Evaluating morpho-physio-biochemical and yield performance of six commercial potato cultivars under a semi-arid agroecosystem.

Evaluating morpho-physio-biochemical and yield performance of six commercial potato cultivars under a semi-arid agroecosystem - PMC Skip to main content An official website of the United States government Here's how you know Here's how you know Official websites use .gov A .gov website belongs to an official government organization in the United States. Secure .gov websites use HTTPS A lock ( Lock Locked padlock icon ) or https:// means you've safely connected to the .gov website. Share sensitive information only on official, secure websites. Search Log in Dashboard Publications Account settings Log out Search… Search NCBI Primary site navigation Search Logged in as: Dashboard Publications Account settings Log in Search PMC Full-Text Archive Search in PMC Journal List User Guide PERMALINK Copy As a library, NLM provides access to scientific literature. 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Learn more: PMC Disclaimer | PMC Copyright Notice Sci Rep . 2026 Mar 4;16:12122. doi: 10.1038/s41598-026-41139-5 Search in PMC Search in PubMed View in NLM Catalog Add to search Evaluating morpho-physio-biochemical and yield performance of six commercial potato cultivars under a semi-arid agroecosystem Syed Mohsin Abbas Syed Mohsin Abbas 1 Department of Horticulture, Faculty of Agricultural Sciences, University of the Punjab, Lahore, 54590 Pakistan Find articles by Syed Mohsin Abbas 1 , Raina Ijaz Raina Ijaz 2 Department of Horticulture, Faculty of Agriculture, University of Poonch Rawalakot, Rawalakot, 12350 Pakistan Find articles by Raina Ijaz 2 , Muhammad Nafees Muhammad Nafees 3 Department of Horticultural Sciences, Faculty of Agriculture and Environment, The Islamia University of Bahawalpur, Bahawalpur, 63100 Pakistan Find articles by Muhammad Nafees 3 , Tanveer Hussain Tanveer Hussain 4 Department of Horticulture, PMAS-Arid Agriculture University, Rawalpindi, 46300 Pakistan Find articles by Tanveer Hussain 4 , Muhammad Tahir Akram Muhammad Tahir Akram 4 Department of Horticulture, PMAS-Arid Agriculture University, Rawalpindi, 46300 Pakistan Find articles by Muhammad Tahir Akram 4 , Alina-Stefania Stanciu Alina-Stefania Stanciu 5 Department of Agriculture-Horticulture, Faculty of Environmental Protection, University of Oradea, Oradea, 3700 Romania Find articles by Alina-Stefania Stanciu 5 , Muhammad Wasim Haider Muhammad Wasim Haider 3 Department of Horticultural Sciences, Faculty of Agriculture and Environment, The Islamia University of Bahawalpur, Bahawalpur, 63100 Pakistan Find articles by Muhammad Wasim Haider 3, ✉ , Paul Nampala Paul Nampala 6 School of Agricultural and Applied Sciences, Bugema University, Kampala, 6529 Uganda Find articles by Paul Nampala 6, ✉ Author information Article notes Copyright and License information 1 Department of Horticulture, Faculty of Agricultural Sciences, University of the Punjab, Lahore, 54590 Pakistan 2 Department of Horticulture, Faculty of Agriculture, University of Poonch Rawalakot, Rawalakot, 12350 Pakistan 3 Department of Horticultural Sciences, Faculty of Agriculture and Environment, The Islamia University of Bahawalpur, Bahawalpur, 63100 Pakistan 4 Department of Horticulture, PMAS-Arid Agriculture University, Rawalpindi, 46300 Pakistan 5 Department of Agriculture-Horticulture, Faculty of Environmental Protection, University of Oradea, Oradea, 3700 Romania 6 School of Agricultural and Applied Sciences, Bugema University, Kampala, 6529 Uganda ✉ Corresponding author. Received 2025 Sep 22; Accepted 2026 Feb 18; 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: PMC13076650  PMID: 41781439 Abstract The potato production in semi-arid regions is often hampered by extreme temperatures, high solar radiation, and low soil fertility, which reduce photosynthetic performance and nutrient absorption. Therefore, it is crucial to identify cultivars that can tolerate such conditions without compromising their yield for sustainable production. To address this challenge, a two-year field trial (2023–2025) was conducted at the Horticulture Experimental Area of the Islamia University of Bahawalpur, Pakistan. Six commercial potato cultivars, including Sante, Musica, Sadaf, Lady Rosetta, Berna, and Kuroda, were evaluated in a randomized complete block design with four replications. The experimental site was characterized by sandy loam soil and moderate salinity in irrigation water. Measurements were taken on morphological, chlorophyll fluorescence, and nutrient uptake attributes and subjected to ANOVA to determine significant genotypic differences ( p ≤ 0.05). Results revealed that cultivar Sadaf produced 25–80% higher tuber yield, and 5–30% greater nitrogen, while 20–60% greater phosphorus-uptake efficiency compared to the rest of cultivars. Musica also demonstrated a stable performance; however, Sante and Berna were constrained in physiological adaptability and nutrient-uptake efficiency. Principal component and cluster analyses confirmed these patterns, grouping Sadaf and Musica with favorable traits and placing Sante and Berna with stress-linked parameters. The improved performance in Sadaf might be linked with enhanced PSII quantum yield, proton flux, and chlorophyll content, indicating improved light energy utilization and nutrient assimilation. The study highlights developing climate-resilient and input-efficient cultivars for sustainable cultivation in semi-arid ecosystems. Supplementary Information The online version contains supplementary material available at 10.1038/s41598-026-41139-5. Keywords: Cultivars evaluation, Nutrient uptake, Solanum tuberosum L., Stress tolerance, Sustainable cultivation Subject terms: Physiology, Plant sciences Introduction Potato ( Solanum tuberosum L.) ranks among the most vital food crops globally, standing as the third most consumed and the fourth most produced staple after maize, rice, and wheat 1 . It is renowned for its high carbohydrate content and balanced nutritional profile, which contribute significantly to caloric intake, mineral availability, and global