Impaginato 303 Adv. Hort. Sci., 2022 36(4): 303­314 DOI: 10.36253/ahsc­13349 Comparison of 18 Iranian caprifig culti­ vars based on some morphological and biochemical parameters B. Jamali 1 (*), H. Amin 2 1 Department of Agriculture, Minab Higher Education Center, University of Hormozgan, Bandar Abbas, Iran. 2 Department of Plant Production, College of Agriculture and Natural Resources of Darab, Shiraz University, Darab, Iran. Key words: Antioxidants, characterization, essential elements, fig, screening. Abstract: Caprifig is a valuable candidate for fig breeding programs as it typically grows naturally under non­optimal conditions. The present study was carried out to evaluate the biochemical/morphological characteristics of 18 caprifig cul­ tivars indigenous to the Darab region/southern Iran with 4 replications in a completely randomized block design. From each cultivar, healthy uniform leaf samples and spring fruits were taken and analyzed. Our results showed that ‘Naneghasem’ had the highest leaf α­tocopherol and polyphenol concentration. The highest leaf ascorbic acid concentration was obtained from the Gol Khengi cultivar (17.03 μg g­1 fresh weight). Also, the contents of chlorophyll, carotenoids, and anthocyanins were significantly different among the studied cultivars. Various cultivars had different absorption potentials for essential ele­ ments as macro and micronutrients concentration in the leaves were statistical­ ly different in various caprifigs; ‘Naneghasem’ had the highest Ca (4.46 mg g­1 dry weight) and Fe (67.71 mg kg­1 dry weight) concentration and the highest leaf K concentration (22.46 mg g­1 dry weight) was observed in ‘Mahali Layzengan’. In conclusion, ‘Naneghasem’ was evaluated as a cultivar which seems to be more morphologically­ and biochemically­distant from other studied caprifig varieties and probably more adaptable/tolerant to environmental conditions. 1. Introduction The fig, a deciduous tree of the Moraceae family and native to the southern Arabian Peninsula in the subtropical region, is an important hor­ ticultural crop that is grown for dry and fresh consumption. Dried figs are an excellent source of amino acids, vitamins, minerals, polyphenols, and crude fiber (5.8%, w/w) (Pourghayoumi et al., 2016). Satisfactory dry fig production is highly dependent on sufficient pollination. Caprifig is the only fig species that has both pistillate and staminate flowers. Growers collect the fruits of caprifigs, place them in cans, and hang the cans on Smyrna­type figs; Blastophaga psenes wasps, which live in the syconium of caprifigs, leave the profichi and enter the female flowers of the edible (*) Corresponding author: babakjamali@ymail.com Citation: JAMALI B., AMIN H., 2022 ­ Comparison of 18 Iranian caprifig cultivars based on some morpho‐ logical and biochemical parameters. ­ Adv. Hort. Sci., 36(4): 303­314. Copyright: © 2022 Jamali B., Amin H. This is an open access, peer reviewed article published by Firenze University Press (http://www.fupress.net/index.php/ahs/) and distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Data Availability Statement: All relevant data are within the paper and its Supporting Information files. Competing Interests: The authors declare no competing interests. Received for publication 5 July 2022 Accepted for publication 14 December 2022 AHS Advances in Horticultural Science https://doi.org/10.36253/ahsc-13349 http://www.fupress.net/index.php/ahs/ http://creativecommons.org/licenses/by/4.0/ http://creativecommons.org/licenses/by/4.0/ http://creativecommons.org/licenses/by/4.0/ Adv. Hort. Sci., 2022 36(4): 303­314 304 figs and pollinate their pistillate flowers (Rahemi and Jafari, 2008). In recent years, fig cultivation has developed rapidly due to the wide adaptability, early fruit and high yield, fewer pests and diseases, and simple man­ agement of this crop in various parts of the world (Zhang et al., 2020). This increases the importance of protecting and developing fig germplasm. Maintaining possible local varieties for commercial cultivation requires characterization of the available germplasm. This type of thermophilic tree could be useful to adapt to the consequences of global warming, as fig trees can grow in dry regions where many other species may not survive (Sugiura et al., 2007). Fig trees have not been subjected to extensive breeding programs; consequently, many fig populations have rich diversity that cannot be fully exploited until cor­ rectly identified (Perez­Jiménez et al., 2012). One of the first steps in breeding is the selection of promising species/cultivars. Previous studies show differences between various common fig cultivars (Aradhya et al., 2010; Perez­Jiménez et al., 2012; Zhang et al., 2020). These differences suggest varied levels of acclimation and/or tolerance potential to abiotic stress conditions. The plant’s enzymatic and non­enzymatic antioxidant potential plays a vital role in this regard (Bonyanpour and Jamali, 2020). Maintenance of high antioxidant capacity to scav­ enge toxic reactive oxygen species (ROS) has been associated with better adaptation of plants to envi­ ronmental conditions (Sharma et al., 2012). In addi­ tion, different cultivars have different abilities to uptake nutrients. The concentration of macro­ and micronutrients in plant organs and tissues correlates with the plant’s potential to tolerate prevailing envi­ ronmental conditions (Jamali et al., 2016; Jamali and Bonyanpour, 2017; Tian et al., 2021). Iran has one of the richest fig germplasm resources in the world; however, the majority of pre­ vious studies have focused on edible commercial cul­ tivars, while studies on caprifig varieties are rare. Caprifig can be a valuable resource for breeding pro­ grams because it grows naturally under less than optimal conditions and is not domesticated. Strengthening research on caprifig germplasm’s genetic/chemical diversity is of scientific importance for germplasm conservation, efficient breeding, and satisfactory production. Typically, cultivar characterization studies involve determining a vast range of morphological, biochemi­ cal, molecular, etc., parameters to find promising varieties for future breeding projects. As part of a series of similar investigations on caprifig cultivars, the present study aimed to compare 18 cultivars indigenous to the Darab/Southern Iran region, focus­ ing on phonological parameters and biochemical characteristics associated with non­optimal growing conditions. 