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Learn more: PMC Disclaimer | PMC Copyright Notice Sci Rep . 2026 Apr 6;16:11927. doi: 10.1038/s41598-026-46146-0 Search in PMC Search in PubMed View in NLM Catalog Add to search Enhancing tuber quality, storage performance and yield response of potato to combined foliar application of zinc sulphate and boric acid Badran S Agha Badran S Agha 1 Department of Horticulture and Landscape Design, College of Agriculture and Forestry, University of Mosul, Mosul, Iraq Find articles by Badran S Agha 1 , Muhanad A Ahmad Muhanad A Ahmad 1 Department of Horticulture and Landscape Design, College of Agriculture and Forestry, University of Mosul, Mosul, Iraq Find articles by Muhanad A Ahmad 1 , Esraa Abd-Alhuseein Jasim Esraa Abd-Alhuseein Jasim 1 Department of Horticulture and Landscape Design, College of Agriculture and Forestry, University of Mosul, Mosul, Iraq Find articles by Esraa Abd-Alhuseein Jasim 1 , Ammar Z A Kassab Bashi Ammar Z A Kassab Bashi 1 Department of Horticulture and Landscape Design, College of Agriculture and Forestry, University of Mosul, Mosul, Iraq Find articles by Ammar Z A Kassab Bashi 1 , Heidar Meftahizade Heidar Meftahizade 2 Department of Horticultural Sciences, Faculty of Agriculture & Natural Resources, Ardakan University, P.O. Box 184, Ardakan, Iran Find articles by Heidar Meftahizade 2, ✉ Author information Article notes Copyright and License information 1 Department of Horticulture and Landscape Design, College of Agriculture and Forestry, University of Mosul, Mosul, Iraq 2 Department of Horticultural Sciences, Faculty of Agriculture & Natural Resources, Ardakan University, P.O. Box 184, Ardakan, Iran ✉ Corresponding author. Received 2025 Nov 19; Accepted 2026 Mar 24; Collection date 2026. © The Author(s) 2026 Open Access This article is licensed under a Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License, which permits any non-commercial use, sharing, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if you modified the licensed material. You do not have permission under this licence to share adapted material derived from this article or parts of it. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by-nc-nd/4.0/ . PMC Copyright notice PMCID: PMC13066163 PMID: 41942596 Abstract Potato ( Solanum tuberosum L.) is a popular food source worldwide, cultivated for tuber consumption, and plays an important role in food security in developing countries. Potato plants (Montreal cultivar) was exposed to Zinc Sulphate (0, 500, and 1000 mg/L) and Boric Acid (0, 50, and 100 mg/L) in a two factorials experiment based on a randomized complete block design (RCBD) with three replications. The experiment was conducted in the Vegetable Field at the University of Mosul during the 2024 spring growing season. Application of Zinc Sulphate (1000 mg L − 1 ) and Boric Acid (100 mg L − 1 ), separately or in combined, increased the plant height, the number of aerial stems, the leaf area, the yield per plant and per hectare. During postharvest storage, the amounts of dry matter (16.14%), starch percentage (18.00%), tuber Boron (38.91%), tuber zinc (17.55%), and tuber firmness (10.19%) were decreased, while the TSS value was increased by 24.17%. Foliar application of combined treatments from Zinc Sulphate and Boric Acid (500 or 1000 mg/L Zinc Sulphate with 50 or 100 mg/L Boric Acid) significantly slow these trends during post-harvest phase. However, the combination of 1000 mg L − 1 Zinc Sulphate and 100 mg L − 1 Boric Acid yielded the greatest values on all of the traits examined associated with tuber quality. This treatment maintained the greatest tuber firmness (18.12 lb in²), dry matter content (19.00%), specific gravity (1.07 g cm-²), starch percentage (12.94%), Zn (24.36 mg/L) and B (22.50 mg/L) contents, and the least amount of weight lost (5.23%) and total accumulation of soluble solids (5.15%) during the period of cold storage. Thus, we recommend the application of 1000 mg/L Zinc Sulphate + 100 g/L Boric Acid to improve yield and quality of potato tubers. Keywords: Potato, Boron, Zinc, Foliar application, cold Storage Subject terms: Biotechnology, Plant sciences Introduction Potato ( Solanum tuberosum L.) is a well-known commonly herbaceous species with fleshy stem and buds in the axils of leaf scars, which cultivated for its tuber 1 . Potato is a major food crop cultivated worldwide, contributing in developing countries food security 2 – 5 . Potato is recognized as one of the chief economic crops which produces more dry matter and protein per hectare than the major cereal crops 2 , 6 , 7 . In addition