1 Introduction

Horticultural crops, including fruits, vegetables, herbs and ornamentals, play a crucial role in ensuring global food and nutritional security, in addition to supporting rural livelihoods and the agroeconomies of both developing and developed countries. These crops are particularly sensitive to environmental fluctuations, making them highly vulnerable to abiotic stressors such as drought, salinity, heat, cold and waterlogging (Borgohain et al., 2019; Pandey et al., 2019). Such stress conditions severely affect seed germination, seedling vigor, and metabolic functioning, ultimately compromising yield quality and quantity. This vulnerability is further exacerbated by the increasing frequency, severity, and unpredictability of these stressors under the influence of climate change (Prasad et al., 2017). Therefore, improving the resilience of horticultural crops to abiotic stresses is critical for achieving sustainable agricultural development and climate-resilient food systems (Fahad et al., 2017).

In recent years, seed priming has emerged as a pre-sowing strategy of high interest due to its ability to enhance stress tolerance without genetic modification or costly inputs. Seed priming refers to the controlled hydration of seeds to initiate pre-germinative metabolic processes, followed by re-drying before radicle protrusion (Paul et al., 2022). This controlled activation leads to faster and more uniform germination, improved seedling vigor and enhanced physiological preparedness to adverse environmental conditions (Singh et al., 2020).

The growing interest in seed priming has led to a surge of research focused on its application across a diverse range of horticultural crops and agroecological contexts. Numerous studies have evaluated the effectiveness of various priming agents, including osmotic solutions, phytohormones, nanomaterials, and microbial biostimulants, in enhancing plant performance under stress (Jisha et al., 2013; de Oliveira and Gomes-Filho, 2016). Osmopriming and hormonal priming, in particular, have shown broad efficacy in improving drought, salinity and cold tolerance (Ulfat et al., 2017; Lei et al., 2021), while nanopriming has demonstrated potential in enhancing antioxidant activity and maintaining membrane stability under heat stress conditions (Khan et al., 2023). Biopriming has also gained attention for its dual role in stress mitigation and the promotion of nutrient acquisition and plant defense mechanisms (Rajendra Prasad et al., 2016).

Despite these advancements, most studies have addressed single-stress conditions under controlled environments. There is a critical knowledge gap in understanding how seed priming may mediate cross-tolerance, the capacity of plants to tolerate multiple concurrent stresses after exposure to a single stimulus (Ramegowda et al., 2020). This is of particular importance given that climate-induced stressors often occur in combination (e.g., drought with high temperature or salinity), presenting more complex physiological challenges than individual stresses. Additionally, the lack of standardized protocols for priming treatments and the limited availability of multi-environment field trials hinders the broader application of these techniques in horticultural production systems.

Therefore, this review aims to comprehensively assess the current state of knowledge on seed priming in horticultural crops, with a special focus on its role in promoting multi-stress resilience. It synthesizes the physiological, biochemical and molecular mechanisms activated by different priming agents, evaluates the potential for cross-tolerance and highlights key research gaps, particularly regarding priming under combined stress scenarios and its scalability to field-level applications. Addressing these gaps is essential to optimize seed priming strategies for real-world use and to enhance the sustainability of horticultural production under climate variability.

2 Key abiotic Stresses and Their Impacts on Horticultural Crops

Horticultural crops are particularly sensitive to abiotic stress due to their delicate physiology, shallow root systems, and high-water content. The most common abiotic stressors include drought, salinity and extreme temperatures (both heat and cold). These factors disrupt physiological processes at multiple stages of development, from germination to postharvest, resulting in significant economic and agronomic losses and their impacts are often magnified when they occur in combination (Rao et al., 2016; Francini and Sebastiani, 2019).

2.1 Drought stress

Drought is one of the most widespread and damaging abiotic stresses affecting agricultural production worldwide (Borgohain et al., 2019). This challenge undermines the sustainability of agricultural systems and is further intensified by the rising global demand for food, driven by the steady growth of the world population (Seleiman et al., 2021). It induces osmotic stress, leading to stomatal closure and reduced leaf expansion, which collectively impair the plant's ability to perform gas exchange and transpiration (Salehi-Lisar and Bakhshayeshan-Agdam, 2016; Fahad et al., 2017). Consequently, photosynthetic rates decline due to alterations in chlorophyll content and disruptions in enzyme activity (Reddy et al., 2004).