food security 2 . It has been adapted across diverse agroecological zones, from temperate to subtropical and even semi-arid regions, far beyond its conventional growing belts 3 . Nonetheless, the average yield of potatoes is comparatively less in semi-arid areas than in temperate areas (USA, France, and Germany), where high-input agriculture attains average tuber yields of 45 tons ha –1 4 . The low production in semi-arid areas is mainly due to a lack of suitable germplasm capable of withstanding abiotic stresses such as temperatures over 25 °C at critical growth stage such as tuber initiation, irradiance beyond 2000 µmol m –2 s –1 , and salinity above 900 mg kg –1 , with low soil fertility prevailing in these environments 5 , 6 . Such harsh environmental circumstances are devastating to the onset of tuber formation, limit photosynthetic rates, and alter the growth physiology of potato plants 7 . The extreme air temperatures and sunlight intensities, particularly at tuber development, can lead to photoinhibition of photosystem (PS) II, which is relatively thermo- and photo-sensitive than PSI 8 – 10 . This photoinhibition impairs the effective N and P absorption by interfering with the plant metabolism, resulting in an imbalance of metabolic processes of the plant and stunted growth 11 . In semi-arid agroecosystems, potato cultivation is constrained not only by temperatures and water scarcity but also by intense solar radiation, saline irrigation, and poor nutrient availability, all of which collectively disrupt plant metabolism and tuber development 2 . The altering temperatures during day and night often impair photosystem II (PSII) stability, reducing chlorophyll efficiency and limiting carbon fixation 12 . Similarly, low soil fertility and nutrient immobilization hinder nitrogen and phosphorus uptake for leaf expansion, enzyme activity, and tuber bulking 13 , 14 . Therefore, traits such as chlorophyll fluorescence stability, leaf thermal regulation, and nutrient uptake efficiency are directly linked to a cultivar’s capacity to withstand these conditions. Evaluating such morpho-physio-biochemical characteristics provides a comprehensive framework for identifying resilient and input-efficient genotypes capable of sustaining yield under semi-arid production systems. Plants exhibit various physiological responses, such as nutrient acquisition efficiency, particularly macronutrients like N and P, which play a pivotal role in defining yield and stress adaptation 15 , 16 . The efficacy of plant nutrient uptake varies across cultivars, necessitating the cultivar screening for a robust PSII system and improved nutrient uptake under such stress environments 17 – 19 . This is especially remarkable when considering the framework of a climate-smart agricultural system, where high-performing cultivars under abiotic stress, on a regional scale, can lead to the resilience of global food supplies 20 . Nitrogen is involved in chlorophyll and protein synthesis, and energy transduction, whereas phosphorus is vital for root growth, ATP formation, and signal transduction 14 , 21 , 22 . Plant N and P uptake becomes even more important under stressed environments since nutrient solubility and bioavailability are frequently affected 23 , 24 . Genotypic variability in nutrient absorption efficiency and its use in semiarid environments can significantly affect the rate of potato crop growth and tuber yield 25 . Therefore, the evaluation of morpho-physiological characteristics and nutrient uptake performance can serve as an integrated approach to cultivar screening. Despite the extensive literature that has been published on the adaptability and yield maximization of potato, with a comparatively small number of studies focusing on the concomitant assessment of morpho-physiological and nutritional dynamics of commercial potato cultivars in high temperature and irradiance 26 – 29 . Also, the breeding strategies of the modern world fail to consider such complex interactions, concentrating on either yield or resistance to stress 30 , 31 . The global introduction of potato production in non-traditional areas and regions with unfavorable climatic conditions has resulted in the need to identify cultivars capable of supporting high production with improved physiological response and nutrient uptake efficiency 32 . This study advances existing potato stress-adaptation research by integrating morphological, chlorophyll fluorescence, and nutrient-uptake traits within two-season field experiments under semi-arid conditions. The previous literature predominately concentrated on individual or independent responses to stress, our approach enables a more holistic understanding of the relationship between physiological efficiency and nutrient acquisition that contributes in yield stability in cultivars under the semi-arid conditions characterized by mean daily temperatures ranging from 13 to 26 °C and average sunshine duration of 7–9 h day –1 during the cropping period. We hypothesized that cultivars exhibiting greater PSII efficiency and nutrient-uptake efficiency would sustain higher yield and adaptability. The specific objectives included: (i) To assess the genotypic variability in morphological, fluorescence, and biochemical traits under semi-arid stress; (ii) To determine nitrogen and phosphorus uptake and uptake efficiencies; and. (iii) To identify cultivars with integrated morpho-physio-biochemical resilience for sustainable production in resource-limited environments. Materials and methods Acquisition of seed tubers The research utilized six commercial cultivars, including Sante, Musica, Sadaf, Lady Rosetta, Berna, and Kuroda (Fig. 1 ), acquired from the Potato Research Institute, Sahiwal, Pakistan. A detailed description of these cultivars, along with parentage and release date, is provided in Table S1 . Fig. 1. Open in a new tab Vegetative growth pattern visually captured in six commercial potato cultivars grown under field conditions. Description of experimental location and site The field experiments were executed at the Horticulture Experimental Area, Department of Horticultural Sciences, Faculty