2. Materials and Methods Plant material Uniform and healthy plants of eighteen mature autochthonous caprifig cultivars were selected. They were planted and grown in a completely randomized block design with 4 replications; each replication had 4 plants with a row spacing of 2.5 m and an inter­row spacing of 3.5 m in a rain­fed collection orchard in the Darab region, Fars province, southern Iran. Average annual climate parameters in the experi­ mental region were: precipitation: 200 mm, relative humidity Max: 55%, Min: 18%, temperature Max: 42°C, Min: 4°C. The soil of the orchard was sampled and analyzed for soil texture, mineral content, organ­ ic matter, pH, and EC (Table 1). Cultivars included: 1­ Mashgholamrezai (MGR), 2­Darabi (DRB), 3­Abbasali (ASA), 4­Khazraie (KZR), 5­Ieji (IJ), 6­Gol Khengi (GK), 7­Rastaghi (RS), 8­Naneghasem (NGM), 9­Mahali Layzengan (MLZ), 10­Cheshmbolboli (CMB), 11­ Pouzehdonbali (PZD), 12­Danesibi (DS), 13­Pasras (PR), 14­Johari (JR), 15­Maseeh (MS), 16­Cho (CO), 17­Pasbehdari (PSD), and 18­Suzu (SZ). Leaf samples were taken from different orientations (north, south, west, and east) of the trees; 25 fully expanded mature leaves from each side of all trees (100 leaves per tree as bulk samples) were transported to the Table 1 ­ Analysis of soil samples in the experimental region Soil depth (cm) Soil texture Soil mineral content (mg kg­1) Organic carbon (%) EC (dS m­1) Nitrate Ca Mg K Fe Zn Mn 0­50 Loamy clay 37 1110 170 220 7.00 1.7 5.50 0.70 0.72 Jamali and Amin ‐ Characterization of caprifig cultivars 305 laboratory. Leaves were of spring bloom, the middle third of the branch, at the height of 1­1.5 m, includ­ ing the petiole. Leaves with abnormal symptoms, including mechanical lesions caused by diseases or pests as well as chlorosis, were avoided. Routine cul­ tural practices were carried out during the experi­ mental period. Measurements The following parameters were determined in caprifig cultivars for two consecutive years, with an average reported. Leaf dry matter content Three uniform leaves were washed with tap and distilled water. After drying with a clean towel, they were weighed with a digital scale to get fresh weight (FW) and then dried in an oven at 70°C for 72 hours and re­weighed to get dry weight (DW). The percent­ age of leaf dry matter was calculated according to the following formula: Leaf dry matters (%) = [Leaf DW (g)/leaf FW (g)] × 100 Leaf relative water content (LRWC) Ten leaf discs from each treatment were weighed (FW), then hydrated to saturation (constant weight) for 48 hours at 5°C in darkness (turgid weight, TW). Leaf discs were dried in an oven (DW). The relative water content was calculated according to the fol­ lowing formula (Jamali and Eshghi, 2015): LRWC (%) = (FW­DW)/(TW­DW) × 100 Leaf electrolyte leakage Leaf electrolyte leakage (EL) was determined by recording leaf leachate’s electrical conductivity (EC) in double­distilled water at 40 and 100°C. Leaf sam­ ples were cut into uniform­sized disks and placed in test tubes containing 10 ml of double­distilled water. The test tubes were kept at 40°C for 30 minutes and at 100°C for 15 minutes, and their respective electri­ cal conductivities (EC1 and EC2) were measured using a conductivity­meter (METROHM Conductometer 644, Switzerland): Electrolyte leakage (%) = (EC1/EC2) × 100 Leaf soluble carbohydrates Leaf samples of 0.5 g (DW) were homogenized in 5 ml ethanol (95%) and centrifuged at 4500 × g for 15 min. The supernatant was removed from the sample, and the residue was resuspended in 5 ml ethanol (70%). The supernatant was centrifuged for final extraction. Both supernatants were combined. The anthrone sulfuric acid assay was used for determina­ tion (Irigoyen et al., 1992). An aliquot of 100 μl was added to 3 ml of the anthrone­sulfuric acid solution; the mixture was shaken, heated in a boiling water bath for 10 min, then cooled at 4°C. The absorbance at 625 nm was determined spectrophotometrically. Glucose (0­100 mg/l, Merk) was used as a standard (Jamali and Bonyanpour, 2017). Leaf chlorophyll and carotenoids concentration Leaf discs of 0.25 g were extracted in 2.5 ml of acetone (80%) and then centrifuged at 6,000 × g for 10 min. The supernatant was used to prepare a final volume of 50 ml of leaf extract. Extraction of the leaf tissue with the buffer was continued until decol­ oration. The absorbance of the extract was measured at 470, 645, and 663 nm spectrophotometrically. Acetone (80%) was used as a blank. Finally, the con­ tent of chlorophyll and carotenoids was calculated using the following equations (Lichtenthaler, 1987): Leaf total chlorophyll concentration (mg. g­1 FW)= [(7.15A663 + 18.71A645) × v /1000 × W] Carotenoids (mg. g­1 FW): 1000A470­ 1.82Chla­ 85.02Chlb / 198 A = absorbance at ƛ (nm), W = sample weight. Leaf anthocyanins concentration Total anthocyanins in the leaves were determined by the pH differential method using a spectropho­ tometer and two buffer systems: potassium chloride buffer, pH 1.0 (0.025 M), and sodium acetate buffer, pH 4.5 (0.4 M). 