to starch, potatoes also provide protein, vitamins, and minerals. Potato tubers are a proven source of vitamin C, vitamin B6, and essential minerals, such potassium, magnesium, phosphorus, manganese, zinc, and even iron 2 , 6 , 8 , 9 . Vitamins and micronutrients are natural bioactive compounds, involving in vital processes in humans, plants, and animals 10 – 12 . Micronutrient fertilization is a valuable pre-harvest treatment which improved physico-chemical and nutritional quality of potato 13 . The micronutrient shortages are typically modified through precise foliar application of the micronutrient. Foliar spraying of micronutrients has been proved to be more effective than other modes because it provides a more instant and adequate supply of nutrients during the vital crop development phases 14 . Foliar Zinc (Zinc Sulphate) application is an eco-friendly, operational and economical way in plant fertilization compared to soil application 15 , 16 . Zinc is a crucial micronutrients in plant fertilization programs for fortifying the crop yield and quality 17 . Zn as a mandatory trace element, is a vital factor for plant physiological processes, such photosynthesis, carbohydrate and protein metabolism, and growth regulation 18 – 24 . Zinc is well-known as a vital activator of several enzymes in plants. Besides, it is directly involved in the biosynthesis of growth substances such as auxin, which produces more plant cells, thus increasing the dry matter 1 . Zinc deficiency causes disruption of plant metabolism, growth restriction, and yield reduction 25 – 28 . Zn involved in tryptophan biosynthesis, a necessary amino acid for biosynthesis of IAA, which controls cell expansion and elongation 19 , 20 . Some studies cleared that Zn deficiency impairs zinc-dependent enzymatic and metabolic systems, and leads to physiological stress 21 , 29 . Guan et al. (2025) stated that Zn boosts N uptake in apple trees by regulating the antioxidant system and C–N metabolism. They observed that dry weight and N accumulation in Zn-treated trees were higher, compared to control plants, about 33.85% and 40.54%, respectively 30 . Disparity fertilization worsen Zinc deficiency by antagonizing its uptake 31 – 35 . Zn deficit is a predominant trouble in crop production. Boron plays a critical role in cis diol complexation with glyco-proteins 36 – 38 , which are structural constituents of the plasma membrane. It forms a diester bond between apiosyl residues and two monomers of pectic polysaccharide rhamnogalacturonan II (RG-II), forming the B-RG-II complex 36 , 39 , which helps cell wall strengthen. Boron deficit changes the membrane potential and reduces the activity of proton-pumping ATPase and thus the proton gradient across the plasma membrane 37 , 40 , 41 . This can lead to disruption in cell wall function and nutrient uptake. Furthermore, B is involved in ion fluxes (H + , K + , PO 4 3− , Rb + , Ca 2+ ) across the membranes, cell division and elongation, nitrogen fixation, nitrogen and carbohydrate metabolism, sugar transport, cytoskeletal proteins, and plasmalemma-bound enzymes, nucleic acid, indoleacetic acid, polyamines, ascorbic acid, and phenol metabolism and transport 37 , 38 , 41 . Imbalanced Boron uptake disrupts the pollination process which consequently reduces flowering, fruit set, yield, and henceforth deteriorates fruit quality by increasing fruit acidity 41 . Application of Boron fertilizers at different rate has shown a significant influence on the yield and quality of fruits 41 . The high concentration of B caused inhibition of root and shoot growth with noticeable signs of stress on leaves in terms of chlorophyll contents, oxidative stress and lipid oxidation of membranes in rice seedlings 42 . Boron improved resistance to drought stress in coffee and its interaction with calcium influences productivity, seed composition, and soil fertility 43 . The previous studies confirmed that micronutrients deficiency reduces quality and yield of potato tubers before and after storage. Improving agronomic practices with micronutrients has noteworthy potential to improve potato yield and postharvest quality, which not only increases food security but also improves the economic returns of producers. Potatoes are very sensitive to B deficiency as well as Zinc. Therefore, we investigated the composition of these micronutrients to assess pre- and post-harvest quantity and quality in potatoes. The findings of this study provide important information about the potential of nutrient management to improve potato yield and storage quality. Materials and methods Experimental site and soil