomerase proteins in seeds may be associated with the loss of seed vigor.[166]
Black poplars
(Populus nigra L.)		NAProteomics
analysis		The excellent storability of seeds at subzero temperatures is related to factors such as the reduced abundance of proteins involved in metabolism, hydrolysis, and protein turnover.[106]
Beech
(Fagus sylvatic L.)		NAProteomics
analysis		Protein Sec61 and glyceraldehyde-3-phosphate dehydrogenase may be used as potential longevity modulators in beech seeds.[167]
Note: NA: natural aging; AA: accelerated aging.

6. Conclusions

Seed storability, which is the ability of seeds to maintain viability during prolonged storage, is a key research focus in forestry resource conservation and germplasm preservation. In the context of global climate change and the continuous decline of wild plant species, optimizing seed storage techniques not only contributes to biodiversity preservation but also plays a critical role in maintaining ecosystem stability. In recent years, scientists have made substantial progress in elucidating the complex mechanisms underlying seed storability through the integration of seed physiology and modern omics technologies. Additionally, efforts to develop improved seeds with extended longevity have achieved partial success. However, despite these breakthroughs, many gaps and challenges remain (Figure 6).
Molecular biology research has provided a new framework for understanding seed storability. As discussed earlier, researchers have identified several genes related to storage tolerance and elucidated their underlying molecular mechanisms, such as DOG1, ABI3, and GA3ox. However, these genes are predominantly associated with seed dormancy, which poses challenges for practical applications that rely on enhancing dormancy to improve storability. Balancing seed viability and storability while maintaining seed dormancy remains a significant focus and challenge for future research. The application of modern omics technologies, such as transcriptomics, proteomics, and metabolomics, has further revealed the regulatory networks involved in seed aging. As described above, metabolic markers such as protein carbonylation and lipid peroxidation have been shown to play essential roles in seed aging. Nevertheless, these molecular markers have only been extensively studied in model plants such as Arabidopsis and rice but remain largely unexplored in forestry tree species, particularly rare and endangered wild plants. Therefore, future research should prioritize the study of seed storability in forest tree species.
Research on seed storability requires a closer integration of fundamental science and practical applications. First, it is essential to further elucidate the molecular mechanisms of seed aging, identify key genes associated with seed storability, and utilize tools such as CRISPR/Cas9 gene-editing technology to precisely regulate the expression of these genes. Building on this foundation, the integration of multi-omics approaches can establish regulatory networks for seed storability, enabling the development of tailored storage strategies for different forest tree seed types. Second, technological advancements have provided valuable tools for seed storability research. High-throughput phenotyping combined with machine learning algorithms offers efficient ways to assess seed vigor by monitoring seed germination curves and seedling vigor dynamics. However, the scalability and practicality of these methods in real-world applications still require further optimization. Additionally future efforts should focus on developing more efficient and cost-effective seed storage technologies. Innovative approaches, such as nano-enabled seed treatment, the introduction of natural antioxidants, and protective coating, could significantly extend seed longevity. Particular attention should be given to endangered forest tree species to develop targeted seed storage techniques that align with the need to preserve genetic diversity and support global ecological conservation.
Currently, breeding programs for many economically important tree species primarily focus on traits such as growth rate, wood quality, and disease resistance while often neglecting seed storability. This oversight leads to a significant decline in the germination rates of high-quality or disease-resistant varieties after long-term storage, thereby reducing their economic and ecological value. By utilizing modern biotechnological tools, such as identifying and introducing storage-tolerant genes, we can significantly enhance the storability of these high-value genotypes while preserving their desirable economic traits. In addition, as mentioned earlier, the overexploitation of forest resources has led to a sharp decline in biodiversity, posing serious challenges to effective protection for these species. To address these challenges, optimizing storage conditions (low RH, low temperature, absence of light and vacuum sealing) and adopting novel seed treatment technologies (protective coatings and oxygen absorbers) are particularly crucial. These methods can effectively maintain the viability of seeds from endangered tree species during long-term storage. Furthermore, in-depth research into the interactions between environmental factors (such as temperature and humidity) and seed storability will provide important theoretical support for future innovations in seed storage technology. By integrating traditional breeding methods with modern biotechnology, this comprehensive approach can not only maximize the delay in seed aging but also enhance the adaptability of trees to ever-changing environmental challenges.
In conclusion, seed storability research plays a critical role in forestry production and resource conservation. With continued advancements in technologies and enhancements in theoretical understanding, this field will provide robust support for addressing climate change and maintaining global ecological balance. In the future, interdisciplinary research that integrates AI and biology may offer new possibilities for seed storability research, paving the way for sustainable forestry and ecological development.

