Impaginato 205 Adv. Hort. Sci., 2025 39(3): 205­214 DOI: 10.36253/ahsc­17508 https://oaj.fupress.net/index.php/ahs Effect of boron priming on germination traits of shallot (Allium ascalonicum L.) Z.K. Zaiful 1, M. Faried 2 (*), E. Syam’un 2, K. Mantja 2, R.W. Putri 3, A. Jalil 4, P. Wijaya 5, C. Cennawati 6 1 Hasanuddin University, Faculty of Agriculture, Agrotechnology Study Program, Makassar, South Sulawesi, Indonesia. 2 Hasanuddin University, Faculty of Agriculture, Department of Agronomy, Makassar, South Sulawesi, Indonesia. 3 West Sulawesi University, Faculty of Agriculture and Forestry, Agroecotechnology Program, Majene, West Sulawesi, Indonesia. 4 Regional Technical Implementation Unit of the Agricultural Extension and Human Resources Development Center, Food, Crops and Horticulture Office, East Kalimantan, Indonesia. 5 Lambung Mangkurat University, Agroecotechnology Program, Agriculture Faculty, Banjarbaru, Indonesia. 6 Muhammadiyah Enrekang University, Faculty of Science and Technology, Agrotechnology Program, Makassar, South Sulawesi, Indonesia. Key words: Early growth, micronutrient, seed treatment. Abstract: Shallot (Allium ascalonicum L.) is a vital horticultural commodity in Indonesia, valued for its culinary and medicinal properties. Seed priming, especially with boron (B), represents a promising approach to enhance germination performance. This study investigated the effects of various boron concentrations on the germination performance of shallot seeds. The experiment was conducted under screen house conditions at the Faculty of Agriculture, Hasanuddin University, in May 2024, using a randomized block design with 11 boron concentrations (0­100 mg L­1) and three replications. Priming with 100 mg L­1 resulted in the best performance across several germination parameters, including as germination percentage (GP), germination speed index (GSI), the time required for 10% of seeds to germinate (T10), the time required for 50% of seeds to germinate (T50), mean germination time (MGT), mean germination rate (MGR), and germination speed coefficient (GSC). Regression analysis showed a strong positive linear relationship between boron concentration and both final germination percentage (r² = 0.84) and germination speed index (r² = 0.76). These findings suggest that boron priming, especially at 100 mg L­1, significantly enhances germination performance of shallot seeds and may as an effective method for improving seedling vigor. (*) Corresponding author: muhfaried@agri.unhas.ac.id Citation: ZAIFUL Z.K., FARIED M., SYAM’UN E., MANTJA K., PUTRI W., JALIL A., WIJAYA P., CENNAWATI C., 2025 ­ Effect of boron priming on germination traits of shallot (Allium ascalonicum L.). ­ Adv. Hort. Sci., 39(3): 205­214. ORCID: ZZK: 0009­0009­8750­0991 FM: 0000­0001­6326­1724 SE: 0000­0001­5875­118X MK: 0000­0001­6522­1689 PRW: 0000­0003­1591­668X JA: 0009­0000­5435­0717 WP: 0009­0003­0604­4100 CC: 0000­0003­4949­4016 Copyright: © 2025 Zaiful Z.K., Faried M., Syam’un E., Mantja K., Putri W., Jalil A., Wijaya P., Cennawati C. This is an open access, peer reviewed article published by Firenze University Press (https://www.fupress.com) and distributed, except where otherwise noted, under the terms of CC BY 4.0 License for content and CC0 1.0 Universal for metadata. Data Availability Statement: All relevant data are within the paper and its Supporting Information files. Competing Interests: The authors declare no conflict of interests. Received for publication 22 March 2025 Accepted for publication 28 May 2025 AHS Advances in Horticultural Science AHS ­ Firenze University Press ISSN 1592­1573 (on line) ­ 0394­6169 (print) http://doi.org/10.36253/ahsc-17508 http://oaj.fupress.net/index.php/ahs mailto:muhfaried@agri.unhas.ac.id http://orcid.org/0009-0009-8750-0991 http://orcid.org/0000-0001-6326-1724 http://orcid.org/0000-0001-5875-118X http://orcid.org/0000-0001-6522-1689 https://doi.org/10.36253/ahsc-13616 http://Orcid.org/0009-0000-5435-0717 http://orcid.org/0009-0003-0604-4100 http://orcid.org/0000-0003-4949-4016 http://orcid.org/0000-0003-4949-4016 http://www.fupress.com http://creativecommons.org/licenses/by/4.0/legalcode http://creativecommons.org/publicdomain/zero/1.0/legalcode Adv. Hort. Sci., 2025 39(3): 205­214 206 1. Introduction Shallot (Allium ascalonicum L.) is a horticultural commodity extensively