0.25 g leaf samples were extracted with 1 ml methanol: water: concentrated HCl solu­ tion (80:20:1 v/v/v). 0.4 ml of leaf extract was mixed with 3.6 ml of corresponding buffers and read against water as blank at 510 and 700 nm. Absorbance (A) was calculated as A = (A515 ­ A700) pH 1.0 ­ (A510 ­ A700) pH 4.5 Then total leaf anthocyanins concentration was cal­ culated using the following equation: Anthocyanin (μg. g­1 FW) = (A × Mw × DF × 1000) / e Where A is the absorbance of the diluted sample, DF is the dilution factor (10), Mw is the molecular weight of cyanidin­3­glucoside (449.2), and e = 26,900 L/mol.cm, the molar extinction coefficient of cyani­ din­3­ glucoside. Leaf α‐tocopherol concentration The α­tocopherol was extracted using Chong et al. (2004) method. 200 mg lyophilized sample was homogenized in 1 ml acetone at 4°C with a pre­ chilled mortar and pestle. 0.5 ml hexane was added, and the homogenate was first vortexed for 30 s, cen­ trifuged at 1000 × g for 10 min. The upper hexane Adv. Hort. Sci., 2022 36(4): 303­314 306 layer was removed while the acetone layer contain­ ing α­tocopherol remained in the vial. A second aliquot of 0.5 ml hexane was added, and the extrac­ tion process was repeated three times. α­tocopherol was determined by the method of Kanno and Yamauchi (1997). To 0.2 ml of pooled extract a 0.4­ml aliquot of 0.1% (w/v) 3­(2­pyridyl)­5,6­ diphenyl­1,2,4­triazine was added. The volume was made up to 3 ml with absolute ethanol, 0.4 ml 0.1% (w/v) ferric chloride (FeCl3.6H2O) was added, and the content was gently mixed under dim light in a dark room to avoid photo­ chemical reduction. After a 4­minute reaction at room temperature, 0.2 ml 0.2 M orthophosphoric acid was added, and the mixture was left for another 30 min. Absorbance was determined spectrophoto­ metrically at 554 nm and reported as μg g­1 FW. The blank was prepared the same way; absolute ethanol was used instead of the sample. α­tocopherol (Sigma Chemical) was used as a standard. Glutathione concentration Two hundred mg of tissue was homogenized in 2 ml of ice­cold 5% TCA. The homogenate was cen­ trifuged at 17,000 × g for 30 min at 4°C. A volume of 75 μl of the supernatant was added to a cuvette con­ taining 300 μl of phosphate buffer (0.2 M, pH 8.0) and 750 μl of 0.6 mM DTNB (5,5ˊ­dithiobis­ 2­ nitrobenzoic acid) in phosphate buffer. The absorbance at 412 nm was read, and glutathione concentration was derived against a standard curve prepared with known amounts of GSH in 5% TCA (Moron et al., 1979). Leaf total polyphenols This parameter was determined using the Folin­ Ciocalteu reagent. One gram of freeze­dried leaf samples was placed in an Eppendorf tube, mixed with 1 ml of methanol (80%), ground at 4°C, then centrifuged at 10000 × g for 15 minutes. The extract was mixed with 0.5 ml of Folin­Ciocalteu reagent; diluted 1:1 with water, and then 1 ml of a 5% sodium carbonate solution was added. Absorbance was mea­ sured at 725 nm and expressed as mg per g FW after 30 minutes. Gallic acid was used as the standard phe­ nolic compound (Bonyanpour and Jamali, 2020). Lipid peroxidation A sample of 0.2 g was homogenized in 2 ml 0.1% trichloroacetic acid (TCAA) solution and then cen­ trifuged at 15,000 × g for 10 min at 4°C, the 0.5 ml of supernatant was added to 1.5 mL 0.5% TBA in 20% TCAA, followed by incubation of the mixture at 95 °C in a shaking water bath for 30 min. The tubes were placed in an ice­water bath to stop the reaction. The samples were re­centrifuged at 10,000 × g for 5 min, and the absorbance of the supernatant was deter­ mined spectrophotometrically at 532 nm and cor­ rected for non­specific turbidity by subtracting the absorbance at 600 nm. The malondialdehyde (MDA) concentration was calculated using the extinction coefficient of 155mM­1 cm­1 (Cakmak and Horst, 1991). Leaves proline concentration Leaf samples were homogenized with 3% sulfosal­ icylic acid. The homogenate was centrifuged at 2500 × g for 20 min. The supernatant was treated with acid ninhydrin and acetic acid, boiled for 60 minutes, and then the absorbance of the solution was record­ ed at 520 nm. Leaves proline contents were expressed as μmol·g­1 FW. Proline amino acid was used as standard (Bates et al., 1973). Leaf ascorbic acid concentration This parameter was measured by the method of Omaye et al. (1979). To 1 g of freeze­dried leaf sam­ ple, 10% ice­cold TCA was added and centrifuged at 3500 × g for 20 min at room temperature. The super­ natant (1 ml) was mixed with 0.2 ml of DTC reagent and incubated at 37°C for 3 h. Then 1.5 ml of ice­cold 65% sulfuric acid was added and mixed, and the solu­ tions were allowed to stand at room temperature for another 30 min. The color developed was read at 520 nm spectrophotometrically and reported as μg g­1 FW. Macro and micronutrients’ concentration Oven­dried leaf samples (0.5 g) were ground and ashed at 550°C in a porcelain crucible for seven h. The white ash was mixed in 2 M hot HCl, filtered, and finally made up to 50 mL with distilled water. Nitrogen (N) concentration was measured using the Kjeldahl digestion method. Potassium (K) concentra­ tion was determined using the flame emission method using a Sherwood Scientific Ltd model 360 flame photometer. An atomic absorption spec­ trophotometer (AA 6200, double beam atomic absorption spectrophotometer Shimadzu, Kyoto, Japan) was used to determine Ca, Mg, and micronu­ trient element, including Fe, Zn, and Mn concentra­ tions. Phosphorus (P) concentration was determined colorimetrically (Kalra, 1998). Leaf biometrics Leaf area was determined using a leaf area meter Jamali and Amin ‐ Characterization of caprifig cultivars 307 (Kaiser RS1) and expressed as mm2; leaf and petiole length by using a digital caliper and expressed as mm. Fruit fresh and dry weight Spring fruits were harvested and weighed using a digital scale and reported as gram. Then they were oven­dried and weighed again. Experimental design and statistical analysis The experiment was carried out in a completely randomized block design with four replications. Data were analyzed by ANOVA test using SPSS (Ver. 9.1); means were compared using Duncan’s multiple range test at 5% probability level. Cluster analysis was also performed according to Ward’s minimum­variance method using SPSS (Ver. 9.1) to classify the cultivars. 