of Agriculture and Environment, The Islamia University of Bahawalpur, Pakistan (29°22′18.4″ N, 72°45′57.6″ E, 214 m above sea level) across two successive growing seasons (2023–24 and 2024–25). The region experiences a subtropical climate. Meteorological data were recorded throughout the trial period. The site’s soil is characterized by salinity and alkalinity issues, while the underlying groundwater is brackish, containing salt concentrations exceeding 900 mg L –1 2 , 33 . Before initiating the experiment, composite soil samples were taken from 0 to 15 cm and 16–30 cm, following the method outlined by Ryan et al. 34 . Both layers of soil were characterized for texture and electrical conductivity 2 , pH 35 , organic matter 36 , available phosphorus 2 , nitrogen 37 , and potassium content 38 . Climatic trends during the experimental period The mean temperature gradually decreased from October (28.3 °C) to January (12.5 °C) during 2023–24 (Fig. 2 ). However, the temperatures above 25 °C were recorded frequently during the tuber initiation stage. Relative humidity fluctuated moderately (Fig. 2 ). Rainfall remained low throughout the season, with the minimum (2.2 mm) recorded in November and slight increases observed in December (4.8 mm) and January (5.1 mm; Fig. 2 ). Sunshine duration showed minor variation, ranging from approximately 8.7 h per day in October to 7.2 h per day in January (Fig. 2 ). A similar trend was evident in the 2024–25 period, where temperature again showed a gradual decline from October (27.9 °C) to January (12.8 °C; Fig. 2 ). Relative humidity followed a comparable pattern, with higher values in October (50%) and December (49%) and a noticeable decrease in November (43%; Fig. 2 ). Rainfall remained low and relatively stable across the months, while sunshine duration exhibited only slight fluctuations, maintaining a consistent pattern with the previous season (Fig. 2 ). Fig. 2. Open in a new tab Monthly variations in relative humidity (R.H; %), temperature (°C), rainfall (mm), and sunshine duration (hr) recorded during experimental duration (2023–2024 and 2024–2025) at the experimental site. Soil physicochemical properties The soil from the two studied depths (0–15 cm and 16–30 cm) was alkaline in nature, with pH values ranging from 8.0 to 8.3 and electrical conductivity between 1.55 and 1.72 dS m –1 (Table 1 ). The soil texture was sandy loam, as it contained 55–58% sand, 14–19% clay, and 26–28% silt (Table 1 ). Organic matter content was relatively low, varying from 0.75 to 0.81%, while the cation exchange capacity ranged between 7.0 and 7.7 cmol kg –1 (Table 1 ). The concentration of total dissolved salts was between 0.65 and 0.79 mg g –1 (Table 1 ). Nutrient analysis indicated that nitrogen content was 0.040–0.051%, phosphorus ranged from 8.5 to 11.0 mg kg –1 , and potassium was 124–150 mg kg –1 (Table 1 ). Table 1. Physicochemical properties of soil during 2023–2025. Particular Mean values obtained for both soil layers 0–15 cm 16–30 cm Silt 26% 28% Sand 55% 58% Clay 19% 14% Soil texture Sandy loam Sandy loam Total dissolved salts 0.65 mg g –1 0.79 mg g –1 pH 8.0 8.3 EC 1.55 dS m –1 1.72 dS m –1 Saturation 34% 31% OC 0.81% 0.75% CEC 7.7 c mol kg –1 7.0 c mol kg –1 Available phosphorus content 8.5 mg kg –1 11.0 mg kg –1 Available potassium content 150 mg kg –1 124 mg kg –1 Total nitrogen content 0.051% 0.040% Open in a new tab Physicochemical analysis of irrigated water The irrigation water used during the 2023–24 and 2024–25 seasons exhibited about 7.5 pH with an electrical conductivity of 1.25 dS m –1 , suggesting moderate salinity within the acceptable range for crop cultivation (Table 2 ). The total dissolved salts content was 1.4 g L –1 , which falls under safe limits for irrigation purposes. The combined calcium and magnesium ions (Ca 2+ + Mg 2+ ) were found to be 7.4 meq L –1 , while sodium concentration was 4.9 meq L –1 (Table 2 ). Bicarbonates (HCO 3 – ) were 2.7 meq L –1 , whereas chloride ions (Cl – ) were comparatively low at 1.1 meq L –1 , with no detectable carbonate (CO₃²⁻; Table 2 ). The sodium adsorption ratio was 2.5, indicating minimal risk of sodicity-related soil problems (Table 2 ). Residual sodium carbonate was observed at 0.31 meq L –1 , indicating that the water may impair soil permeability or structure (Table 2 ). Table 2. Irrigation water quality during 2023–2025. Particular Mean value pH 7.5 EC 1.25 dS m –1 Total dissolved salts 1.4 g L –1 Ca + Mg 7.4 Meq L –1 Na 4.9 Meq L –1 CO 3 – HCO 3 2.7 Meq L –1 Cl 1.1 Meq L –1 Sodium adsorption ratio 2.5 Residual sodium carbonate 0.31 Meq L –1 Open in a new tab Experimental design and crop management Potato crops were cultivated during two consecutive seasons (2023–2024 and 2024–2025). Planting was done in October, and harvesting was done in January for both years. Uniform, healthy tubers weighing 65 ± 5 g were manually planted at a depth of 10 cm and spaced 15 cm apart in ridges prepared with a tractor-mounted ridger, maintaining a 75 cm distance between rows. In the first season, sowing was completed on 4 October 2023, and harvesting on 25 January 2024, following haulm killing with “Desica” (Syngenta Pvt. Ltd.). In the second season, planting was completed on October 2, 2024, with harvesting on January 28, 2025. Each experimental plot measured 14.6 m 2 and consisted of two rows, each containing 60 tubers, for a total of 120 tubers per cultivar. The crop was fertilized at a rate of 185 kg P ha –1 , 250 kg P 2 O 5 ha –1 , and 125 kg K 2 O ha –1 , following regional recommendations for potato production. The application of P and N was made in the form of diammonium phosphate and urea fertilizers, while K was applied in the form of sulphate of potash taken from Fauji Fertilizer Company Ltd. The split number in this experiment was determined on the basis of the total number of irrigations (Table S2). A randomized complete block design was adopted with four repeats for each treatment. Observed attributes The morphological, fluorescent, and nutrient dynamics of six cultivars of potato were assessed from 25 randomly selected potato plants to ensure statistical robustness. Morphological