cultivated by farmers across various regions in Indonesia. In addition to its wide­ spread use as a culinary spice, shallots contain numerous bioactive compounds with potential medi­ cinal properties, such as quercetin, flavonoids, saponins, tannins, glycosides, polyphenols, and alka­ loids (Devika et al., 2021). The ongoing development of shallot cultivation is driven by its various advan­ tages, including meeting national consumption needs, providing a source of income for farmers, and its potential to contribute to foreign exchange earn­ ings. These attributes position shallots as a strategi­ cally important commodity with high economic value. Shallot production in Indonesia reached 1,985,233 tons in 2023, an increase from 1,982,360 tons in 2022 but a decrease from 2,004,590 tons in 2021 (C e n t r a l B u r e a u o f S t a t i s t i c s o f I n d o n e s i a , 2024). Shallot production still experiences fluctua­ tions, often resulting in imbalances between supply and demand in the market. National demand for shallot has been increasing at a rate of approximately 5.0­6.67% annually (Shrestha et al., 2019). Therefore, efforts to boost production must continue, including improving the quality of planting materials by transi­ tioning to true shallot seed (TSS). However, chal­ lenges are often encountered during the seedling process when cultivating shallots from TSS. The nurs­ ery process, which typically lasts 30­45 days before planting, extends the time required for crop produc­ tion and delays harvest (Nciizah et al., 2020). Additionally, shallot seeds are highly susceptible to losings viability due to various factors during storage, leading to reduced germination ability and speed (Simatupang and Pangaribuan, 2022). Germination is a crucial stage in the plant life cycle that determines the success of subsequent growth stages (Atabakia et al., 2022). Seed priming is an effective method for producing high­quality seedlings (Tanjung et al., 2021). This technique can improve initial seed germination and seedling growth (Shimizu et al., 2023), accelerate germination, and enhance germination performance even under extreme environmental conditions and poor soils (Chookhampa et al., 2023). Several factors influence seed priming, including water potential, priming agents, duration, temperature, seed vigor, and storage conditions (Farooq et al., 2011; Faisal et al., 2023). Among these, the choice of priming agent is crucial for the success of seed priming. Micronutrients, such as boron (B) can be used as priming agents to enhance plant productivity and quality (Mansouri et al., 2022). Boron priming enhances seed vigor and promotes better growth and seedling development compared to untreated seeds (Chakraborty and Dwivedi, 2022). As a priming agent, boron contributes to improving and stabilizing cell wall structure and has a positive impact on reducing disease severity (Pangestuti et al., 2021). Research by Rehman et al. (2022) reported that seed priming with 0.01% boron significantly acceler­ ated seedling emergence and the time to reach 50% seedling emergence, improved germination index, leaf number, fresh and dry root weight, chlorophyll content, various yield parameters, and increased car­ bohydrate, protein, and fiber content in mung bean plants (Vigna radiata L.). Research by Nciizah et al. (2020) also revealed that seed priming with 0.01% boron reduced the number of days to seedling emer­ gence by 94% and increased seedling fresh and dry weight, as well as the chlorophyll index by 29%, 47%, and 58%, respectively, compared to the control in maize plants (Zea mays L.). Mansouri et al. (2022) found that seed priming with boron enhanced chlorophyll and carotenoid pigment content and increased the levels of enzymatic and non­enzymatic antioxidants such as anthocyanin, superoxide dismu­ tase, and peroxidase in wheat seeds. Based on the above reviews, this study aims to determine the effect of different boron concentrations as a priming agent on the germination performance of shallot seeds. 