preparation The experiment was conducted in the vegetable field of Department of Horticulture and Landscape engineering College of Agriculture and Forestry, University of Mosul, spring of 2024. Before planting, surface soil was sampled at a depth of up to 30 cm in a composite sampling procedure to measure the physical and chemical characteristics of the soil. The specification of farm soil presented in Table 1 . No fertilizer was used except for foliar application of micronutrients. Table 1. The chemical and physical properties of the farm soil (analyzed at Central Laboratory -College of Agriculture and Forestry- University of Mosul). Acidity (%) EC (dS.m⁻¹) O.M. (%) N (mg/L) P (mg/L) K (mg/L) Fractions (%) Texture Sand Clay Silt 7.0 0.50 2.25 430 0.27 2.20 38.05 32.7 29.25 Clay Loam Open in a new tab Experimental design and treatments The factorial experiment was done in a randomized complete block design (RCBD) with three replications. Two factors were considered in the study including foliar application of Zinc sulfate (36% Zn) foliar-applied at three concentrations (0, 500 and 1000 mg L − 1 ) (Factor A) and Boric Acid (17% B) in three levels (0, 50, 100 mg L − 1 ) (factor B). The micronutrient levels applied in this experiment were selected based on the results of a previous study on the potato cultivar Argana 44 . Planting and agronomic practices Agronomic activities were standardized in the experiments plots, such as weeding, ridging and preventive and treatment of pests, diseases and weeds. The size of each experimental plot was 4 × 0.75 m (equal to 3 m 2 ). The tubers were planted in mid-February, 13 plants per ridge, at a depth of 15 cm, using a drip irrigation system. The potato cultivar used was Montreal, a Dutch variety developed from the hybridization of Amora × Amalia. This cultivar is characterized by early maturity, large, oval, uniform tubers with light yellow skin and yellow flesh. It is widely cultivated in region and does not require any special permit for cultivation. Treatments schedule The first spray was applied after transplanting in the 4–6 leaves stage, followed by two additional applications every 15 days. Each plant received approximately 100 mL of foliar solution per application without surfactant agent. following commonly reported volumes in similar studies to ensure proper nutrient uptake 44 . In the combined treatment, Zinc Sulphate and Boric Acid were applied together as a single foliar spray solution to each plant 44 . Postharvest assays For postharvest assays, after harvesting, the tubers were dried for two weeks in a shaded area at 15 °C. After drying, the tubers were packed in bags (5 kg per treatment per replicate) and stored for five months in a cold storage at 4.4 °C and 90–95% relative humidity 44 . Storage conditions, including temperature and humidity, were continuously monitored using a calibrated thermohygrometer to ensure stable conditions throughout the storage period. Trait measurements Plant height (cm) was measured with measuring tape from soil surface to the end of the tallest leaf for 30 plants. Number of aerial stems (stems plant⁻¹) were counted for 30 random plants and a mean was calculated. Single leaf area (cm²) was measured from means of 30 leaves using a Light Box (ADC, UK) 45 . Single plant yield (kg plant⁻¹) was calculated from dividing the total yield of the plot by the number of plants on the plot. Total yield (ton ha⁻¹) was measure by multiplication of plant yield × number of plants/ha. The percentage of weight loss (%) was calculated using the Eq. 1 . 1 Total soluble solids (TSS) before and after storage were determined by a hand refractometer in A.O.A.C. (2000). Dry matter (%) was calculated by Eq. 2 . 2 Starch percentage in tubers for before and after storage (%) were estimated according to AOAC (1970) using the Eq. 3 . 3 Tuber specific gravity before and after storage (g cm⁻³) was measured with Eq. 4 . 4 Potatoes firmness was measured using UC. Firmness-tester and calculated as Ib in − 2 using Eq. 5 . 