attributes On the 65th day after sowing (DAS), seedling emergence percentage, plant height, number of stems plant –1 , number of branches plant –1 , leaf area index, and crop growth rate were determined. Leaf area was calculated using the method described by Haider et al. 2 , and the leaf area index was derived from leaf area covering the ground over the area of the ground. The assessment of crop growth rate was carried out using the methodology adopted by Haider et al. 4 . Yield attributes After harvesting, the number of tubers plant –1 , average tuber weight, tuber yield, marketable yield, tuber diameter, and specific gravity were recorded. Yield data recorded per plot (14.6 m 2 ) were converted to tons per hectare (t ha –1 ) based on the harvested area. The extrapolation from small plots may introduce a minor degree of estimation error due to potential border effects and spatial variability. To minimize this, uniform border plants were maintained, and yield data were collected from the central rows to ensure representativeness. The tuber specific gravity was assessed by the water displacement method (Haider et al., 2024; Haider, 2018) 2 , 3 . Fluorescence-related attributes On the 65th DAS, from 09:00 to 11:00 AM, measurements were taken on intact leaves for attributes including leaf angle, leaf temperature, leaf thickness, photosynthetically active radiation, linear electron flow, proton flux, proton conductivity, quantum yield of photosystem II, non-photochemical quenching, non-regulatory energy dissipation, chlorophyll content, and total magnitude of electrochromic shift. Data were recorded using a MultispeQ-Beta device linked with the PhotosynQ platform, Michigan State University, USA 39 . Each leaf was dark-adapted for 20 min prior to measurement to ensure the stabilization of PSII reaction centers. Nutrient uptake For the assessment of N and P uptake, 10 plants were randomly chosen from the experimental plot. The haulm biomass from each plant was harvested, weighed, and labeled accordingly. Additionally, 30 tubers were collected from the harvested section, weighed, and cut into strips of approximately 10 mm in width. Both the initial and final weights were recorded before and after oven-drying 500 g samples of tubers and haulm biomass at 70 °C for 72 h. The dried materials were ground to a fine level using a mortar and pestle to conduct N and P analysis. The concentration of P was measured through the colorimetric method as outlined by Haider et al. 2 with the help of the UV-vis spectrophotometer. The amount of N was determined by the use of the Kjeldahl method, as described by Bremner 37 . The uptake of the nutrients to haulms and tubers was determined by multiplying the dry weights of the samples by the respective concentrations of the nutrients. The total N and P uptake by the plants was calculated as the sum of the values of haulms and tubers 58 . 2 The nutrient uptake efficiency of the potato plants was computed as the ratio of the total nutrient uptake to the amount of nutrient supplied, as given in the following equation (Haider et al., 2024). 3 Data analysis Both years of data were averaged using Microsoft Excel 2016. Prior to analysis, assumptions of analysis of variance (ANOVA; normality of residuals and homogeneity of variances) were tested using the Shapiro–Wilk and Levene’s tests, respectively, and all datasets satisfied these assumptions after appropriate transformations. Where necessary, percentage data were arcsine-transformed, and count data were log-transformed to meet ANOVA assumptions. To evaluate the impact of the year factor, a preliminary two-way analysis of variance (ANOVA) of cultivars and years was performed. As the impact of year and its interaction with cultivar were not significant ( p > 0.05), both years of data were pooled, and means were subjected to final analysis. The least significant difference (LSD) test was used to compare the mean values at ( p ≤ 0.05). Principal component analysis (PCA) was performed with XLSTAT 2022.1. Results Morphological attributes Significant differences were found among potato cultivars for morphological attributes (Figs. 3 and 4 ). The highest emergence percentage (83%) was recorded in cultivar Sadaf, while the lowest (68%) was noted in Sante (Fig. 3 A). Plant height also followed a similar trend, with Sadaf attaining the greatest height (35 cm), followed by Lady Rosetta (30.3 cm), Kuroda (30), and Musica (25 cm), whereas Sante (23 cm) remained comparatively shorter (Fig. 3 B). Regarding the number of stems plant –1 , Sadaf attained the maximum number of stems (5.2), while the least stems (2.7) were observed in Sante (Fig. 3 C). The number of branches plant –1 was significantly greater (5.5) in Sadaf, closely followed by Musica (5.2), while the minimum branches were observed in Kuroda (3.7) and Sante (3.8; Fig. 3 D). A similar response was noted for leaf area index, where Sadaf exhibited the maximum value (3.6), while the lowest index (2) was found in Sante (Fig. 3 E). Likewise, the crop growth rate was highest (0.505 g m –2 day –1 ) in Sadaf, followed by Musica (0.455 g m –2 day –1 ), whereas Sante recorded the lowest values (0.215 g m –2 day –1 ; Fig. 3 F). Fig. 3. Open in a new tab Comparison of six commercial potato cultivars for emergence percentage ( A ), plant height ( B ), number of stems plant –1 ( C ), number of branches plant –1 ( D ), leaf area index ( E ), and crop growth rate ( F ). Error bars represent standard error (SE) averaged over two years, and bars sharing the same letter within each graph are not significantly different at p ≤ 0.05 according to least significant difference (LSD) test. Since no significant year effect was observed, the data were pooled across years. Sample size (n) = 6 × 4 (cultivar × replication) = 24. Fig. 4. Open in a new tab Comparison of six commercial potato cultivars for number of tubers plant –1 ( A ), average tuber weight ( B ), tuber yield ( C ), marketable yield ( D ), tuber diameter ( E ), and specific gravity ( F ). Error bars represent standard error (SE) averaged over two years, and bars sharing the same letter within each graph are not