Additionally, drought stress disrupts hormonal balance, particularly by increasing abscisic acid (ABA) synthesis, which further reinforces stomatal closure and inhibits cell elongation (Souza and Cardoso, 2003). In crops like tomato (Solanum lycopersicum L.) and pepper (Capsicum annuum L.), prolonged water stress is also associated with increased susceptibility to blossom-end rot due to reduced calcium mobility within the plant (Saure, 2014). At the metabolic level, drought stress often triggers the accumulation of osmoprotectants (e.g., proline, trehalose) and the upregulation of antioxidant enzymes like superoxide dismutase (SOD) and catalase (CAT) as defensive responses (Rejeb et al., 2014).

2.2 Salinity stress

Soil salinization is among the most critical abiotic constraints limiting agricultural productivity on a global scale. It may occur naturally (referred to as primary salinization) or arise from anthropogenic activities, particularly poor irrigation management, known as secondary salinization (Paz et al., 2023). Currently, salinity affects approximately 20% of the world’s irrigated land (around 45 million hectares) posing a substantial threat to crop yields in areas responsible for nearly one-third of global food production (Machado and Serralheiro, 2017).

From a physiological standpoint, salt-stressed plants experience an increase in reactive oxygen species (ROS) and membrane lipid peroxidation (Yadav et al., 2011; Hao et al., 2021). Additionally, stomatal regulation becomes compromised, reducing photosynthetic efficiency and carbon assimilation (Li et al., 2024). In fruit-bearing vegetables, like cucumber (Cucumis sativus L) and melon (Cucumis melo L.), salinity stress can hinder normal fruit development, often resulting in misshapen or poor-quality fruits (Joshi et al., 2013; Pinheiro et al., 2019).

2.3 Heat stress

The increasing concentration of greenhouse gases in the atmosphere is projected to raise the global average temperature by approximately 1.0 °C-1.8 °C by the end of the 21st century under very low emissions scenarios, or 2.1 °C-3.5 °C under intermediate scenarios, according to recent projections by the Intergovernmental Panel on Climate Change (Lee et al., 2023). In plants, heat stress is defined as a rise in temperature above a critical physiological threshold, sustained over a period of time long enough to cause irreversible alterations in growth and developmental processes (Wahid et al., 2007).

High temperatures interfere with the entire physiological machinery of crops. When temperatures exceed critical thresholds (often 35 °C-38 °C), enzymes involved in photosynthesis, respiration and reproductive development begin to denature or lose functionality (Wahid et al., 2007).

Pollen tube elongation, ovule fertilization, and embryo development are particularly heat-sensitive stages (Prasad et al., 2017). In tomato (Solanum lycopersicum L.), pepper (Capsicum annuum L.) and eggplant (Solanum melongena L.), this results in reduced fruit set, flower drop and malformed fruits (Abou-Hussein, 2012).

Heat stress also accelerates respiration rates, which consumes the sugars produced by photosynthesis and decreases the net energy available for growth and storage in edible parts. Leaf senescence may be triggered prematurely, reducing the effective photosynthetic area (Tan et al., 2023). Furthermore, the synthesis of pigments (e.g., anthocyanins and carotenoids) is suppressed, leading to color defects in fruits such as tomatoes and peppers (Espley and Jaakola, 2023).

2.4 Cold stress

Cold stress, including both chilling (0 °C-15 °C) and freezing (<0 °C), affects subtropical and tropical horticultural crops by damaging membranes and inhibiting vital metabolic pathways. During germination, cold conditions delay radicle emergence, reduce uniformity and increase seedling mortality (Aslam et al., 2022). In leafy crops, like lettuce (Lactuca sativa L.), chard (Beta vulgaris L. var. cicla), spinach (Spinacia oleracea L.) and arugula (Eruca vesicaria L.), low temperatures reduce photosynthetic activity and may cause water-soaked lesions and tissue collapse due to ice crystal formation (Zhou et al., 2019; Jahed et al., 2023). These effects are associated with oxidative stress and membrane damage, leading to growth inhibition and yield losses (Devireddy et al., 2021).