2. Materials and Methods Research location and experimental design This research was conducted under screen house conditions at the Experimental Farm of the Faculty of Agriculture, Hasanuddin University, in May 2024. The study was designed as a randomized block experi­ ment with a single treatment factor. The treatment factor was boron concentration, which included 11 levels: 0, 10, 20, 30, 40, 50, 60, 70, 80, 90, and 100 mg L­1. Each treatment level was replicated three times, resulting in a total of 33 experimental units. Priming and germination assay The seed variety used in this study was ‘Maserati’. Zaiful et al. ‐ Influence of boron as a priming agent 207 RStudio version 4.2.1 (Wickham, 2016; Shimizu et al., 2023; R Core Team, 2024). 3. Results Germination parameters The germination parameters of shallot seeds exhibited a significant response to increasing boron concentrations (Table 2 and Table 3). The FGP rose from 80.67% at 0 mg L⁻¹ to 96.00% at 100 mg L⁻¹, reflecting a 15.33% increase, which highlights boron’s positive influence on germination efficiency. Similarly, GSI improved by 63.79%, from 6.96 to 11.40, indicating enhanced metabolic activity. T10 decreased from 4.03 days to 2.40 days, showing a 40.45% reduction, while T50 decreased by 33.39%, from 6.08 days to 4.05 days. MGT also improved, dropping by 28.25% from 6.30 days to 4.52 days, reflecting faster and more uniform germination under higher boron concentrations. The application of boron significantly influenced several germination parameters of shallot seeds. MGR and GSC increased significantly with higher boron concentrations, with the highest values observed at 100 mg L⁻¹, indicating faster germination compared to the control. Although VGT and GS were not significantly affected (p > 0.05), lower VGT values at 20, 50, 80, and 100 mg L⁻¹ suggest more uniform germination, while GS tended to improve at 20, 50, and 80 mg L⁻¹. The CVGT was significantly reduced at 20 and 50 mg L⁻¹, reflecting more consistent germina­ tion timing. Overall, boron concentrations between 50 and 100 mg L⁻¹ were most effective in enhancing germination performance by improving speed, rate, and uniformity. Regression analysis The impact of boron concentration on various ger­ mination parameters of shallot seeds is depicted in the series of linear regression analyses (Table 4, Fig. 1). The analysis demonstrates that boron application significantly enhances several key germination para­ meters. For instance, FGP shows a strong positive correlation with boron concentration (R² = 0.84), indicating that higher boron levels lead to a substan­ tial increase in germination efficiency. This suggests that boron plays a vital role in breaking seed dorman­ cy and supporting metabolic processes essential for successful germination. Similarly, the GSI exhibited a positive trend (R² = The seeds were primed with a boron solution follow­ ing the specified treatment protocols. They were then placed in plastic jars containing the solution at a seed­to­solution ratio of 1:5, with an aerator added to ensure an adequate oxygen supply. The priming process lasted for 20 hours at room temperature (Ghassemi­Golezani et al., 2010; Farahzety et al., 02023; Katriani et al., 2023; Faried et al., 2024). Afterward, the seeds were removed and air­dried at room temperature for 48 hours before further use. Germination was evaluated using a soil and com­ post growth medium in a 1:1 (v/v) ratio. The medium was thoroughly mixed and placed into transparent plastic boxes with drainage holes at the bottom. Each plastic box had a volume of 500 cm³ and was filled with 500 g of the growth medium. The primed seeds were then planted by creating furrows 1 cm deep. Each plastic box contained 50 shallot seeds. After sowing, the seeds were lightly irrigated twice daily (morning and evening) with approximately 30 mL of water per application to maintain adequate soil mois­ ture under the warm ambient conditions of the experiment. Watering volumes were carefully con­ trolled to prevent waterlogging while providing opti­ mal conditions for germination. The seedling emer­ gence was observed daily until day 14. Data collection and analysis The parameters evaluated in this study included final germination percentage (FGP) (%), germination speed index (GSI), the time required for 10% of seeds to germinate (T10) (d), the time required for 50% of seeds to germinate (T50) (d), mean germination time (MGT) (d), mean germination rate (MGR), variance of germination time (VGT), coefficient of variation of germination time (CVGT), germination synchrony (GS), and germination speed coefficient (GSC) (Table 1). These