significantly different at p ≤ 0.05 according to least significant difference (LSD) test. Since no significant year effect was observed, the data were pooled across years. Sample size (n) = 6 × 4 (cultivar × replication) = 24. Yield attributes The highest number of tubers plant –1 (11.5) was recorded in Sadaf, which was significantly higher than that of Musica, while the lowest number of tubers (8) was obtained from Kuroda (Fig. 4 A). For average tuber weight, the maximum value (341.2 g) was found in Sadaf, which was at par with Musica (327.5 g), whereas Lady Rosetta (175 g) and Berna (179.5 g) produced significantly smaller tubers (Fig. 4 B). Sadaf again ranked first in terms of tuber yield (30.7 t ha –1 ), followed by Musica (24.7 t ha –1 ) and Kuroda (24.5 t ha –1 ). The minimum yield (16.8 t ha –1 ) was noted in Berna (Fig. 4 C). Marketable yield percentage was greatest in Sadaf (60.5%), statistically higher than Musica (48.1%) and Kuroda (40.3%), while Lady Rosetta displayed the lowest marketable yield (22.3%) (Fig. 4 D). Regarding tuber diameter, Sadaf produced the largest tubers (46.1 mm), closely followed by Musica (41.1 mm), whereas Lady Rosetta had the smallest diameters (26.1 mm; Fig. 4 E). Tuber specific gravity was highest in Sadaf (1.15), followed by Berna (1.1), Kuroda (1.08), and Lady Rosetta (1.07). The lowest specific gravity (0.90) was recorded in Musica (Fig. 4 F). Fluorescence-related attributes Significant variations were observed among potato cultivars for fluorescence-related attributes (Fig. 5 ). The widest leaf angle (46.5°) was recorded in Sadaf, while the narrowest (35.5°) was found in Sante (Fig. 5 A). For leaf temperature, the maximum value (31.6 °C) was noted in Sante, statistically comparable with Berna (31.1 °C), whereas the lowest (28.8 °C) was measured in Sadaf (Fig. 5 B). Leaf thickness was greatest in Sadaf (1.57 μm), significantly exceeding other cultivars, while the thinnest leaves (0.81 μm) were produced by Berna (Fig. 5 C). Linear electron flow (LEF) was highest in Sadaf (180 µmol electrons m –2 s –1 ), followed by Musica (166.5 µmol electrons m –2 s –1 ), while the lowest (155 µmol electrons m –2 s –1 ) was observed in Sante (Fig. 5 D). Photosynthetically active radiation (PAR) uptake was also maximum in Sadaf (1398.8 µmol photons m –2 s –1 ), followed by Musica (1345 µmol photons m –2 s –1 ), and minimum in Sante (1185 µmol photons m –2 s –1 ) (Fig. 5 E). For proton flux (gH + ), the highest value (275) was recorded in Sadaf, while the lowest (242.5) was found in Berna (Fig. 5 F). Proton conductivity (vH + ) followed a similar pattern, being greatest in Sadaf (0.53) and smallest in Berna (0.30) (Fig. 5 G). The quantum yield of photosystem II (Φ II ) reached its maximum in Sadaf (0.8), followed by Musica (0.75), whereas Sante (0.64) and Berna (0.67) showed significantly lower values (Fig. 5 H). Non-photochemical quenching (Φ NPQ ) was maximum in Sante (0.25), statistically at par with Berna (0.24), while the lowest (0.17) was observed in Sadaf (Fig. 5 I). Non-regulated energy dissipation (Φ NO ) was also highest in Sante (0.09), whereas the lowest (0.03) was noted in Musica (Fig. 5 J). Chlorophyll content (SPAD values) was greatest in Sadaf (48.5), followed by Musica (45), while the lowest (38) was found in Sante (Fig. 5 K). Electrochromic shift (ECSt), indicating the capacity for proton motive force, was highest in Sadaf (0.87 × 10 3 mAU) and lowest in Sante (0.62 × 10 3 mAU) (Fig. 5 L). Fig. 5. Open in a new tab Comparison of six commercial potato cultivars for leaf angle ( A ), leaf temperature ( B ), leaf thickness ( C ), linear electron flow (LEF; D ), photosynthetically active radiations (PARs; E ), proton flux through the chloroplast ATP synthase (gH + ; F ), proton conductivity of the chloroplast ATP synthase (vH + ; G ), quantum yield of photosystem II (Φ II ; H ), non-photochemical quenching (Φ NPQ ; I ), non-regulatory energy dissipation (Φ NO ; J ), chlorophyll content ( K ), electrochromic shift (ECSt; L ). Error bars represent standard error (SE) averaged over two years, and bars sharing the same letter within each graph are not significantly different at p ≤ 0.05 according to least significant difference (LSD) test. Since no significant year effect was observed, the data were pooled across years. Sample size (n) = 6 × 4 (cultivar × replication) = 24. Nutrient uptake There was also a significant difference between potato cultivars in terms of nutrient uptake (Fig. 6 ). The highest nitrogen uptake (149.8 kg ha –1 ) was recorded in cultivar Sadaf, followed by Musica (140.2 kg ha –1 ), while the lowest values (103.9 kg ha –1 ) were found in Sante (Fig. 6 A). A similar pattern was observed for nitrogen uptake efficiency (NUE), where Sadaf peaked (0.46 kg uptake kg –1 N supply), whereas Berna exhibited the lowest efficiency (0.35 kg uptake kg –1 N supply) (Fig. 6 B). In the case of phosphorus uptake, Sadaf again was found to have significantly greater P uptake (22.9 kg ha –1 ) than all other cultivars, while the lowest (12.7 kg ha –1 ) was noted in Berna (Fig. 6 C). Phosphorus uptake efficiency (PUE) also peaked (0.37 kg uptake kg –1 P supply) in Sadaf, followed by Musica (0.31 kg uptake kg –1 P supply), while the minimum efficiency (0.23 kg P uptake kg –1 P supply) was observed in Berna, comparable with Sante (0.25 kg uptake kg –1 P supply; Fig. 6 D). Fig. 6. Open in a new tab Comparison of six commercial potato cultivars for plant N uptake ( A ), nitrogen uptake efficiency (NUE; B ), plant P uptake ( C ), and P uptake efficiency (PUE; D ). Error bars represent standard error (SE) averaged over two years, and bars sharing the same letter within each graph are not significantly different at p ≤ 0.05 according to least significant difference (LSD) test. Since no significant year effect was observed, the data were pooled across years. Sample size (n) = 6 × 4 (cultivar × replication) = 24. Multivariate analysis The PCA biplot provided insight into the associations between cultivars and morpho-physio-biochemical traits (Fig. 7 ). Sadaf was strongly associated with favorable traits, including crop growth rate, tuber yield, PAR