Author Contributions

H.C.: Writing—original draft. J.S.: Methodology. Y.S.: Writing—review and editing. All authors have read and agreed to the published version of the manuscript.

Funding

This work is was supported by a long-term scientific research base for the in vitro conservation of ray native tree germplasm resources in Jiangsu Province, LYKJ[2021]03; and the Priority Academic Program Development of Jiangsu Higher Education Institutions (PAPD).

Data Availability Statement

No data were used for the research described in the article.

Conflicts of Interest

The authors declare that there are no conflicts of interest regarding the publication of this paper.

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Figure 1. Key factors influencing seed storability. This figure was summarized and modified according to Zhou et al. [7] and Choudhary et al. [33].
Figure 1. Key factors influencing seed storability. This figure was summarized and modified according to Zhou et al. [7] and Choudhary et al. [33].
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Figure 2. Major reactions that generate and eliminate ROS occur during seed aging under humid conditions (RH above 60%). I, NADH dehydrogenase; II, succinate dehydrogenase; III, cytochrome bc1 complex; VI, cytochrome c oxidase; V, ATP synthase. The figure visualization was created with BioRender.com.
Figure 2. Major reactions that generate and eliminate ROS occur during seed aging under humid conditions (RH above 60%). I, NADH dehydrogenase; II, succinate dehydrogenase; III, cytochrome bc1 complex; VI, cytochrome c oxidase; V, ATP synthase. The figure visualization was created with BioRender.com.
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Figure 3. Lipid peroxidation during seed aging under humid conditions (RH above 60%). The figure visualization was created with BioRender.com.
Figure 3. Lipid peroxidation during seed aging under humid conditions (RH above 60%). The figure visualization was created with BioRender.com.
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Figure 4. Conceptual diagram of molecular regulatory mechanisms to enhance seed storability. The figure visualization was created with BioRender.com.
Figure 4. Conceptual diagram of molecular regulatory mechanisms to enhance seed storability. The figure visualization was created with BioRender.com.
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Figure 5. Endogenous hormonal pathways regulating seed storability.
Figure 5. Endogenous hormonal pathways regulating seed storability.
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Figure 6. Past practices and future innovations in enhancing seed longevity and storability.
Figure 6. Past practices and future innovations in enhancing seed longevity and storability.
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Cai, H.; Shao, J.; Shen, Y. Seed Storability in Forest Trees: Research Progress and Future Perspectives. Forests 2025, 16, 467. https://doi.org/10.3390/f16030467

AMA Style

Cai H, Shao J, Shen Y. Seed Storability in Forest Trees: Research Progress and Future Perspectives. Forests. 2025; 16(3):467. https://doi.org/10.3390/f16030467

Chicago/Turabian Style

Cai, Hao, Jun Shao, and Yongbao Shen. 2025. "Seed Storability in Forest Trees: Research Progress and Future Perspectives" Forests 16, no. 3: 467. https://doi.org/10.3390/f16030467

APA Style

Cai, H., Shao, J., & Shen, Y. (2025). Seed Storability in Forest Trees: Research Progress and Future Perspectives. Forests, 16(3), 467. https://doi.org/10.3390/f16030467

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