5 Tuber Boron content before and after storage (mg L⁻¹) were assayed using the dry burned method 46 . The 0.5 g of ground plant material was burned in an electric furnace at 550 °C, the ash was dissolved in HCl, the volume adjusted to 50 mL with distilled water, and the solution was analyzed for Boron using an atomic absorption spectrophotometer. For tuber Zinc content (mg L⁻¹) estimation, samples were dried in an oven at 75 °C until constant weight, ground, and 0.5 g of each sample was digested with a mixture of H₂SO₄ and HClO₄. After digestion, the volume was adjusted to 50 mL with distilled water, and Zinc content was determined using an atomic absorption spectrophotometer 46 . The 30 plants from each treatment in each replication taken randomly and tagged for records of the total yield and tubers quality parameters. Number of tubers per plant was determined just after harvesting time (120 days from planting) using the average number of tubers of 30 plants. Average tuber fresh weight (g) was determined immediately after harvesting, by dividing the weight of tubers yield by tuber’s number of 30 plants. The number of tubers/10 kg, was determined by taking a random sample of 10 kg of tubers from the yield of each treatment and then counted. Average tuber yield per plant (g/plant) was calculated using the average weight of tubers of 30 plants. Tubers dry weight (%) was carried out via randomly tuber samples of 100 g of fresh weight which were dried in an electrical oven at 70 till the constant weight, then the obtained value of tuber dry weight was calculated as a percentage. Total soluble solids (TSS %), or degrees Brix (Bx) is numerically equal to the percentage of sugar and others dissolved in a solution 47 was estimated in the juice of the fresh tubers using a hand refractometer according to AOAC (1992). Statistical analysis Data were analyzed using a combination of univariate and multivariate statistical methods including Tow way- ANOVA, Tukey’s HSD test, Principal Component Analysis (PCA), Pearson’s Correlation Estimation, and linear regression using R software (Ver. 4.5). Results Results exhibited that application of all combination treatments of Zinc Sulphate and Boric Acid significantly enhanced morpho-physiological, agronomic and qualitative traits of potato except TSS and specific gravity. Increasing Zinc Sulphate or Boric acid concentration alone or in combined enhanced tuber growth, performance, and quality. However, the most pronounced synergistic effect was observed for combined dose of 1000 mg/L Zinc Sulphate + 100 mg/L Boric Acid. So that, the highest values for morpho-physiological, agronomic, physical, and biochemical attributes were recorded in response to application of 1000 mg/L Zinc Sulphate + 100 mg/L Boric acid, while the lowest of most traits were observed in control plants. Furthermore, application of Zinc Sulphate and Boric acid inhibited the loss of tuber weight during post-harvest storage. The interaction effect of Zinc Sulphate × Boric Acid was significant for all studied traits ( p < 0.01). This combination substantially influenced plant height, shoot number, leaf area, dry matter, and total tubers per plant (Fig. 1 a–f). The lowest values for plant height (44.67 cm), shoot number (2.80), leaf area (28.45 cm²), dry matter before storage (15.10%) and after storage (12.43%), and total tubers per plant (7.40) were observed in control plants (respectively, Fig. 1 a– f). While, the highest plant height (64.67 cm), shoot number (6.13), leaf area (59.60 cm²), dry matter before storage (21.37%), dry matter after storage (19%), total tubers/Plant (13.43 n) (respectively, Fig. 1 a and f) were obtained in response to 1000 mg/L Zinc Sulphate + 100 mg/L Boric Acid (Fig. 1 a– f). Fig. 1. Open in a new tab The effect of combined levels of Zinc Sulphate and Boric Acid on plant height ( a ), leaf area ( b ), shoot number ( c ) of potato plants, and dry matter before storage (BS) ( d ) and dry matter after storage (AS) of potato tubers ( e ), in addition, total number of tubers per plant ( f ). Tukey’s test (α = 0.05, n = 30). The combined application of Zinc sulphate and Boric Acid substantially ( p < 0.01) influenced potato growth and tuber quality parameters, including dry matter, specific gravity, starch content, mineral accumulation, firmness, and total soluble solids (TSS). The interaction effect of Zinc Sulphate × Boric Acid was significant for yield per plant, total yield, Tuber firmness before storage and after storage, and tuber weight loss ( p < 0.01). The highest yield per plant (1.58 kg) (Fig. 2 a) and total yield (70.09-ton ha⁻¹) (Fig. 2 b) were obtained in plants treated with 1000 mg/L Zinc Sulphate and 100 mg/L Boric Acid, while the lowermost were gained in control plants. Tuber firmness in before storage stage (BS) was improved significantly from 15.17 lb in⁻² in the control tubers (minimum) to 19.33 lb in⁻² (highest) in plants treated with combination of 1000 mg/L Zinc Sulphate + 100 mg/L Boric Acid, suggesting stronger tissue structure and storage stability (Fig. 2 c). Correspondingly, tuber firmness, in after storage stage (AS), was improved significantly in response of micronutrient levels, so that the least firmness (12.50 Ib