Chilling-sensitive fruit vegetables such as tomato (Solanum lycopersicum L.), bell pepper (Capsicum annuum L.) and melon (Cucumis melo L.), commonly exhibit symptoms such as interveinal chlorosis, leaf curling, and delayed flowering when exposed to low, non-freezing temperatures (Zhang et al., 2017; Huang et al., 2022). Cold stress also impacts reproductive development by impairing the activity of floral enzymes and limiting pollen tube growth, ultimately reducing fruit set and productivity. The accumulation of soluble sugars, which can offer partial protection, varies greatly by genotype and is often insufficient to prevent injury (Huang et al., 2022).

3 Seed Priming

3.1 Concepts and mechanisms

Seed priming is defined as a pre-sowing technique that involves the controlled hydration of seeds to initiate early metabolic processes associated with germination, without permitting radicle protrusion. After hydration, seeds are re-dried to their original moisture content, preserving viability until sowing (Sumita and Simanta, 2018). This approach enhances seed performance under both optimal and stress conditions by synchronizing germination, improving emergence uniformity and strengthening seedling vigor (de Oliveira and Gomes-Filho, 2016).

These priming techniques can be tailored to specific crop species and anticipated stress conditions. Their success depends on key variables such as priming duration, temperature, seed quality, and post-priming storage (Paparella et al., 2015). This technique follows a defined sequence of physiological stages: initial imbibition under controlled conditions, metabolic activation of enzymes and gene expression, and a drying phase to revert the seed to a storable state. This cycle induces a physiological “primed state” that improves the seed’s readiness to respond to environmental challenges and accelerates early growth under stress conditions (Wojtyla et al., 2016). Optimizing these parameters is essential to ensure consistency and maximize benefits across genotypes and environments.

3.2 Types of priming

Several seed priming techniques have been developed to enhance the resilience of horticultural crops under abiotic stress. These methods vary in complexity and mode of action, but all aim to improve the physiological and biochemical preparedness of seeds before germination.

“Hydropriming”, which consists of soaking seeds in water, is the most basic and accessible technique (Koushal et al., 2024). The main disadvantage of hydropriming lies in the difficulty of precisely controlling the degree of seed hydration, which often results in uneven water uptake among seeds and consequently leads to non-uniform germination (Lutts et al., 2016).

“Osmopriming” involves the application of osmotic agents to regulate seed water uptake by lowering the osmotic potential (Ψo) and has been widely applied to enhance drought and salinity tolerance through osmotic adjustment and antioxidant activation(de Oliveira and Gomes-Filho, 2016). Polyethylene glycol (PEG) is the most widely used osmotic agents due to its non-toxic nature and high molecular weight, which prevents cellular penetration (Lei et al., 2021). However, the high viscosity of PEG solutions can restrict oxygen diffusion during treatment and complicate the retrieval of seeds post-priming (Paparella et al., 2015). Alternatively, inorganic salts such as NaCl, CaCl₂ and KNO₃ have been employed as cost-effective and user-friendly priming agents (Castañares and Bouzo, 2018). Yet, their application demands careful calibration of both concentration and duration, as the release of ions into the solution may induce phytotoxicity and negatively impact seed viability (Paparella et al., 2015).

“Hormonal priming” uses phytohormones such as gibberellic acid (GA₃), abscisic acid (ABA), salicylic acid (SA) and jasmonic acid (JA) to activate hormonal signaling pathways related to stress tolerance (Farooq et al., 2019). The main limitation of this technique lies in the variation of treatment conditions required for different plant species (Rhaman et al., 2020).

“Biopriming”, an approach gaining prominence in sustainable agriculture, incorporates beneficial microbes like plant growth-promoting rhizobacteria (PGPR), arbuscular mycorrhizal fungi or Trichoderma spp (Chakraborti et al., 2022). These organisms not only promote germination and nutrient uptake but also bolster plant immunity and stress tolerance. However, their practical deployment is constrained by factors such as microbial viability during storage, formulation stability and varying performance under field conditions (Singh et al., 2023). “Nanopriming”, an emerging and technologically advanced method, uses nanoparticles such as ZnO, TiO₂ or silver to stimulate antioxidant enzyme activity, improve membrane stability, and enhance signaling under stress conditions (Khan et al., 2023). Redox priming involves priming agents such as hydrogen peroxide (H₂O₂) or nitric oxide (NO) that precondition the seed’s redox state, thereby strengthening antioxidant systems and improving tolerance to oxidative stress (Hussain et al., 2022).