uptake, Φ II , and nutrient-uptake efficiencies, reflecting its robust physiological and biochemical adaptability (Fig. 7 ). Musica also clustered near high-performance traits, particularly those linked with photosynthetic efficiency (Fig. 7 ). In contrast, Sante and Berna grouped with parameters such as higher Φ NPQ and Φ NO , suggesting that their physiological response relied more on energy dissipation rather than efficient photochemistry (Fig. 7 ). This separation of cultivars along the first principal component indicates the contrasting adaptability strategies under semi-arid conditions (Fig. 7 ). Fig. 7. Open in a new tab Principal Component Analysis (PCA) biplot showing the distribution of six commercial potato cultivars and their association with 28 morpho-physio-biochemical traits under a semiarid agroecosystem. PC1 and PC2 explain 73.76% and 9.86% of the total variability, respectively. Chl chlorophyll content, CGR crop growth rate, vH + proton conductivity of the chloroplast ATP synthase, NUE nutrient uptake efficiency, PhiNPQ non-photochemical quenching, PUE phosphorus uptake efficiency, PhiNO non-regulatory energy dissipation, ECSt electrochromic shift, Phi2 quantum yield of photosystem II, ATW average tuber weight, gH + proton flux through the chloroplast ATP synthase, LEF linear electron flow, PARs photosynthetically active radiations. The heatmap analysis further validated these trends, with Sadaf and Musica forming a distinct cluster associated with superior morphological, physiological, and biochemical performance (Fig. 8 ). Conversely, Sante and Berna clustered together, reflecting their limited adaptability and stress-prone characteristics (Fig. 8 ). Such clustering not only confirms the individual trait analysis but also demonstrates the potential of multivariate tools to identify genotypes with complementary or divergent trait combinations (Fig. 8 ). Fig. 8. Open in a new tab Heatmap depicting the clustering pattern of six commercial potato cultivars based on 28 morphological, physiological, and biochemical traits. Chl chlorophyll content, CGR crop growth rate, vH + proton conductivity of the chloroplast ATP synthase, NUE nutrient uptake efficiency, PhiNPQ non-photochemical quenching, PUE phosphorus uptake efficiency, PhiNO non-regulatory energy dissipation, ECSt electrochromic shift, Phi2 quantum yield of photosystem II, ATW average tuber weight, gH + proton flux through the chloroplast ATP synthase, LEF linear electron flow, PARs photosynthetically active radiations. Discussion The present two-year study evaluates six commercial potato cultivars based on their adaptability under a semi-arid agroecosystem, aiming to pinpoint candidates with optimized photosynthetic systems that can thrive in regions with frost stress during autumn. Low temperature during the cropping season (Fig. 2 ), particularly at tuber initiation stage, which initiates within the 50 days after sowing 4 , can impair photosystem II (PSII) function and reduce chlorophyll content, leading to lower fluorescence yield and reduced photosynthetic efficiency 40 – 43 . The statistical comparison of six potato cultivars showed that there was significant genotypic diversity in the emergence, height of the plant, pattern of branching, and general growth rate of the crop. The results are consistent with those of Rykaczewska 44 and Demirel et al. 45 , who reported substantial variations among potato genotypes. Another study by Kleinwechter et al. 46 also reported a cultivar difference in growth and yield attributes in contrasting stress environments. Similarly, Paul et al. 47 found substantial variation in the potato cultivars in terms of carbon assimilation, antioxidant enzyme activities, and tuber yield. Furthermore, in a field study by Angmo et al. 48 , potato genotypes exhibited a significant variability for morpho-anatomical attributes under frost stress. In the present study, cultivar Sadaf consistently outperformed others, exhibiting 5–20% higher emergence, 15–65% taller plants, 5–90% greater number of stems, and 5–50% more number of branches plant –1 , and 10–80% higher leaf area index (Fig. 3 ). Sadaf also excelled in the yield-related attributes such as 5–40% higher number of tubers plant –1 , 5–95% heavier average tuber weight, and 10–75% greater tuber diameter, hence leading to 25–80% greater tuber yield and 1.3–2.7 times higher marketable percentage (Fig. 4 ). In contrast, Berna and Lady Rosetta recorded markedly lower yields, largely due to reduced tuber size and lower marketable fraction (Fig. 4 ). These findings support earlier observations that genotypes with greater vegetative vigor and expansive canopy development are better able to assimilate carbon and allocate resources to tuber bulking under challenging environments 46 , 47 . The yield-related characteristics of Sadaf and Musica indicate that they have a more effective dry matter partitioning physiological system, which is essential in semi-arid environments. On the other hand, low performance of Sante indicates its low adaptability, which is congruent with reports of genotype-dependent differences in growth and yield stability at stress-prone conditions 48 . The cultivars also differed markedly in their chlorophyll fluorescence and associated physiological traits, reflecting underlying variation in photosynthetic performance. Sadaf exhibited the widest leaf angle, lowest leaf temperature, and thickest leaves (Fig. 5 ), which collectively enhanced its photosynthetic efficiency. The positive association of leaf thickness with LEF, PAR uptake, and Φ II observed in this study aligns with earlier reports indicating that thicker leaves typically contain more chloroplasts per unit area, thereby supporting higher rates of carbon assimilation 49 , 50 . Conversely, thinner leaves, as observed in Berna, were associated with reduced photosynthetic activity and higher dissipation of absorbed energy. Among the fluorescence parameters, Sadaf consistently showed higher Φ II , PAR uptake, electron transport, and proton flux (gH + ), while maintaining lower Φ NPQ and Φ NO (Fig. 5 ). This indicates a greater ability to channel absorbed