ten parameters were calculated using observational data on the number of seeds germinat­ ing daily. Seeds were considered to have germinated when the plumule emerged above the surface of soil. An analysis of variance (ANOVA) was conducted to test significant differences among treatments. When significant effects were found, Duncan’s Multiple Range test was applied to separate means. Additionally, regression analysis was conducted to assess the relationship between boron concentration and the germination parameters. Calculations for the parameters were performed using the Seedcalc pack­ age, while data analysis and visualization were con­ ducted using the ggplot2 and AgroR packages in Adv. Hort. Sci., 2025 39(3): 205­214 208 0.76), indicating that higher boron concentrations accelerated germination rate. The reductions in T10 (R² = 0.43) and T50 (R² = 0.47) further supported this, as boron shortened the time needed to reach early and intermediate germination stages. MGT showed a negative correlation (R² = 0.45), reflecting a decrease Table 1 ­ Overview of the measured parameters related to seed germination No Parameters Formula 1 Final germination percentage (FGP) FGP = (n/N) x 100 n is the number of seeds germinated, and N is the total number of seed. 2 Germination Speed Index (GSI) n is the number of seeds germinated on each day of daily counting up to the last count, and t is the number of days after the beginning of the test in each count. 3 T10 Germination (T10) Days at which 10% cumulative germination is reached. 4 T50 Germination (T50) N is the final number of seeds germinated, and ni and nj are the total number of seeds germinated in adjacent counts in time ti and tj, respectively, when 5 Mean Germination Time (MGT) ni is the number of seeds germinated per day (not the accumulated number, but the number corresponding to the i­th observation), and ti is the time since the beginning of the germination test up to the i­th observation. 6 Mean Germination Rate (MGR) MGR = CoVg / 100 = 1/t t is the mean germination time, and CoVg is the germination speed coefficient. 7 Variance of Germination Time (VGT) t is the mean germination time, ti is the time between the beginning of the experiment and the i­th observation (day or hour), ni is the number of seeds germinated in time i, and k is the last count of the germination test. 8 Coefficient of Variation of Germination Time CVGT= (St / t) 100 (CVGT) St is standard deviation of the germination time, and t is mean germination time. 9 Germination Synchrony (GS) Sinc = ∑Cni, 2 / N Cni, 2 = ni (ni – 1) / 2 N = ∑ni (∑ni ­1) / 2 Cni is the combination of the seeds germinated in time i, two by two, and ni is the number of seeds germinated in time i. 10 Germination Speed Coefficient (GSC) fi is the number of newly germinated seeds on day i, and xi is the number of days from sowing. Zaiful et al. ‐ Influence of boron as a priming agent 209 in total seedling emergence time, while MGR increased (R² = 0.47), highlighting improved metabol­ ic efficiency and faster development. In contrast, VGT (R² = 0.01), CVGT (R² = 0.12), and GS (R² = 0.03) all revealed weak or negligible correlations, suggesting that boron had limited effects on uniformity, consis­ tency, and synchronization of germination. Conversely, GSC demonstrated a moderate posi­ tive correlation (R² = 0.47), emphasizing boron’s role in enhancing the speed of germination completion. Overall, these findings suggested that boron effec­ tively improved germination performance, particular­ ly in terms of percentage and speed, while having minimal impact on uniformity and synchronization. Correlation analysis The correlation matrix illustrates the relationships between different germination parameters of shallot seeds under varying boron concentrations. GP showed a strong positive correlation with GSI (r = Table 2 ­ Germination performance parameters of shallot seeds under different boron concentrations (x) Different letters within the same column indicate statistically significant differences according to Duncan's Multiple Range Test. Boron concentration (mg L­1) Final germination percentage (%) x Germination speed index x Gemination time T10 (d) x Gemination time T50 (d) x Mean germination time (d) x 0 80.67 d 7.26 de 3.52 bc 5.19 bc 5.92 ab 10 82.00 d 7.42 de 3.39 bcd 5.57 ab 5.86 abc 20 82.00 d 8.10 