Despite the promising results of these priming methods, especially under controlled environments, there are growing concerns about their ecological and food safety implications. In particular, nanopriming raises issues related to the potential accumulation of nanoparticles in plant tissues, which could pose toxicological risks to humans and animals if residues persist in edible parts (Chaithanya and Rao, 2023). Furthermore, nanoparticles released into agricultural soils may alter microbial communities and affect non-target organisms, potentially disrupting ecosystem dynamics (Shelar et al., 2023). Biopriming, although generally considered safe, field applications require rigorous evaluation of microbial stability, environmental persistence and unintended ecological effects (Mahmood et al., 2016).

Taken together, the diversity of seed priming strategies highlights their considerable potential in improving abiotic stress resilience in horticultural crops, but also underscores the need for species-specific optimization, safety assessments and regulatory clarity to ensure their practical applicability under real-world agricultural conditions.

3.3 Biochemical and physiological changes induced by priming

Seed priming triggers a wide range of biochemical and physiological modifications that collectively enhance the plant’s ability to tolerate abiotic stress. One of the primary effects is the activation of the antioxidant defense system, including enzymes such as superoxide dismutase (SOD), catalase (CAT) and ascorbate peroxidase (APX), which reduce oxidative damage caused by reactive oxygen species (ROS) during stress episodes (Islam et al., 2015).

Priming also improves membrane stability by increasing the synthesis of membrane-stabilizing proteins and protecting lipids from peroxidation. This is critical during desiccation and rehydration phases, and particularly beneficial in temperature and drought stress conditions (Wojtyla et al., 2016).

At the osmotic level, priming induces the accumulation of compatible solutes such as proline, glycine betaine, and soluble sugars (Ashraf and Foolad, 2005). These osmoprotectants help maintain cell turgor, stabilize enzymes and proteins and reduce water loss under saline or drought conditions (Zouari et al., 2019).

Moreover, priming enhances the repair of damaged nucleic acids and proteins, increases ATP production, and accelerates the expression of genes related to stress tolerance. This prepares seeds for faster germination, improved energy metabolism, and early seedling vigor (Wojtyla et al., 2016). Seed priming has also been shown to modulate hormonal balance, particularly enhancing the sensitivity to abscisic acid (ABA) and fine-tuning interactions with salicylic acid (SA), jasmonic acid (JA) and ethylene. This hormonal crosstalk is crucial for coordinating defense and growth processes during early seedling development under stress conditions (Wojtyla et al., 2016).

Recent studies have identified that priming can leave epigenetic marks, such as histone modifications and DNA methylation changes, which may contribute to “stress memory” and transgenerational tolerance effects, although these mechanisms are still under active investigation in horticultural species (Liu et al., 2022).

Collectively, these biochemical and molecular changes induced by seed priming result in faster germination, improved seedling establishment, and greater resilience to abiotic stressors, forming the foundation of the cross-tolerance mechanisms (Figure 1) that will be explored in the following section.

Figure 1 Molecular effects of priming

Moreover, many of the biochemical and physiological changes triggered by seed priming are not stress-specific, but rather enhance the plant’s general defensive capacity. This leads to the phenomenon known as cross-tolerance, in which primed plants exhibit improved resilience not only to the target stress but also to a broad range of abiotic challenges, such as drought, salinity, and heat (Liu et al., 2022). This multifaceted response results from the activation of interconnected signaling networks and defense pathways that allow plants to cope more efficiently with complex stress environments (Wojtyla et al., 2016; Johnson and Puthur, 2021).

In conclusion, seed priming initiates a cascade of biochemical and physiological events that significantly improve plant performance under abiotic stress. By enhancing early vigor, activating defense pathways, and inducing metabolic readiness, primed seeds gain a competitive advantage in challenging environments. These responses, coupled with the capacity for cross-tolerance, position seed priming as a strategic, low-cost intervention to promote resilience in horticultural crops.