3. Results The leaf glutathione concentration in NGM (67.01 μg g­1 FW) was nearly 50% higher than MS (32.32 μg g­1 FW). CO (64 μg g­1 FW) was not statistically differ­ ent when compared with NGM. The leaf MDA con­ centration in KZR (12.12 μg g­1 FW) was lower than MS and CO cultivars. Other caprifig trees were not statistically different (Table 2). The leaf ascorbic acid concentration in the GK cul­ tivar (17.03 μg g­1 FW), was 59% higher than DRB (7.14 μg g­1 FW). RS, CMB, PZD, JR, MS, and CO culti­ vars were not statistically different in comparison with GK. The leaf proline concentration in MS (6.77 μg g­1 FW) was not statistically different in compari­ son with GK, NGM, CMB, PR, JR, and MS cultivars. The leaf total polyphenols concentration in NGM (26.71 mg g­1 FW), was 56% higher when compared with the MGR cultivar. PZD, JR, and PSD were not sta­ tistically different compared to the NGM cultivar (Table 2). The leaf dry weight in NGM was 26% higher when compared with MGR. CMB, PZD, MS, CO, PSD, and SZ cultivars were not statistically different in comparison with NGM. Leaf total sugars were not statistically dif­ ferent in all cultivars. LRWC in the NGM cultivar was 8.4% higher when compared with MS . IJ, GK, MLZ, DS, CO, and SZ were not statistically different in com­ parison with NGM. Leaf EL in MGR (22.63%) was sig­ nificantly higher than DS (17.62%). This parameter was not statically different in other cultivars (Table 3). Leaf nitrogen concentration was not statistically different in all cultivars. The leaf P concentration in DS (3.32 mg g­1 DW) was higher in comparison with PR, JR, and CO; other cultivars were not different. Table 2 ­ The concentration of some non­enzymatic antioxidants and MDA in leaves of studied cultivars Z Means followed by the same letters within columns are not different at 5% probability using Duncan’s test. Cultivar Glutathione (µg g­1 fresh weight) MDA (µg g­1 fresh weight) α­Tocopherol (µg g­1 fresh weight) Ascorbic acid (µg g­1 fresh weight) Proline (µg g­1 fresh weight) Polyphenol (mg g­1 fresh weight) MGR 45.02 fg z 15.00 abc 92.92 d 7.40 c 2.83 d 17.06 e DRB 44.30 fg 16.00 abc 110.41 ab 7.14 c 3.65 cd 17.17 e ASA 53.3 de 14.07 bc 102.13 bc 8.88 c 3.33 d 18.19 e KZR 57.20 cd 12.12 c 69.82 e 10.12 bc 4.41 bcd 18.88 e IJ 50.10 ef 17.07 abc 97.04 cd 10.21 bc 3.45 cd 22.09 bcd GK 61.11 bc 15.19 abc 72.20 e 17.03 a 5.55 abc 18.29 e RS 53.44 d 15.41 abc 105.45 bc 12.00 abc 4.14 bcd 17.21 e NGM 67.01 a 12.14 c 122.18 a 8.22 c 6.71 a 26.71 a MLZ 48.15 ef 13.25 c 114.12 a 9.06 c 2.92 d 19.82 de CMB 40.22 gh 17.15 abc 115.45 a 16.33 a 6.20 ab 18.85 e PZD 52.36 de 18.02 abc 105.31 bc 12.14 abc 4.38 bcd 25.42 ab DS 61.10 bc 14.14 bc 100.05 bc 7.63 c 2.78 d 23.41 bc PR 45.16 fg 13.14 c 77.14 e 10.47 bc 5.61 abc 22.22 bcd JR 37.35 hi 14.20 bc 98.32 cd 12.42 abc 5.99 ab 24.56 ab MS 32.32 i 20.37 ab 77.11 e 15.15 ab 6.77 a 20.37 cde CO 64.00 ab 21.07 a 100.07 cd 15.71 ab 4.70 abcd 20.17 cde PSD 40.05 gh 15.3 abc 105.50 bc 9.16 c 3.77 cd 20.22 abc SZ 41.07 gh 13.33 c 115.00 a 8.05 c 3.42 cd 22.15 bcd 308 Adv. Hort. Sci., 2022 36(4): 303­314 The leaf K concentration in MLZ (22.46 mg g­1 DW) which was not statistically different compared with ASA, KZR, GK, RS, NGM, PZD, DS, PR, JR, MS, CO, PSD, and SZ. The leaf Ca concentration (4.46 mg g­1 DW) in NGM was 25% higher than PR. CMB had the highest leaf Mg concentration (2.48 mg g­1 DW); KZR had 42% lower leaf Mg concentration. Other cultivars were not statistically different (Table 4). The leaf Fe concentration in NGM (67.71 mg kg­1 DW) was significantly higher than KZR, MLZ, PR, JR, MS, PSD, and SZ. The leaf Zn concentration in NGM (27.31 mg kg­1 DW) was not statistically different in comparison with DRB, MLZ, PSD, and SZ. Leaf Mn concentration was not statistically different in all cul­ tivars (Table 5). The leaf total chlorophyll concentration in NGM (1.43 mg g­1 FW) was not statistically different com­ pared with DRB, ASA, KZR, MLZ, DS, MS, PSD, and SZ (Fig. 1). The leaf concentration of carotenoids in PSD was significantly higher than ASA, KZR, GK, CMB, PZD, RS, JR, and MS. Other cultivars were not statistically different in comparison with PSD (Fig. 2). The concen­ tration of leaf anthocyanins in MGR was significantly higher than PSD. This parameter was not significantly different in other cultivars (Fig. 3). Table 3 ­ Leaf relative water content, electrolyte leakage, dry matter, and total sugars in studied caprifig cultivars Z Means followed by the same letters within columns are not dif­ ferent at 5% probability using Duncan’s test. Cultivar Leaf dry matter (%) Leaf total sugars (mg g­1 dry weight) Leaf relative water content (%) Electrolyte leakage (%) MGR 26.22 b z 27.33 a 77.92 d 22.63 a DRB 28.30 b 26.05 a 81.21 abc 22.04 ab ASA 27.33 b 27.17 a 78.89 bcd 21.87 ab KZR 26.55 b 29.55 a 77.92 cd 19.12 ab IJ 27.44 