light energy into photochemistry rather than dissipating it as heat or non-regulated energy. Similar physiological patterns have been described by Fini et al. 51 and Fujii et al. 52 , yet those studies lacked combined nutrient-use data. Our results, therefore, extend existing evidence by linking PSII efficiency directly with nitrogen and phosphorus assimilation, offering a more holistic explanation of stress tolerance. The elevated SPAD values and stronger electrochromic shift in Sadaf further confirmed its robust photosynthetic apparatus, suggesting a well-developed proton motive force and superior light energy utilization 53 . On the other hand, Sante and Berna exhibited higher Φ NPQ and Φ NO , accompanied by lower chlorophyll content and Φ II , reflecting a stress-prone photosynthetic system. Such responses are often linked with photoinhibition, where excess absorbed energy cannot be effectively utilized, leading to enhanced non-photochemical dissipation 4 , 54 , 55 . The contrasting patterns among cultivars indicate the physiological basis for yield differences observed, as those with higher photosynthetic efficiency and lower energy losses (Sadaf and Musica) produced greater biomass and yield compared with Sante and Berna. The yield differences observed among cultivars can be directly attributed to their contrasting physiological and biochemical efficiencies. Cultivars such as Sadaf and Musica, which maintained higher Φ II , greater gH + , and enhanced nutrient uptake efficiencies, effectively translated these physiological advantages into higher tuber yield (Figs. 4 and 5 ). Their ability to sustain greater light energy conversion efficiency and nutrient assimilation under stress conditions likely supported more robust source–sink dynamics, ensuring continuous carbohydrate translocation toward developing tubers 56 . Conversely, Sante and Berna exhibited higher non-photochemical quenching (Φ NPQ ) and non-regulated energy dissipation (Φ NO ), indicating that a substantial portion of absorbed energy was lost as heat rather than utilized for photochemistry, resulting in weaker vegetative growth and lower yields (Figs. 4 and 5 ). These findings reinforce the concept that physiological resilience and biochemical efficiency are key determinants of productivity in stress-prone environments, providing a mechanistic basis for the yield hierarchy observed among cultivars. The highest yield performance of cultivar Sadaf can be mechanistically ascribed to its improved photosystem II functionality, as reflected by higher Φ II , LEF, gH⁺, and chlorophyll content. The strongest fluorescence parameters measured, including Φ II and LEF, had the highest association with tuber yield, since they loaded positively and close to yield, rate of crop growth and nutrient-uptake efficiency along the first dimensional component of the PCA biplot (Fig. 7 ). This multivariate analysis implies that the cultivars that were able to maintain greater photochemical energy conversion and electron transport efficiency during semi-arid stress, were more efficient in converting the absorbed light to produce biomass and tuber. In contrast, non-photochemical quenching (Φ NPQ ) and non-regulated energy dissipation (Φ NO ) loaded oppositely to yield and Φ II , particularly in stress-prone cultivars such as Sante and Berna, suggesting a greater reliance on energy dissipation mechanisms rather than productive photochemistry. There was a substantial genotypic difference in the nutrient uptake and uptake efficiency of the experimental potato cultivar, indicating their varying adaptability to the semi-arid environment (Fig. 6 ). The highest values of both nitrogen and phosphorus uptake and their respective efficiencies were invariably obtained in cultivar Sadaf, and the lowest in the cultivars Berna and Sante (Fig. 6 ). These findings emphasize the pivotal role of nutrient acquisition and utilization as determinants of growth, yield, and overall adaptability of potato cultivars under suboptimal field conditions 57 , 58 . The superiority of Sadaf in nitrogen uptake (149.8 kg ha –1 ) over all other cultivars is closely linked to its vigorous morphological development, including higher leaf area index and crop growth rate (Fig. 3 ), which together enhanced its photosynthetic capacity and assimilate production. The strong canopy of Sadaf not only enabled greater interception of light but also promoted higher carbon assimilation, thereby increasing its demand and uptake for nitrogen. Previous studies have similarly highlighted that cultivars with well-developed foliage maintain higher nitrogen uptake due to greater root growth and canopy vigor 59 , 60 . In contrast, Sante, which had weaker vegetative growth, absorbed only 103.9 kg ha –1 of nitrogen, indicating its reduced ability to sustain photosynthetic activity and biomass accumulation under water- and nutrient-limiting conditions. NUE further revealed marked genotypic differences, with Sadaf attaining the maximum value (0.46 kg uptake kg –1 N supply) and Berna the lowest (0.35 kg uptake kg –1 N supply). These differences suggest that nutrient uptake alone does not guarantee efficiency; rather, it is the effective assimilation and utilization of absorbed nitrogen that determines productivity. Efficient cultivars such as Sadaf are likely characterized by a well-developed root system for nutrient acquisition and a strong sink capacity for assimilating absorbed nitrogen into protein synthesis and tuber growth. Such variation in NUE among genotypes has been previously documented, where superior performers utilized nitrogen more efficiently to sustain higher yields in marginal environments 2 , 61 . Phosphorus is a critical element in plant metabolism, being integral to ATP synthesis, energy transfer, and signal regulation. Genotypes that can acquire greater phosphorus are better equipped to maintain metabolic stability under stress. The phosphorus uptake, in this study, also displayed a parallel trend, with Sadaf recording the highest value (22.9 kg ha –1 ), followed by Musica (19.7 kg ha –1 ), while Berna was at the bottom with 12.7 kg ha –1 (Fig. 6 C). The higher