in⁻² in the control tubers (minimum) and the highest (18.12 lb in⁻²) was observed in those treated with 1000 mg/L Zinc Sulphate + 100 mg/L Boric Acid (Fig. 2 d). The maximum tuber weight loss (9.83%) during storage period was observed in control plants, while the minimum (5.223%) was observed in plants treated with 1000 mg/L Zinc Sulphate + 100 mg/L Boric Acid (Fig. 2 e). Fig. 2. Open in a new tab The effect of combined levels of Zinc Sulphate and Boric Acid on yield per plant ( a ), total yield ( b ), tuber firmness before storage (BS) ( c ), tuber firmness after storage (AS) ( d ), and tuber weight loss after storage ( e ). Tukey’s test (α = 0.05, n = 30). Also, the B and Zn contents of tubers (before and after storage) and starch percentage of tubers (before and after storage) were positively affected by foliar application of Zinc Sulphate and Boric Acid ( p < 0.01). The application of Zinc Sulphate and Boric Acid considerably affected mineral accumulation and starch percentage in tubers ( p < 0.01). Both Zinc and Boron concentrations in tubers were increased distinctly with increasing treatment levels. The uppermost Boron value (32.67 mg/L) and Zinc content (29.00 mg/L) in pre-storage phase were assayed in response to application of 1000 mg/L Zinc Sulphate + 100 mg/L Boric Acid (Fig. 3 a and c). Furthermore, in post storage stage (AS), the highest B and Zn content also, were observed in tubers treated with 1000 mg/L Zinc Sulphate + 100 mg/L Boric Acid (22.50 mg L⁻¹ and 24.36 mg L⁻¹, respectively (Fig. 3 b and d). Starch percentage also increased notably, from 9.47% in control tubers to 15.05% in plants treated with 1000 mg/L Zinc Sulphate + 100 mg/L Boric Acid (before storage) and from 8.13% (control) to 12.94% in response to 1000 mg/L Zinc Sulphate + 100 mg/L Boric Acid (after storage) (Fig. 3 e and f). Conversely, tuber weight loss during storage decreased from 9.83% in the control to 5.23% under the highest combined treatment (1000 mg/L Zinc Sulphate + 100 mg/L Boric Acid), implying better physiological integrity and reduced dehydration Fig. 3 . Fig. 3. Open in a new tab The effect of combined levels of Zinc Sulphate and Boric Acid on tuber B content in pre storage stage (BS) ( a ), tuber B content after storage (AS) ( b ), tuber Zn content in pre storage stage (BS)( c ), tuber Zn content after storage (AS) ( d ), and tuber starch percentage before storage (BS) ( e ) and after storage (AS) ( f ), Tukey’s test (α = 0.05, n = 30). Total soluble solids of tubers showed a slight decrease with higher nutrient levels. The maximum TSS before storage (5.17%) and after storage (7.00%) were evaluated in control plants, however the least TSS before storage (3.97%) was measured in response to Zinc Sulphate at 0 mg/L + Boric Acid at 100 mg/L and the its lowest after storage (5.15%) in response to Zinc Sulphate at 1000 mg/L plus Boric Acid at 100 mg/L (Fig. 4 a, b). The specific gravity of tubers was not influenced meaningfully by application of treatments, however the minimum specific gravity before storage (1.06 g cm-²) and after storage (1.04 g cm-²) was assessed in control plants, while the supreme amounts by 1.09 and 1.07 g cm-² in tubers gained from those plants which treated with 1000 mg/L Zinc Sulphate + 100 mg/L Boric Acid (Fig. 4 a, b). Fig. 4. Open in a new tab The effect of combined levels of Zinc Sulphate and Boric Acid on total soluble solids of potato tuber before storage (BS) ( a ) and after storage (AS) ( b ), in addition, tuber specific gravity before storage (BS) ( c ) and after storage (AS) ( d ), Tukey’s test (α = 0.05, n = 30). Principal component analysis In the principal component analysis (PCA), the first two principal components (PC1 and PC2) accounted for 79.4% and 9.4% of the total variance, respectively, explaining 88.8% of the overall variability among the measured traits (Fig. 5 ) The treatment 1000 mg L⁻¹ Zinc Sulphate + 100 mg L⁻¹ Boric Acid closely associated with yield-related and quality traits, including total tuber yield, tuber Boron and Zinc content after storage, starch accumulation, specific gravity, and dry matter content. In contrast, treatments without Zinc Sulphate and Boric Acid supplementation (0 mg L⁻¹) correlated more strongly with vegetative traits such as plant height, shoot number, and leaf area. These observations indicated that increasing Zinc Sulphate and Boric Acid concentrations acted synergistically to enhance tuber yield, quality, and mineral composition in potato. Fig. 5. Open in a new tab The graph of Principal Component Analysis (PC1 and PC2) for different studied traits of Potato in response to different Zinc