b 26.77 a 83.04 a 20.00 ab GK 28.15 b 27.37 a 80.08 abcd 22.03 ab RS 27.61 b 28.47 a 79.52 bcd 21.43 ab NGM 33.05 a 29.14 a 83.60 a 17.96 ab MLZ 28.15 b 30.13 a 81.11 abc 19.06 ab CMB 30.31 ab 28.45 a 79.63 bcd 22.33 ab PZD 29.55 ab 29.02 a 77.32 d 22.34 ab DS 27.22 b 27.37 a 80.29 abcd 17.62 b PR 29.71 ab 28.02 a 77.14 d 20.77 ab JR 28.44 b 30.00 a 78.32 cd 19.72 ab MS 29.81 ab 28.88 a 77.11 d 19.75 ab CO 28.88 ab 30.11 a 81.00 abc 21.72 ab PSD 29.02 ab 27.14 a 79.67 bcd 19.55 ab SZ 28.87 ab 28.64 a 82.22 ab 19.15 ab Table 4 ­ Leaf macronutrients in studied caprifig cultivars Z Means followed by the same letters within columns are not different at 5% probability using Duncan’s test. Cultivar N (mg g­1 dry weight) P (mg g­1 dry weight) K (mg g­1 dry weight) Ca (mg g­1 dry weight) Mg (mg g­1 dry weight) MGR 22.12 a z 2.60 ab 16.33 c 3.76 bc 2.00 ab DRB 20.13 a 3.10 ab 17.16 bc 3.77 bc 2.22 ab ASA 20.15 a 3.05 ab 19.65 abc 3.97 abc 2.02 ab KZR 21.41 a 2.52 ab 18.27 abc 4.44 a 1.55 b IJ 21.05 a 2.97 ab 17.05 bc 3.65 bc 1.95 ab GK 21.77 a 2.55 ab 21.20 abc 3.80 bc 1.77 ab RS 22.30 a 2.92 ab 18.85 abc 4.20 ab 1.67 ab NGM 22.12 a 3.00 ab 20.38 abc 4.46 a 2.35 ab MLZ 19.64 a 3.14 ab 22.46 a 3.95 abc 2.20 ab CMB 20.51 a 2.46 ab 16.44 c 3.67 bc 2.48 a PZD 22.35 a 2.81 ab 18.71 abc 3.72 bc 2.36 ab DS 19.19 a 3.32 a 18.22 abc 3.90 abc 1.83 ab PR 22.66 a 2.45 b 19.70 abc 3.55 c 2.11 ab JR 20.44 a 2.37 b 19.00 abc 3.63 bc 2.07 ab MS 20.11 a 2.88 ab 20.77 abc 4.18 ab 1.61 ab CO 22.00 a 2.44 b 18.00 abc 3.76 bc 2.18 ab PSD 21.02 a 3.11 ab 21.56 ab 4.02 ab 2.14 ab SZ 21.48 a 3.23 ab 22.07 ab 3.89 abc 2.25 ab Jamali and Amin ‐ Characterization of caprifig cultivars 309 The highest fruit FW and DW were obtained from DS. NGM had the highest leaf length (154.66 mm). PR, MS, PZD, CMB, RS, and KZR were not statistically different. The leaf area (11581.2 mm2) and petiole length (66.69 mm) in NGM were significantly higher than all other cultivars (Table 6). The cluster analysis based on Ward’s method divided the cultivars into three major clusters, includ­ ing 8 cultivars in cluster 1, 9 cultivars in cluster 2, and 1 cultivar in cluster 3. The first cluster consisted of two subclusters: subcluster 1 (MGR, IJ, ASA, and DS) and subcluster 2 (DRB, PSD, SZ and MLZ). Cluster 2 also had 2 subclusters: subcluster 1 (KZR, MS, and RS) and subcluster 2 (GK, CO, CMB, JR, PZD, and PR). Cluster 3 had only one cultivar: NGM (Fig. 4). 4. Discussion and Conclusions Non­enzymatic antioxidants were statistically dif­ ferent among investigated cultivars in our study, which was consistent with previous studies. Jamali Fig. 2 ­ Concentration of leaf carotenoids in studied caprifig cul­ tivars. Columns with the same letters are not statistically different at 5% probability using Duncan’s test. Vertical bars indicate standard errors (n=4). Fig. 3 ­ Concentration of leaf anthocyanins in studied caprifig cultivars. Columns with the same letters are not statisti­ cally different at 5% probability using Duncan’s test. Vertical bars indicate standard errors (n=4). Fig. 1 ­ Total chlorophyll concentration in studied caprifig culti­ vars. Columns with the same letters are not statistically different at 5% probability using Duncan’s test. Vertical bars indicate standard errors (n=4). Table 5 ­ Leaf micronutrients in studied caprifig cultivars Z Means followed by the same letters within columns are not dif­ ferent at 5% probability using Duncan’s test. Cultivar Fe (mg kg­1 dry weight) Zn (mg kg­1 dry weight) Mn (mg kg­1 dry weight) MGR 60.66 ab z 23.33 bcd 47.22 a DRB 60.47 ab 26.50 ab 50.16 a ASA 62.23 ab 22.74 cd 49.44 a KZR 58.36 b 22.46 cd 50.33 a IJ 63.33 ab 23.00 bcd 49.31 a GK 64.44 ab 21.20 d 48.56 a RS 60.96 ab 22.85 cd 47.91 a NGM 67.71 a 27.31 a 48.05 a MLZ 57.22 b 25.44 abc 50.20 a CMB 61.22 ab 22.07 cd 50.27 a PZD 63.42 ab 23.08 bcd 49.22 a DS 60.75 ab 22.77 cd 48.71 a PR 59.29 b 23.12 bcd 48.05 a JR 57.33 b 21.90 cd 49.66 a MS 59.40 b 22.35 cd 50.23 a CO 60.05 ab 22.68 cd 51.44 a PSD 58.63 b 24.88 abcd 50.63 a SZ 59.34 b 25.14 abc 47.33 a Adv. Hort. Sci., 2022 36(4): 303­314 310 and Bonyanpour (2017) determined leaf mineral composition and also some biochemical parameters, i.e., concentration of non­enzymatic antioxidants in leaves, in seven Iranian pomegranate cultivars for selecting probable more tolerant cultivars. They found that cultivars with a higher concentration of non­enzymatic antioxidants (e.g., glutathione, α­ Tocopherol, proline, etc.) had better adaptability to the prevailing environmental conditions. As men­ tioned earlier, various species and cultivars have inherently different potential to tolerate non­optimal growth conditions. One of the possible reasons for this difference is their enzymatic/non­enzymatic antioxidant responses and/or their ability to absorb macro/micro nutrients. Therefore, determination of these characteristics can be used as biochemical markers for cultivar comparison/screening. Gholami et al. (2012) compared four fig cultivars (Deyme Ahvaz, Sabz Estahban, Siah, and Shahanjir) under normal, drought, and recovery conditions. According to their results, some of the non­enzymat­ ic and enzymatic antioxidants in leaves differed sig­ nificantly under optimal conditions. Their findings show that the fig cultivars demonstrated a clear dif­ ference in their response to water stress and recov­ ery. They evaluated Deyme Ahvaz as a more tolerant cultivar than Sabz Estahban. Deyme Ahvaz had higher leaf concentration of glutathione and anthocyanin in comparison to Sabz Estahban under non­stress condi­ tions. Samec et al. (2021) reviewed the role of phenolic compounds in inducing tolerance to non­optimal conditions