## 4 Discussion
### ROS Production and Lipid peroxidation  
Aging during storage is a major cause of decreased seed quality [22; 62] in terms of germinability and longevity. During seed imbibition, lipids are engaged in membrane reorganization while facing free-radical-mediated oxidative injury [63]. Lipid peroxidation causes imbibition damage [64] when dry seeds rapidly uptake water, affecting seed quality [65]. An increasing number of hydration-rehydration cycles also results in decreased seed viability [66]. Furthermore, biochemical and cellular events triggered by water uptake are accompanied by the generation of ROS, especially H\({}_{2}\)O\({}_{2}\)[67], one of the primary ROS [68] present in plant tissues within certain normal limits [69]. It has two roles, i.e., beneficial [68] and deleterious [51; 67; 70]. Synthesis of H\({}_{2}\)O\({}_{2}\) results in an oxidative burst upon application of related elicitors such as high temperature or mechanical damage [68]. At high levels, H\({}_{2}\)O\({}_{2}\) may attack a variety of susceptible enzymes and ultrastructural cell components with detrimental effects [71; 72], which may affect the growth axis potential and eventually lead to cell death. In this study, H\({}_{2}\)O\({}_{2}\) may have possibly been stimulated by water imbibition during seed priming resulting from increased respiration. In a similar study, [PERSON] et al. [67] indicated that an intense increase in respiratory activity spurs H\({}_{2}\)O\({}_{2}\) production, which continues due to the storage condition stresses and the reduction in the antioxidative mechanisms. Furthermore, there was a positive correlation between the production of H\({}_{2}\)O\({}_{2}\) and MDA accumulated in the seed, which was highest after 120 days of storage. H\({}_{2}\)O\({}_{2}\)-producing and consuming processes are active early in germination [73]. Predominantly, catalase and other H\({}_{2}\)O\({}_{2}\)-consuming enzymes, peroxididases, superoxide dismutase, and glutathione peroxidase have been recorded as active in the early stages of the imbibition phase. [PERSON] et al. [74] observed a high decrease in catalase and glutathione reductase activity on the first day of accelerated aging and a slight transient increase in the activity of  
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the enzymes SOD, CAT, and GR. Moreover, genes associated with cell wall synthesis and degradation were also activated during the process of imbibition, and most cell-wall-related genes were upregulated under H\({}_{2}\)O\({}_{2}\) treatment [70]. The increased antioxidant capacity of the primed seed maintained the H\({}_{2}\)O\({}_{2}\) at levels comparable to the non-primed seed (Table 4), and the reduction in the antioxidant capacity may have led to the increasing levels of H\({}_{2}\)O\({}_{2}\). In this study, for hydropriming, the changes in MDA were insignificant until after 60 days and 90 days for halopriming and osmopriming. A significant change occurred for the non-primed seeds at 120 days of storage, but the levels were higher than the primed seeds. Similarly, MDA content was enhanced in primed Chinese cabbage seeds [29] and sunflowers [74] with increased storage duration and temperature. Moreover, in the non-primed seeds of barley exposed to stress [42] and in pepper [59], this was accompanied by increased EC and a decrease in ROS antioxidants and seed vigor. Additionally, MDA is often employed as an indicator of lipid peroxidation, an end-product that easily penetrates biological membranes and affects other cellular components. This study documented a decline in seed vigor and viability in storage after 120 days. In a similar study, conventional or traditional storage of rice seed at 25 \({}^{\circ}\)C recorded a reduction in germination parameters [10]. Furthermore, high temperature and relative air humidity during storage periods [35; 75] affect seed quality and are considered critical environmental factors that directly influence deterioration.