Sulphate + Boric Acid levels. BS: before storage, AS: after storage. Pearson’s correlation estimation Pearson’s correlation analysis identified statistically significant associations among agronomic, physiological, and tuber quality traits before storage (BS) and after storage (AS). Total yield (ton ha⁻¹) exhibited strong positive correlations with tuber Boron before storage ( r = 0.927 ** ), tuber Boron after storage ( r = 0.917 ** ), total tubers per plant ( r = 0.876 ** ), leaf area ( r = 0.807 ** ), dry matter before storage ( r = 0.829 ** ), dry matter after storage ( r = 0.832 ** ), specific gravity before storage ( r = 0.829 ** ), specific gravity after storage ( r = 0.832 ** ), starch before storage ( r = 0.829 ** ), starch after storage ( r = 0.832 ** ), and tuber firmness after storage ( r = 0.836 ** ), while showing moderate associations with, tuber Zinc before storage ( r = 0.656 ** ), and tuber Zinc after storage ( r = 0.654 ** ). Tuber Boron before storage was strongly correlated with tuber Boron after storage ( r = 0.979 ** ), tuber firmness after storage ( r = 0.876 ** ), shoot number ( r = 0.861 ** ), and leaf area ( r = 0.765 ** ), and negatively with tuber weight loss ( r = − 0.855 ** ). Tuber Zinc after storage showed high positive correlations with tuber Zinc before storage ( r = 0.975 ** ), shoot number ( r = 0.825 ** ), tuber firmness after storage ( r = 0.845 ** ), and total tubers per plant ( r = 0.760 ** ), and a strong negative link with tuber weight loss ( r = − 0.743 ** ). Leaf area was strongly associated with total tubers per plant ( r = 0.933 ** ), dry matter after storage ( r = 0.829 ** ), specific gravity after storage ( r = 0.829 ** ), starch after storage ( r = 0.829 ** ), and tuber firmness after storage ( r = 0.764 ** ), and negatively with tuber weight loss ( r = − 0.918 ** ) and TSS BS ( r = − 0.823 ** ). Dry matter before storage, specific gravity before storage, and starch before storage were correlated with total tubers per plant ( r = 0.918 ** , 0.918 ** , 0.918 ** ) and yield per plant ( r = 0.893 ** , 0.893 ** , 0.893 ** ). Tuber weight loss displayed strong negative correlations with shoot number ( r = − 0.876 ** ), tuber firmness after storage ( r = − 0.825 ** ), and TSS after storage ( r = 0.614 ** ). Tuber firmness after storage was positively correlated with plant height ( r = 0.763 ** ), leaf area ( r = 0.764 ** ), and tuber Boron before storage ( r = 0.876 ** ). TSS before storage and TSS after storage were highly intercorrelated ( r = 0.977 ** ) and negatively associated with dry matter before storage and after storage, specific gravity before storage and after storage, and starch before storage and after storage ( r = − 0.789 ** to − 0.862 ** ). Plant height showed a high positive correlation with total tubers per plant ( r = 0.879 ** ) and dry matter before storage ( r = 0.867 ** ), while shoot number was strongly linked to tuber Zinc before storage ( r = 0.872 ** ) and tuber Boron before storage ( r = 0.861 ** ) Fig. 6 . Fig. 6. Open in a new tab The graph of Pearson’s correlation Estimation for different studied variables of Potato in response to different Zinc Sulphate and Boric Acid levels. BS: before storage, AS: after storage. Regressions between traits Stepwise regression analysis (Table 2 ) identified application of tuber Boron concentration before storage (BS) as the strongest single predictor of total yield (ton/ha), with a slope of 1.84 ( p < 0.01), explaining 86.0% of the variance (R² = 0.86, adjusted R² = 0.854). Leaf area emerged as the second most influential predictor, yielding a slope of 1.32 ( p < 0.01) and accounting for 77.0% of yield variation (R² = 0.77, adjusted R² = 0.758). It provided robust prediction with moderate error. Dry matter (BS) was a significant contributor (slope = 5.78, p < 0.01), explaining 68.8% of the variance (R² = 0.69, adjusted R² = 0.675) with RMSE = 7.726 and MAE = 6.139. Total tubers per plant showed the lowest explanatory power among significant predictors (slope = 5.45, p < 0.01; R² = 0.62, adjusted R² = 0.604), with the highest prediction errors (RMSE = 8.532, MAE = 6.913). Collectively, the results underscored tuber Boron concentration before storage as the dominant driver of total yield, followed by leaf area, with dry matter and tuber number offering supplementary but less precise predictive value. Table 2. The model summary of stepwise regression for total yield for dependent and predictor variables. Model Slope P value Sig. R2 Adjusted R 2 RMSE MAE Total yield d and Leaf area p 1.32 0.000 ** 0.77 0.758 6.669 5.788 Total yield d and tubers/Plant p 5.45 0.000 ** 