in plants. They reported that the potential tolerance of a plant species to stress conditions can be associated with this diverse family of chemicals which is universally present in plants and includes Fig. 4 ­ Cluster analysis dendrogram of 18 Iranian caprifig culti­ vars based on Ward's method. Table 6 ­ Fruit fresh and dry weight and leaf and petiole length, and leaf area in studied caprifig cultivars Z Means followed by the same letters within columns are not different at 5% probability using Duncan’s test. Cultivar Fruit fresh weight (g) Fruit dry weight (g) Leaf length (mm) Leaf area (mm2) Petiole length (mm MGR 4.77 hi z 0.90 fg 78.97 g 1879.5 j 23.45 h DRB 5.77 gh 1.09 ef 89.70 fg 3217.3 efg 29.36 fgh ASA 6.48 efg 1.20 de 83.54 g 2232.6 hij 23.08 h KZR 7.60 cde 1.21 de 121.63 bc 6352.9 b 37.46 cd IJ 4.94 hi 0.95 fg 98.97 ef 2005.9 ij 28.60 fgh GK 8.06 cd 1.48 bc 107.57 de 4433.1 d def 10/33 RS 8.64 c 1.76 a 118.04 bcd 7090.9 b 50.72 b NGM 6.63 efg 0.94 fg 154.66 a 11581.2 a 66.69 a MLZ 4.01 i 0.72 g 79.28 g 2814.7 fgh 26.57 gh CMB 7.12 def 1.09 ef 116.12 bcd 3698.2 def 38.30 cd PZD 5.92 fgh 0.96 f 117.25 bcd 4373.1 d 35.95 de DS 12.40 a 1.71 a 98.90 ef 2522.8 ghij 30.79 efg PR 6.39 efg 1.03 ef 113.67 bcd 4554.1 d 37.77 cd JR 6.37 efg 1.07 ef 99.53 ef 3980.3 de 32.84 defg MS 10.88 b 1.58 ab 125.09 b 6518.2 b 42.68 c CO 6.76 efg 1.02 ef 112.09 cd 5459.2 c 43.36 c PSD 6.67 efg 1.36 cd 81.50 g 3334.1 efg 28.24 fgh SZ 4.94 hi 0.9 fg 87.78 fg 2965.6 fgh 29.02 fgh Jamali and Amin ‐ Characterization of caprifig cultivars 311 more than 8000 molecules. Phenolic compounds play an essential role in plant responses, especially in defense mechanisms. They have strong antioxidant properties and neutralize the detrimental effects of reactive oxygen species (Balasundram et al., 2006). The absolute concentrations of these chemicals in plant organs vary significantly depending on the determination method, cultivar, or tissue, but it can be concluded that higher phenol concentrations cor­ relate with increased stress tolerance. Arteaga et al. (2020) have proposed the use of proline as a biochemical marker for rapid and simple large­scale screening of different genotypes for drought and salt tolerance. Transgenic plants, espe­ cially those overexpressing genes for proline accumu­ lation, show higher adaptation to abiotic stresses (El Moukhtari et al., 2020; Ghosh et al., 2022. Similarly, endogenous proline concentration has been linked to a relatively higher stress tolerance when comparing different cultivars by various authors in different species (Kapuya et al., 1985; Misra and Gupta, 2005; Goharrizi et al., 2020). The lower MDA content in NGM, RS, or MLZ may be due to the higher content of antioxidants such as α­tocopherol in these two caprifigs. This is consistent with previous studies (Assaha et al., 2015; Amoah et al., 2019; Nawaz and Wang, 2020). Except for leaf N and Mn concentration, other macro and micronutrients were significantly different in various caprifig cultivars. This was in agreement with previous studies on different fig cultivars (Anac et al., 1982; Aksoy et al., 1987; Askin et al., 1998; Hakerlerler et al., 1998; Bougiouklis et al., 2020). The higher leaf DW or LRWC in NGM could be attributed to the higher leaf concentration of K, Ca, Fe, or Zn in this caprifig in comparison with other studied cultivars. This was in agreement with previ­ ous studies as various cultivars absorb macro/micronutrients differently. Hegwood (1972) reported a significant varietal effect on leaf mineral composition during full bloom and harvest in eleven snapbean cultivars. Jordão et al. (1999) studied 15 olive cultivars and found that the mean effect of cul­ tivar on leaf concentrations of essential elements, including N, P, K, Ca, Mg, S, Mn, Zn, and B, was signif­ icant. There are also many other examples of the effects of cultivar or root system of different species on leaf mineral composition (Tsipouridis and Thomidis, 2005; North and Cook, 2006; Tomala et al., 2008; Kviklys et al., 2012). This difference in endogenous macro/micronutri­ ent composition can affect numerous aspects of plant growth and development. In other words, the presence of a particular macro­ or micronutrient (e.g., Ca) above a critical concentration can alter the plant’s response to non­ optimal growing conditions (Pilbeam and Morley, 2007; Taiz and Zeiger, 2010). For instance, Fageria (2013) reported an increased root DW in 12 lowland rice genotypes after adding K fertilizer. They found that increase in root DW at high K levels was 246% compared to the low level of K. Leaf chlorophyll and carotenoids concentration were significantly different in studied cultivars. Chlorophyll concentration is a sensitive biochemical marker indicating cellular metabolic state (Chutipaijit et al., 2011). Previous studies have shown that chlorophyll concentration and stability correlate with plants’ high tolerance potential against abiotic stress­ es such as salinity or drought (Hasanuzzaman et al., 2013). Carotenoids protect the photosynthesis appa­ ratus by scavenging free radicals keeping its integrity against photo­oxidative damages (Dall’Osto et al., 2007; Andrade­Souza et al., 2011). As an ABA precur­ sor, higher carotenoid concentration means lower photo­oxidative damage and elevated potential for regulating plant growth under stress conditions (Götz et al., 2002; Han et al., 2008). Caprifig cultivars in our study were clustered into three main groups indicating lower level of variability within clusters and higher variability levels between clusters. Morphological and biochemical markers have long been used for screening and characterizing different