The concept of cross-tolerance opens new avenues for sustainable horticulture, especially under the threat of climate variability and stress combinations. Although the molecular basis of cross-tolerance is still being unraveled, it represents a practical, low-cost intervention to increase the resilience of high-value crops without genetic modification.

4 Practical Applications and Case Studies of Seed Priming Under Abiotic Stress in Horticultural Crops

4.1 Drought stress

Water scarcity is a major limiting factor in horticultural production, particularly in arid and semi-arid regions. Seed priming has emerged as a promising technique to improve drought tolerance in vegetables by enhancing seed germination, seedling vigor and early stress adaptation (Jisha et al., 2013).

In carrot (Daucus carota L.), priming with agents such as PEG and GA₃ increased germination and seedling vigor by up to 28% under drought stress, with genotype-specific antioxidant responses playing a key role in tolerance (Nowicki et al., 2025). In melon (Cucumis melo L.), salicylic acid priming significantly enhanced germination, physiological performance, and fruit yield, with optimal results at 50% field capacity (Alam et al., 2022). Similarly, in green gram (Vigna radiata L.), halopriming with NaCl alleviated the negative effects of PEG-induced drought and salinity stress by maintaining shoot growth, chlorophyll content and membrane stability.

4.2 Salinity stress

Seed priming has shown significant potential to mitigate the adverse effects of salinity in several vegetable species by enhancing osmotic adjustment, antioxidant activity and ion homeostasis (Ibrahim, 2016).

In salt-sensitive lettuce (Lactuca sativa L.), hydropriming improved germination synchronization, fresh and dry biomass, and reduced membrane damage under salinity stress, outperforming KNO₃ and GA₃ treatments (Adhikari et al., 2022). Similarly, in melon (Cucumis melo L.), seed priming with NaCl or CaCl₂ for two days improved germination and early seedling growth under high salinity (8.0 dS/m), along with enhanced chlorophyll content, water balance and antioxidant response (Castañares and Bouzo, 2020).

In tomato (Solanum lycopersicum L.), seed priming, particularly with PEG, enhanced root development and ionic balance under salt stress, while reducing oxidative and osmotic damage in root tissues (Habibi et al., 2025). In turnip (Brassica rapa L.), selenium priming at 100 μmol/L increased seed germination, biomass and photosynthetic activity under salinity stress by upregulating antioxidant gene expression and reducing levels of ROS-related markers (Hussain et al., 2023). These findings support the use of seed priming as a practical and adaptable tool to enhance salinity tolerance in horticultural crops, especially during germination and early growth stages.

4.3 Heat stress

Seed priming has been explored as a strategy to improve thermotolerance by stabilizing cell membranes, enhancing antioxidant defenses, and protecting key proteins involved in stress response (Chakraborty and Dwivedi, 2021).

In spinach (Spinacia oleracea L.), osmopriming with CaCl₂ and PEG6000 significantly improved germination rates at elevated temperatures. Trials with three cultivars showed up to 50% higher germination at 20 °C in primed seeds compared to controls, while additional biopriming with Azospirillum brasilense did not provide further benefits (Breit et al., 2025). In lettuce (Lactuca sativa L.), priming with a combination of plant growth regulators (GA₃, ABA and ethylene) enhanced germination under heat stress (30 °C) by promoting endo-β-mannanase activity and weakening endosperm resistance, resulting in faster and more uniform seedling emergence (Park et al., 2022).

Similarly, in garden pea (Pisum sativum L.), osmopriming with CaCl₂ and hormopriming with salicylic acid improved germination energy, seedling vigor and physiological performance under high temperature, supporting their use in enhancing thermotolerance in leguminous vegetables (Tamindžić et al., 2023).The results confirm that seed priming is a practical and effective approach to mitigate heat stress in thermosensitive horticultural species, particularly during the critical stages of germination and early growth.

4.4 Cold stress

Under chilling stress, seed priming has emerged as a promising strategy to alleviate damage by stabilizing cellular membranes, activating antioxidant defense mechanisms, and enhancing metabolic preparedness for stress conditions (Hussain et al., 2016).

Zinc seed priming significantly enhanced spinach (Spinacia oleracea L.) germination rate and total emergence under chilling conditions (8  °C), with evidence of enhanced Zn uptake and root translocation contributing to early seedling development (Imran et al., 2021).