d 3.34 bcd 4.94 bcd 5.22 bcdef 30 82.00 d 6.96 e 3.66 ab 6.08 a 6.3 0a 40 82.00 d 8.10 d 3.18 cde 4.67 cde 5.36 bcde 50 86.00 c 7.95 de 4.03 a 5.00 bcd 5.5 9bcd 60 88.67 c 9.65 bc 2.84 efg 4.34 de 4.89 def 70 88.67 c 9.40 c 2.82 efg 4.23 de 5.21 bcdef 80 92.67 b 10.52 ab 2.67 fg 4.23 de 4.65 ef 90 95.33 a 9.99 bc 2.95 def 4.41 cde 5.14 cdef 100 96.00 a 11.40 a 2.40 g 4.05 e 4.52 f p‐value <0.01 <0.01 <0.01 <0.01 <0.01 Table 3 ­ Metabolic and synchronization parameters of shallot seed germination under different boron concentrations (x) Different letters within the same column indicate statistically significant differences according to Duncan's Multiple Range Test. Boron concentration (mg L­1) Mean germination rate (x) Variance of germination time Coefficient of variation of germination Germination synchrony Germination speed coefficient (x) 0 0.169 de 2.38 25.04 bc 0.24 16.95 de 10 0.170 de 2.11 24.73 bc 0.25 17.05 de 20 0.191 bcd 0.87 17.91 c 0.34 19.12 bcd 30 0.158 e 2.57 25.31 bc 0.22 15.88 e 40 0.187 cd 2.09 24.81 bc 0.25 18.79 cd 50 0.179 cde 1.04 17.42 c 0.39 17.98 cde 60 0.204 abc 1.74 26.31 abc 0.30 20.48 abc 70 0.192 bcd 3.05 32.00 ab 0.24 19.26 bcd 80 0.215 ab 1.06 22.27 bc 0.33 21.59 ab 90 0.194 bcd 4.07 36.78 a 0.29 19.48 bcd 100 0.220 a 1.64 27.81 abc 0.28 22.08 a p‐value <0.01 >0.05 <0.01 >0.05 <0.01 210 Adv. Hort. Sci., 2025 39(3): 205­214 Table 4 ­ Regression analysis of parameters No. Paramaters Equation R2 1 Final germination percentage (%) Y = 78.6 + 0.165 0.84 2 Germination speed index Y = 6.77 + 0.040 0.76 3 Time to 10% seed germinated Y = 3.69 + 0.010 0.43 4 Time to 50% seed germinated Y = 5.55 ­ 0.015 0.47 5 Mean germination time Y = 5.99 – 0.013 0.45 6 Mean germination rate Y = 0.166 + 0.000466 0.47 7 Variance of germination time Y = 1.82 + 0.00482 0.01 8 Coefficient of variation of germination time Y = 21.4 + 0.0819 0.12 9 Germination synchrony Y = 0.27 + 0.000365 0.03 10 Germination speed coefficient Y = 16.6 + 0.0466 0.47 Fig. 1 ­ Regression analysis of each germination trait to boron concentration. Zaiful et al. ‐ Influence of boron as a priming agent 211 0.90), suggesting that higher germination percent­ ages are associated with faster germination rates. However, GP negatively correlates with T10 (r = ­ 0.64), T50 (r = ­0.69), and MGT (r = ­0.71), indicating that increased boron concentration reduces the time needed for germination (Fig. 2). GSI also exhibited significant positive correlations with MGR (r = 0.93) and VGT) (r = 0.93), showing that faster germination is linked to higher consistency in seed performance. Conversely, GSI has a strong neg­ ative correlation with MGT (r = ­0.91), emphasizing that a higher speed index leads to shorter germina­ tion times. T10, T50, and MGT parameters displayed high intercorrelations, with T10 and T50 showing r = 0.78 and T50 and MGT showing r = 0.87. Parameters related to uniformity, such as VGT and GS, showed weaker and mostly non­significant correlations with the other considered parameters. 4. Discussion and Conclusions Seed priming is a crucial technique in agriculture aimed at enhancing seed germination and seedling establishment under various stress conditions. By promoting pre­germination metabolic activity through controlled hydration and drying, seed prim­ ing can significantly improve germination rates and seedling vigor, helping plants to overcome abiotic stresses such as salinity and drought (Chatterjee et al., 2018; Hameed et al., 2025; MacDonald and Mohan, 2025). The positive impacts of seed priming are not limited to increasing germination and vigor, but also extend to improving crop yields and quality, making it an essential practice for sustainable agricul­ ture (Gour et al., 2023; Jarrar et al., 2024; Zhang et al., 2025). The significant improvement in shallot seed ger­ mination parameters in response to increasing boron concentrations highlights boron’s essential role in seed physiology. The germination percentage increased by 15.33%, from 80.67% at 0 mg L⁻¹ to 96.00% at 100 mg L⁻¹, demonstrating boron’s positive impact on germination efficiency. This improvement is likely due to biochemical and physiological changes induced by priming, which enhance water uptake, activate metabolic pathways, and initiate early growth processes. Bajwa et al. (2018) similarly found that priming wheat seeds increased