genotypes, as they are the first steps in describing/classifying any germplasm (Cantini et al., 1999). Previous studies on figs show the usefulness of these markers in documenting variability among genotypes (Salhi­Hannachi et al., 2006; Saddoud et al., 2008; Podgornik et al., 2010). NGM had the highest leaf glutathione, α­ Tocopherol, proline, polyphenols, Ca, and Zn concen­ trations. Leaf length, leaf area, and petiole length were the highest in this cultivar. Cluster analysis indi­ cated that NGM was the only cultivar in cluster 3. In conclusion, NGM was evaluated as a genotype which seems to be more distant (morphologically and bio­ chemically) from other studied caprifigs and probably more adaptable/tolerant. Our findings can be used as an overture for complementary studies for selecting promising genotypes for breeding and introducing new tolerant fig cultivars. Adv. Hort. Sci., 2022 36(4): 303­314 312 References AKSOY U., ANAC D., ERYUCE N., YOLTAS T., 1987 ­ Determination and evaluation of the nutrients status of fig orchards in the Aegean Region. ­ J. Ege Univ. Fac. Agric., 24(2): 21­35. AMOAH J.N., KO C.S., YOON J.S., WEON S.Y., 2019 ­ Effect of drought acclimation on oxidative stress and tran‐ script expression in wheat (Triticum aestivum L.). ­ J. Plant Int., 14(1): 492­505. ANAC D., AKSOY U., HAKERLERLER H., DUZBASTOLAR M., 1982 ­ Nutritional status of fig orchard in the small meander basin and relationships between soil and leaf nutrients and some yield and quality attributes. ­ Taris Research and Development Centre Project No. 4. ANDREADE­SOUZA V., COSTA M.G.C., CHEN C.X., GMITTER F.G., COSTA M.A., 2011 ­ Physical location of the carotenoid biosynthesis genes Psy and b‐Lcy in Capsicum annuum (Solanaceae) using heterologous probes from Citrus sinensis (Rutaceae). ­ Gen. Mol. Res., 10: 404­409. ARADHYA M.K., STOVER E., VELASCO D., KOEHMSTEDT A., 2010 ­ Genetic structure and differentiation in cultivat‐ ed fig (Ficus carica L.). ­ Genetica, 138: 681­694. ARTEAGA S., YABOR L., DÍEZ M.J., PROHENS J., BOSCAIU M., VICENTE O., 2020 ­ The use of proline in screening for tolerance to drought and salinity in common bean (Phaseolus vulgaris L.) genotypes. ­ Agronomy, 10(6): 817. ASKIN A., CEYLS S., YENER, H., 1998 ­ A study on the national status of fig orchards in Birgiirimagzi. ­ Acta Horticulturae, 480: 239­246. ASSAHA D.V., MEKAWY A. M. M., UEDA A., SANEOKA H., 2015 ­ Salinity‐induced expression of HKT may be cru‐ cial for Na+ exclusion in the leaf blade of huckleberry (Solanum scabrum Mill.), but not of eggplant (Solanum melongena L.). ­ Biochem. Biophys. Res. Commun., 460: 416­421. BALASUNDRAM N., SUNDRAM K., SAMMAN S., 2006 ­ Phenolic compounds in plants and Agri‐industrial by‐ products: antioxidant activity, occurrence, and poten‐ tial uses. ­ Food Chem., 99: 191­203. BATES L.S., WALDERN R.P., TEARE I.D., 1973 ­ Rapid deter‐ mination of free proline for water‐stress studies. ­ Plant Soil., 39: 205­207. BONYANPOUR A.R., JAMALI B., 2020 ­ Seasonal enzymatic and non‐enzymatic antioxidant responses in seven Iranian pomegranate cultivars. ­ Adv. Hort. Sci., 34(3): 265­276. BOUGIOUKLIS J.N., KARACHALIOU Z., TSAKOS J., KALKANIS P., MICHALAKOS A., MOUSTAKAS N., 2020 ­ Seasonal variation of macro‐ and micro‐ nutrients in leaves of fig (Ficus carica L.) under Mediterranean conditions. ­ Agron. Res., 18(4): 2328­2339. CAKMAK I., HORST J.H., 1991­ Effects of aluminium on lipid peroxidation, superoxide dismutase, catalase, and per‐ oxidase activities in root tips of soybean (Glycine max). ­ Physiol. Plant., 83: 463­468. CANTINI C., CIMATO A., SANI G., 1999 ­ Morphological evaluation of olive germaplasm present in Tuscany region. ­ Euphytica., 109: 173­181. CHONG T.M., ABDULLAH M.A., FADZILLAH N.M., LAI O.M., LAJIS N.H., 2004 ­ Anthraquinones production, hydro‐ gen peroxide level and antioxidant vitamins in Morinda elliptica cell suspension cultures from intermediary and production medium strategies. ­ Plant Cell Rep., 22: 951­958. CHUTIPAIJIT S., CHAUMS S., SOMOPORNAILIN K., 2011 ­ High contents of proline and anthocyanin increase pro‐ tective response to salinity in Oryza sativa L. spp. ­ Indica. Aust. J. Crop. Sci., 5: 1191­1198. DALL’OSTO L., FIORE A., CAZZANIGA S., GIULIANO G., BASSI R., 2007 ­ Different roles of alpha and beta branch xan‐ thophylls in photosystem assembly and photoprotec‐ tion. ­ J. Biol. Chem., 282: 35056­35068. EL MOUKHTARI A., CABASSA­HOURTON C., FARISSI M., SAVOURÉ A., 2020 ­ How does proline treatment pro‐ mote salt stress tolerance during crop plant develop‐ ment?. ­ Front. Plant Sci., 11: 1127. FAGERIA N.K., 2013 ­ The Role of plant roots in crop pro‐ duction. ­ CRC Press, Boca Raton, Florida, USA, pp. 461. GHOLAMI M., RAHEMI M., KHOLDEBARIN B., RASTEGAR S., 2012 ­ Biochemical responses in leaves of four fig culti‐ vars subjected to water stress and recovery. ­ Sci. Hort., 148: 109­117. GHOSH U.K., ISLAM M.N., SIDDIQUI M.N., CAO X., KHAN M.A.R., 2022 ­ Proline, a multifaceted signalling mole‐ cule in plant responses to abiotic stress: understanding the physiological mechanisms. ­ Plant Biol., 24(2): 227­ 239. GOHARRIZI K., BAGHIZADEH A., AFROUSHTEH M., AMIRMAHANI F., KERMANI, S., 2020 ­ Effects of salinity stress on proline content and expression of Δ1‐pyrro‐ line‐5‐carboxylate synthase and vacuolar‐type H sub‐ unit E genes in wheat. ­ Plant Genet. Resour., 18(5): 334­342. GÖTZ T., SANDMANN G., RÖMER S., 2002 ­ Expression of bacterial carotene hydroxylase gene (crtZ) enhances UV tolerance in tobacco. ­ Plant Mol. Biol., 50: 127­140. HAKERLERLER H., SAATÇI N., HEPAKSOY S., AKSOY U., 1998 ­ Fruit and leaf nutritional status of some Fig clones and cultivars and relationships with some