In chickpea (Cicer arietinum L.), seed priming with GA₃ improved emergence and reduced chilling injury under low temperatures, with effects on water retention, electrolyte leakage and early growth varying by cultivar and GA₃ dose (Aziz and Pekşen, 2020). Seed priming demonstrates notable versatility and effectiveness as a pre-sowing strategy to enhance cold tolerance in horticultural crops. By modulating key physiological and biochemical processes, such as membrane stability, antioxidant activity, nutrient uptake, and hormone signaling, it enables seedlings to better withstand chilling stress during early developmental stages. Customizing priming agents and protocols to specific crop species and environmental conditions can therefore significantly enhance germination performance and seedling vigor under low-temperature stress.

To complement the case studies described above, Table 1 summarizes the most relevant applications of seed priming under different abiotic stress conditions in horticultural crops, highlighting the priming agents used, target crops, and observed benefits.

Table 1 Overview of seed priming benefits under various abiotic stresses in horticultural crops

5 Advantages and Limitations of Multi-stress Priming

Seed priming as a technique to enhance tolerance against multiple abiotic stresses has gained significant attention in horticultural crop production. While the practical benefits of priming have been demonstrated across numerous studies and crop species, it is important to critically evaluate both the advantages and the limitations of this approach to inform its effective implementation.

5.1 Agronomic and physiological benefits

Multi-stress seed priming improves key agronomic traits such as germination rate, seedling vigor, and early establishment under adverse environmental conditions. By pre-conditioning seeds, priming triggers a cascade of physiological and biochemical changes, including enhanced antioxidant enzyme activity, osmotic adjustment, membrane stabilization, and improved nutrient uptake (de Oliveira and Gomes-Filho, 2016; Wojtyla et al., 2016). These responses collectively confer increased resilience to drought, salinity, heat, and cold stresses (Zulfiqar, 2021). Furthermore, priming can reduce the lag phase during germination, leading to more uniform and synchronous seedling emergence, a desirable trait in commercial horticulture (Jisha et al., 2013).

Studies have also shown that primed seeds exhibit better root architecture and shoot development, which translates into improved water and nutrient acquisition, ultimately enhancing yield stability under stress (Wojtyla et al., 2016).

Agronomically, seed priming offers a cost-effective, simple, and scalable method that can be easily integrated into existing seed treatment protocols (Nowicki et al., 2025). It reduces the need for expensive chemical inputs and can complement other management practices such as irrigation scheduling and fertilization (Paul et al., 2022).

5.2 Challenges: duration of effect, genetic variability, and environmental conditions

Despite its advantages, multi-stress seed priming presents several challenges that need consideration. One major limitation is the transient nature of the priming effect. The physiological enhancements induced by priming may diminish over time during seed storage, thus necessitating optimized storage conditions and timing of sowing to maintain efficacy (Hussain et al., 2015; Adhikari et al., 2024).

Genetic variability among crop species and cultivars also influences the responsiveness to priming treatments. What proves effective for one genotype may be less so for another, making it imperative to develop tailored priming protocols based on genotype-specific responses (Ashraf and Foolad, 2005).

Environmental conditions during and after priming further impact the effectiveness of treatment. Factors such as temperature, humidity and soil properties can modulate the stress level experienced by seedlings and influence the degree to which priming confers protection. Moreover, interactions between multiple stresses may be complex and priming designed to mitigate one stress could have neutral or even negative effects under combined stress scenarios (de Oliveira and Gomes-Filho, 2016).

Scaling seed priming techniques from laboratory or controlled environments to large-scale agricultural settings requires standardization and quality control to ensure consistent outcomes (Ashraf and Foolad, 2005).

6 Future Perspectives

As climate change intensifies the frequency and complexity of abiotic stress factors affecting horticultural crops, the strategic use of seed priming must evolve to meet future agricultural demands. While current studies have shown promising results, several areas require further exploration to optimize priming technologies for consistent, scalable and sustainable implementation.