0.62 0.604 8.532 6.913 Total yield d and Dry matter (BS) p 5.78 0.000 ** 0.69 0.675 7.726 6.139 Total yield d and Tuber Boron (BS) p 1.84 0.000 ** 0.86 0.854 5.184 4.301 Open in a new tab D dependent variable, p predictor, BS before storage. A strong positive relationship ( p < 0.001) between tuber Boron concentration (mg L⁻¹) and total yield (ton/ha) was observed. The regression formula was y = 12.661 + 1.038x and R² was 0.859 and 85.9% of the variability in total yield is accounted for by Boric Acid levels. The association between dry matter before storage (%) and total yield (ton/ha) was significant and positive ( p < 0.01). The regression relation was y = -5.5899 + 5.777x and R² = 0.688, explaining 68.8% of the yield variance. The linear fit was solid, with residuals generally under 12 ton/ha and no apparent patterns, supporting the validity of the regression assumptions. The number of total tubers per plant (n) displayed a positive correlation ( p < 0.05) with total yield (ton/ha), so that the regressions formula was y = -1.724 + 5.447x and adjusted R² = 0.619, capturing 61.9% of the variation. Leaf area (cm²) was positively and significantly ( p < 0.01) correlated to the total yield (ton/ha) with formula as y = -3.343 + 1.32x and R² = 0.767, which accounted 76.7% of yield variability Fig. 7 . Fig. 7. Open in a new tab The linear regression for total tuber yield (dependent variable) and predictors: tuber B content ( a ), tuber dry matter ( b ), the numbers of tuber per plant ( c ), and leaf area ( d ) of potato in response to different Zinc Sulphate + Boric Acid levels. Discussion Our findings demonstrated that combined Zinc Sulphate and Boric Acid application exerts a synergistic effect on vegetative growth, tuber yield, and tuber quality, and the 1000 mg/L Zinc Sulphate + 100 mg/L Boric Acid treatment produced more optimal physiological and yield responses. Zinc is an essential micronutrient involved in the plant physiological processes, including the activation of more than 300 enzymes, the synthesis of chlorophyll, auxin metabolism (via tryptophan), carbohydrate metabolism and maintenance of membrane integrity 48 – 51 . Furthermore, Zinc is required for many physiochemical-biochemical processes in plants and can play a role in plant growth and development. It has already been confirmed that the activity of various ZIP family members is governed by transcription factors from the basic leucine zipper (bZIP) family, which are linked to the Zn deficiency response element (ZDRE) of many genes that are prone to Zn deficiency 50 . In addition, the transcriptional factors such Zinc finger proteins (ZFPs) are involved in plant growth regulation and responses to biotic and abiotic stresses, which may have contributed to the growth and quality of potato plants in the present study 49 . Furthermore, Zn upsurges water uptake and leaf relative water content through up-regulation of the expression of aquaporin genes (MdPIP1;1, MdPIP1;2). Also, Zn promoted the transport of photosynthetic products (sucrose, sorbitol) from the leaves to roots. In a study, regarding N metabolism, the activities of NR, NiR, GS, and GOGAT in Zn-treated plants were significantly higher than those in non-treated plants. Zn-treated plants also exhibited significantly higher expression of nitrate transporter genes (MdNRT1.1, MdNRT2.4) and NO3 − ion influx flow rate in root 30 . All these potential physio–biochemical evidences confirm our findings that Zinc application helps improve potato quality and quantity. In line with our findings, foliar application of a mixture of Zn and B (3 kg Zn/ha + 2 kg B/ha) significantly improved potato growth, sizes and number of tuber, and total yield compared to control and separate application of Zn or B 52 . Also, application of Zinc Sulphate and Boron at 25 and 5 kg/ha significantly has improved the potato tuber yield (36.5 t/ha) and Zinc and Boron content of tubers 53 . In a study, the combination of B and Zn has increased plant height (33.5%), number of aboveground stems (112.6%), leaf area (60.3%), tuber yield (128.7%), in pre–harvest period and dry matter (18%) and starch percentage (57.7%) after storage 44 . Also, foliar application of 60 mg L − 1 Zinc caused uppermost height of plants (41.13 cm), branches per plant (3.85), and leaf area (1337.6 cm 2 plant − 1 ), in leaves of faba bean (116.58 mg kg − 1 dry matter) 54 . Application of 50 and 100 ppm ZnO NPs enhanced growth fruit yield of eggplant by 12.2% and 22.6%, respectively, compared with control 55 . Foliar spraying of 0.1 mg/L B facilitated fruit ripening of tomato, and the 0.5 mg/L B increased the