P uptake in Sadaf and Musica suggests superior root architecture and an enhanced ability to mobilize phosphorus from the rhizosphere. Conversely, the restricted P uptake in Berna and Sante could be attributed to limited root activity or reduced efficiency of phosphorus transport mechanisms. Earlier research has similarly reported substantial cultivar variation in phosphorus uptake, largely explained by differences in root traits and genotypic capacity to mobilize soil P 62 , 63 . PUE was also found to be the highest in Sadaf (0.37 kg uptake kg –1 P supply), followed by Musica (0.31 kg uptake kg –1 P supply), while Berna exhibited the lowest efficiency (0.23 kg uptake kg –1 P supply). This indicates that Sadaf not only absorbed greater quantities of phosphorus but also utilized it more effectively for biomass and yield formation. Efficient P utilization is particularly valuable in semi-arid agroecosystems, where low soil fertility and high temperature stress often limit crop productivity. Genotypes capable of converting limited phosphorus resources into higher yields are more desirable for sustainable production systems 64 . The lower PUE of Berna and Sante points out their limited adaptability, as inefficient use of phosphorus compromises their growth and productivity. The multivariate analyses provided further evidence for these observations. The PCA biplot positioned Sadaf and Musica in close association with favorable traits, including nitrogen and phosphorus uptake, NUE, and PUE (Fig. 7 ). In contrast, Sante and Berna clustered with traits indicative of stress, such as higher Φ NPQ and Φ NO , reflecting their reliance on energy dissipation rather than efficient nutrient utilization (Fig. 8 ). Similar findings have been documented in earlier studies, where nutrient uptake efficiencies were among the most influential traits explaining genotypic variability under stress-prone ecosystems 31 , 61 , 62 . This study provides a novel, integrative framework for evaluating potato cultivars under a semi-arid environment by simultaneously linking morphological, fluorescence, and nutrient-use traits with yield performance. Unlike previous research focusing on individual physiological or agronomic aspects, our findings demonstrate that combining chlorophyll fluorescence indices with nutrient-uptake efficiency traits offers a more precise understanding of stress adaptability. Future studies should expand this approach by incorporating molecular traits to identify the genetic basis of nutrient and photosynthetic efficiency. From a breeding perspective, traits such as PSII quantum efficiency, proton flux capacity, and nutrient-uptake efficiency can serve as reliable selection indices for developing climate-smart, resource-efficient potato cultivars. Multi-location trials across diverse semi-arid regions are also recommended to validate the stability of high-performing cultivars such as Sadaf and Musica for broader climate-resilient breeding and sustainable production strategies. Conclusion The evaluation of six potato cultivars under semi-arid conditions showed the importance of integration of morphological, fluorescence, and nutrient-uptake traits for identifying resilient cultivars. Sadaf and Musica proved stronger adaptability, emphasizing that future improvement programs should prioritize physiological and nutrient-use traits alongside tuber yield. Broader adoption of such cultivars can lower input dependency and enhance sustainability. Further research linking field performance with molecular mechanisms will be essential for guiding breeding and management strategies aimed at resilience in stress-prone environments. Supplementary Information Below is the link to the electronic supplementary material. Supplementary Material 1 (22.8KB, docx) Acknowledgements The authors sincerely thank the Higher Education Commission (HEC) of Pakistan for their support in providing the necessary research materials under project No. SRGP/NAHE/HEC/2020/286, entitled “Improving Potato Production through Varietal and Fertilizer Management under Cholistan Region”. The authors are also thankful to the Islamia University of Bahawalpur, Pakistan for providing space to conduct this trial under the project “Establishment of Commercial Multistorey Orchard and Fruit Research Unit” which mainly focused on maximizing sustainable fruit production per unit area. Author contributions Syed Mohsin Abbas: Conceptualization, Investigation, Methodology, Resources, Validation, Writing - original draft. Raina Ijaz: Conceptualization, Investigation, Methodology, Resources, Validation, Writing - review & editing. Muhammad Nafees: Conceptualization, Investigation, Methodology, Project administration, Validation, Writing - review & editing. Tanveer Hussain: Investigation, Validation, Writing - review & editing. Muhammad Tahir Akram: Software, Validation, Visualization, Writing - review & editing. Alina-Stefania Stanciu: Resources, Validation, Writing - review & editing. Muhammad Wasim Haider: Investigation, Resources, Project administration, Validation, Writing - review & editing. Paul Nampala: Conceptualization, Investigation, Methodology, Resources, Supervision, Validation, Writing - review & editing. Funding This research received no specific grant from any funding agency. Data availability All the data related to this work can be sourced from the corresponding authors. 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. Contributor Information Muhammad Wasim Haider, Email: wasim.haider@iub.edu.pk. Paul Nampala, Email: nampalap@yahoo.co.uk, Email: pnampala@bugemauniv.ac.ug. References 1. FAOSTAT. Food and Agriculture Organization of the United Nations database. (2023). Available at: http://faostat.fao.org 2. Haider, M. W. et al. Rejuvenating potato growth and yield in challenging semiarid and saline sandy Cholistan: Harnessing PGPB-coated N and P application strategies. BMC Plant Biol. 24 , 386. 10.1186/s12870-024-05056-x (2024). [ DOI ] [ PMC free article ] [ PubMed ] [ Google Scholar ] 3. Haider, M. W. Management strategies for breaking tuber dormancy in potato. 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