6.1 Need for multifactorial assays

Most current studies focus on single-stress conditions under controlled environments, which do not fully replicate the multifaceted nature of field conditions. Horticultural crops are often subjected to combinations of drought, salinity, heat, and cold, making it imperative to develop multifactorial experimental designs that evaluate cross-tolerance mechanisms under simultaneous stress exposures. Multifactorial assays should also consider temporal aspects, such as stress duration and intensity, and their interaction with developmental stages of the plant. These comprehensive studies will be crucial for unraveling synergistic or antagonistic effects of priming treatments and tailoring protocols for real-world applications.

6.2 Role of priming combined with the microbiome, bio inputs and gene editing

The integration of seed priming with biological inputs such as beneficial microbes (e.g., plant growth-promoting rhizobacteria, endophytes and mycorrhizae) holds promise to boost plant resilience. Biopriming, which involves coating seeds with microbial consortia, has been shown to enhance nutrient uptake, hormone production, and systemic resistance under stress (Singh et al., 2023). Future research should aim to identify specific microbial strains that synergize with chemical or hormonal priming agents for enhanced multi-stress tolerance.

6.3 Importance of selecting cultivars with a good response to priming

One of the most critical factors influencing the success of priming strategies is genotypic variability. Not all cultivars respond equally to the same priming treatment; some may show enhanced germination and stress tolerance, while others may not benefit or could even experience negative effects (Ashraf and Foolad, 2005). Therefore, screening and selecting cultivars that exhibit high responsiveness to priming, especially under multi-stress conditions, should become a standard part of breeding and production programs.

To enhance the reliability of priming strategies in horticulture, future research should aim to develop cultivars with consistent responses to priming. This requires integrating priming compatibility into breeding programs, both conventional and molecular, so that treatments align with crop genetics, improving stress resilience and field performance.

6.4 Economic, policy and social dimensions of adoption

Beyond biological optimization, the broader adoption of seed priming will depend on its economic feasibility, regulatory clarity, and social acceptance among growers. Cost-benefit analyses are essential to determine the profitability of priming protocols, particularly for smallholder and resource-limited farmers. For example, studies in carrot and lentil (Lens culinaris L.) have demonstrated that seed priming can yield favorable benefit-cost ratios and increase income under stress conditions (Sharma et al., 2020; Ceritoglu et al., 2024). On-farm trials and meta-analyses also indicate that low-cost priming enhances emergence, yield and farmer returns under challenging environments (Sissoko et al., 2022).

However, concerns related to initial input costs, lack of infrastructure and access to reliable priming agents may limit adoption in certain contexts (Carrillo-Reche et al., 2018). In addition, the use of nanomaterials or microbial consortia in seed treatment may face trade barriers and regulatory constraints, especially in international markets (Ram et al., 2022; Shelar et al., 2023). Harmonization of standards, labeling requirements, and safety evaluations will be essential to facilitate trade and scale-up.

Social dimensions, including farmers' risk perception, trust in new technologies, and willingness to adopt alternative practices, are equally critical. Participatory trials, extension outreach, and localized demonstrations can help foster awareness, build confidence and tailor priming protocols to farmer needs and agroecological conditions. Ultimately, adoption will depend not only on the biological effectiveness of priming, but on its economic rationality and alignment with farmer knowledge systems.

7 Conclusions

Seed priming has emerged as a promising and practical strategy to enhance tolerance to multiple abiotic stresses in horticultural crops. By modulating physiological, biochemical, and molecular processes during the early stages of development, priming treatments can improve germination, seedling vigor, and adaptive responses to drought, salinity, heat, and cold. The review highlights the effectiveness of diverse priming agents, ranging from osmotic and hormonal solutions to micronutrients and bio stimulants, demonstrated through numerous case studies in economically important vegetables.

Despite its potential, the success of priming is influenced by several factors, including stress intensity, environmental conditions, and genotype-specific responses. Challenges such as the duration of priming effects, scalability, and variability across cultivars need to be addressed through multifactorial studies and integration with emerging technologies. Promising avenues include the combination of priming with beneficial microbes, gene editing and targeted breeding for priming-responsive cultivars.

Overall, seed priming represents a low-cost, eco-friendly tool with significant potential to improve crop performance under climate-induced stress scenarios. Its integration into holistic crop management strategies can contribute to sustainable horticulture and global food security.

Conflict of Interest

Author declares that there are no conflicts of interests.

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