alpha­amylase activity, facilitating carbohydrate breakdown and providing energy for embryo development. A compa­ rable enzymatic activation may contribute to the enhanced germination observed in shallot seeds. Boron also accelerated germination times, as indi­ cated by the 40.45% reduction in time to 10% germi­ nation, a 33.39% decrease in time to 50% germina­ tion, and a 28.25% reduction in mean germination time. The shortest durations were recorded at 2.40 days (T10), 4.05 days (T50), and 4.52 days (MGT), all significantly lower than the control. These findings align with previous research by Rehman et al. (2022), which demonstrated that boron priming accelerates the germination process. However, it is important to consider that at certain concentrations, boron can become toxic to plants (Hussain et al., 2011; Atique­ ur­Rehman et al., 2020). Additionally, boron priming enhanced the germi­ nation speed index. At 100 mg L⁻¹, the highest GSI (11.40) was recorded, whereas the control treatment had the lowest value (6.96). This suggests that boron plays a crucial role in expediting germination, likely due to its involvement in cell wall formation, mem­ brane integrity, and metabolic regulation. Similar results were reported by Farooq et al. (2011), who found that boron priming significantly increased the germination index of rice seeds. Kaya and Ergin (2023), in a study conducted on Carthamus tinctorius seeds, also observed a linear increase in germination index with higher boron concentrations, reinforcing boron’s beneficial effects on seed germination. Interestingly, the coefficient variation of germina­ tion time increased by 105.46%, reflecting improved Fig. 2 ­ Correlation analysis among parameters. synchronization of the germination process. This sug­ gests that boron not only accelerates germination but also promotes more uniform seedling establish­ ment. However, the lack of significant differences in germination synchrony and the stability of the germi­ nation speed coefficient indicate that boron’s impact is more pronounced in accelerating germination rather than reducing variability. Chookhampa et al. (2023) also reported that seed priming on Andrographis paniculata, Sesamum indicum, and Abelmoschus esculentus improves the germination performance, leading to more synchronized germina­ tion. The limited effect of boron on germination uni­ formity may be influenced by inherent genetic fac­ tors and environmental conditions. Beyond improving germination rates and speed, priming has also been linked to enhanced seedling vigor and quality (Biradar et al., 2023; Faried et al., 2023). Overall, this study demonstrates that increas­ ing boron concentrations positively influence germi­ nation percentage, speed index, and synchronization, while also accelerating the overall germination process. These findings underscore boron’s potential as a seed priming agent to enhance germination effi­ ciency and metabolic activity in shallot seeds. However, the effects observed are indeed influenced by multiple factors. The success of seed priming depends on the choice of priming agent, its concen­ tration, the duration of priming, and the specific response of the plant species or variety (Siyar et al., 2020; Corbineau et al., 2023; Fu et al., 2024; Jarrar et al., 2024). Moreover, priming cannot overcome issues arising from genetic factors; seeds lacking in genetic purity, will result in poorly growing plants (Reed et al., 2022). Priming with 100 mg L­1 produced the most favor­ able outcomes on several germination parameters, such as final germination percentage, germination speed index, time required for 10% of seeds to ger­ minate, time required for 50% of seeds to germinate, mean germination time, mean germination rate, and germination speed coefficient. Regression analysis showed a strong positive linear relationship between boron concentration and final germination percent­ age (r² = 0.84) as well as germination speed index (r² = 0.76). The study concludes that boron priming, par­ ticularly at 100 mg L­1, significantly improves germi­ nation performance in shallot seeds, highlighting its potential to enhance seedling vigor and productivity. The results of this research offer valuable insights for both shallot researchers and farmers. Adv. Hort. 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