chlorophyll content of leaves by 25% 56 . Boron 0.04% significantly has enhanced large tubers plant − 1 (2.1), total yield (18.7 t ha − 1 ), survival (75%), dry matter content (18.2%) and starch percentage of potato (14.9%) compared to control 57 . In additions, foliar application of (0.06% Ca) + (0.02% B) resulted in the supreme total yield (20.5 t ha − 1 ), dry matter content (19.5%) and starch percentage (16.2%) 57 . Mahmoud et al. (2020) reported that the nano-form of Zn and B significantly has increased plant height, shoot dry weight, number of stems per plant, chlorophyll content, and potato tuber yield compared to control 1 . Furthermore, soil application of nano Zn and B has increased the concentration of nutrients (N, P, K, Ca, Zn, and B), protein, carbohydrates, and antioxidant enzymes in tubers, and leaf gibberellic acid hormone (GA3) 1 . In another study, 50 mg L − 1 Nano-Boron (foliar application) has enhanced the tuber yield and 50 and 100 mg L − 1 has boosted N and B contents of tuber 14 . The applying Boron at 30 mg l − 1 significantly has increased potato tuber firmness, tuber dry weight, starch, sugars, TSS, and ascorbic acid, and specific gravity compared to the control 58 . In a study by Lerna et al. (2017), the micronutrients fertilization improved quality characteristics of potato tuber including firmness (+ 17%), TS (+ 16%), TSS (+ 12%), sugars (+ 12%), and ascorbic acid contents (+ 40%) 13 . Another study conducted by this author revealed micronutrients fertilization improves that mineral composition like Fe (+ 70%), Zn (+ 27%), N (+ 23%), and Mn (+ 18%) in potato tubers 59 . Lenka & Das (2019) reported that application of both Zinc and Boron improved the growth and yield of potato significantly, while Zinc and Boron acted synergistically in increasing the tuber yield, so the highest total tuber yield (30.12 t/ha) has recorded with foliar application of 0.1% Boron + 0.1% Zinc along with recommended dose of NPK 60 . As the other examples, the foliar application nano-Boron (50 mg L − 1 ) has improved the total tuber yield, and shoot and tuber N and B contents 14 . Furthermore application of B 100 mg/ L + Se 5 mg/L significantly has increased the plant growth and yield of potato, plant height, tuber dry weight, number of tubers/plant, tuber yield/plant, tuber yield/feddan, tuber TSS %, ascorbic acid, tubers sugars, and nutrient content of tubers 61 . The improvements in growth, yield and potato quality in present experiment can be attributed to the activity of Boron to enhance cell division and cell elongation, carbohydrate transport, and membrane and wall stability in cells 62 – 64 and also, Zinc in auxin biosynthesis, enzyme activation, metabolic regulation, and cell wall integrity 1 , 18 , 19 . Conclusion We found that Zinc Sulphate at 500 or 1000 mg/L and Boric Acid at 50 or100 mg/L, separately or in combination, improved potato morpho-physiological attributes, tuber yield and quality before and after storage stages, however 1000 mg/L Zinc Sulphate in combination with 100 mg/L Boric Acid was the more effective treatment, so that it caused the highest tuber yield, dry matter content, micronutrient contains, specific gravity, starch content, firmness, and Zn and B contents. Furthermore, the minimum TSS value, and the least amount of weight lost during the period of storage were observed in tubers which treated with this combination. These improvements can be attributed to the activity of Boron to enhance cell division and cell elongation, carbohydrate transport, and membrane and wall stability in cells and Zinc in auxin biosynthesis, enzyme activation, metabolic regulation, and cell wall integrity. Overall, present results demonstrated that the combined application of Boron and Zinc, especially a combination of 1000 mg/L Zinc Sulphate and 100 mg/L Boric Acid, enhances tuber quality, storage behavior, and textural strength, resulting the favorable postharvest characteristics. Thus, we recommend the application of 1000 mg/L Zinc Sulphate + 100 mg/L Boric Acid to improve yield and quality of potato tubers. Author contributions Badran S. Agha, Muhamad A Ahmad, Esraa Abd-Alhuseein Jasim and Ammar Z. A. Kassab Bashi performed the experiment, contributed to data collection. Heidar Meftahizade analyzed the data and wrote the manuscript. All authors read and approved the final version of the manuscript. Data availability Data is provided within the manuscript. Declarations Competing interests The authors declare no competing interests. Footnotes Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. 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