sugar fractions. ­ Acta Horticulturae, 480: 247­252. HAN H., LI H., ZHOU S., 2008 ­ Overexpression of phytoene synthase gene from Salicornia europaea alters response to reactive oxygen species under salt stress in transgenic Arabidopsis. ­ Biotech. Lett., 30(8): 1501­ 1507. HASANUZZAMAN M., NAHAR K., FUJITA M., 2013 ­ Plant response to salt stress and role of exogenous protec‐ Jamali and Amin ‐ Characterization of caprifig cultivars 313 tants to mitigate salt‐induced damages, pp. 25­77. ­ In: AHMAD P., P.P. AZOOZ, AND M.N.V. PRASAD (eds.). Ecophysiology and Responses of Plants under Salt Stress. Springer Verlag, New York, NY, USA, pp. 510. HEGWOOD D.A., 1972 ­ Cultivar effects on leaf and fruit mineral composition in snapbeans, Phaseolus vulgaris. ­ Commun. Soil Sci. Plant Anal., 3(2): 123­139. JAMALI B., BONYANPOUR A.R., 2017 ­ Evaluation of adapt‐ ability potential of seven Iranian pomegranate cultivars in southern Iran, Arsenjan region. ­ Adv. Hort. Sci., 31(2): 97­105. JAMALI B., ESHGHI S., 2015 ­ Salicylic acid‐induced salinity redressal in hydroponically grown strawberry. ­ Comm. Soil Sci. Plant Anal., 46: 1482­1493. JAMALI B., ESHGHI S., KHOLDEBARIN B., 2016 ­ Changes in antioxidant activities of strawberry cv. Selva as affected by salicylic acid application timing under saline condi‐ tions. ­ J. Berry Res. 6: 291­301. JORDÃO P.V., MARCELO M.E., CENTENO M.S.L., 1999 ­ Effect of cultivar on leaf mineral composition of olive tree. ­ Acta Horticulturae, 474: 349­352. KALRA Y.P., 1998 ­ Handbook of reference methods for plant analysis. ­ CRC Press, New York, NY, USA, pp. 287. KANNO C., YAMAUCHI K., 1997 ­ Application of new iron reagent, 3‐(2‐pyridyl)‐5,6‐diphenyl‐1,2,4‐triazine, to spectrophotometric determination of tocopherols. ­ Agric. Biol. Chem., 41: 593­596. KAPUYA J.A., BARENDSE G.W.M., LINSKENS H.F., 1985 ­ Water stress tolerance and proline accumulation in Phaseolus vulgaris L. ­ Acta Bot. Neerl., 34: 293­300. KVIKLYS D., KVIKLIENE N., BITE A., LEPSIS J., UNIVER T., UNIVER N., USELIS N., LANAUSKAS J., BUSKIENE L., 2012 ­ Baltic fruit rootstock studies: Evaluation of 12 apple rootstocks in North‐East Europe. ­ Hortic. Sci., 39: 1­7. LICHTENTHALER H.K., 1987 ­ Chlorophylls and carotenoids: Pigments of photosynthetic bio membrane. ­ Methods Enzymol., 148: 350381. MISRA N., GUPTA A.K., 2005 ­ Effect of salt stress on pro‐ line metabolism in two high yielding genotypes of green gram. ­ Plant Sci., 169: 331­339. MORON M.S., DEPIERRE J.W., MANNERVIK B., 1979 ­ Levels of glutathione, glutathione reductase and glu‐ tathione S‐transferase activities in rat lung and liver. ­ Biochim. Biophys. Acta, 582: 67­78. NAWAZ M., WANG Z., 2020 ­ Abscisic acid and glycine betaine mediated tolerance mechanisms under drought stress and recovery in Axonopus compressus: A New Insight. ­ Sci. Rep. 10: 6942. NORTH M., COOK N., 2006 ­ Effect of six rootstocks on ‘Forelle’ pear tree growth, production, fruit quality and leaf mineral content. ­ Acta Horticulturae, 772: 97­103. OMAYE S.T., TURNBALL J.D., SAUBERLICH H.E., 1979 ­ Selected methods for the determination of ascorbic acid in animal cells, tissues and fluids. ­ Methods Enzymol., 62: 3­11. PEREZ­JIMÉNEZ M., LÓPEZ B., DORADO G., PUJADAS­ SALVÁ A., GUZMÁN G., HERNANDEZ P., 2012 ­ Analysis of genetic diversity of southern Spain fig tree (Ficus car­ ica L.) and reference materials as a tool for breeding and conservation. ­ Hereditas, 149: 108­113. PILBEAM D.J., MORLEY P.S., 2007 ­ Calcium, pp. 121­145. ­ In: BARKER A.V., and D.J. PILBEAM (eds.) Handbook of Plant Nutrition, CRC Press, NY, USA, pp. 632. PODGORNIK M., VUK I., VRHOVNIK I., MAVSAR D.B., 2010 ­ A survey and morphological evaluation of fig (Ficus car­ ica L.) genetic resources from Slovenia. ­ Sci. Hortic., 125: 380­389. POURGHAYOUMI M., BAKHSHI D., RAHEMI M., NOROOZISHARAF A., JAFARI M., SALEHI M., CHAMANE R., HERNÁNDEZ F., 2016 ­ Phytochemical attributes of some dried fig (Ficus carica L.) fruit cultivars grown in Iran. ­ Agric. Cons. Sci., 81(3): 161­166. RAHEMI M., JAFARI M., 2008 ­ Effect of Caprifig type on quantity and quality of Estahban dried fig Ficus carica cv. Sabz. ­ Acta Horticulturae, 798: 249­252. SADDOUD O., BARAKET G., CHATTI K., TRIFI M., MARRAKCHI M., SALHI­HANNACHI A., MARS M., 2008 ­ Morphological variability of fig (Ficus carica L.) culti‐ vars. ­ Int. J. Fruit Sci., 8: 35­51. SALHI­HANNACHI A., CHATTI K., SADDOUD O., MARS M., RHOUMA A., MARRAKCHI M., TRIFI M., 2006 ­ Genetic diversity of different Tunisian fig (Ficus carica L.) collec‐ tion revealed by RAPD fingerprints. ­ Hereditas, 143: 15­22. SAMEC D., KARALIJA E., ŠOLA I., BOK VV., SALOPEK­SONDI B., 2021 ­ The role of polyphenols in abiotic stress response: The Influence of molecular structure. ­ Plants, 10(118): 1­24. SHARMA P., JHA A.B., SHANKER D.R., PESSARAKLI M., 2012 ­ Reactive oxygen species, oxidative damage and antioxidative defense mechanism in plants under stressful conditions. ­ J. Bot., 2012: 1­26. SUGIURA T., KURODA H., SUGIURA, H., 2007 ­ Influence of the current state of global warming on fruit tree growth in Japan. ­ Hortic. Res., 6: 257­263. TAIZ L., ZEIGER E., 2010 ­ Plant physiology. ­ Sinauer Associates Inc., New York, NY, USA, pp. 782. TIAN X.Y., HE D.D., BAI S., ZENG W.Z., WANG Z., WANG M., 2021 ­ Physiological and molecular advances in magne‐ sium nutrition of plants. ­ Plant Soil, 468: 1­17. TOMALA K., ANDZIAK J., JEZIOREK K., DZIUBAN R., 2008 ­ Influence of rootstock on the quality of ‘Jonagold’ apples at harvest and after storage. ­ J. Fruit Ornam. Plant Res., 16: 31­38. TSIPOURIDIS C., THOMIDIS T., 2005 ­ Effect of 14 peach rootstocks on the yield, fruit quality, mortality, girth expansion and resistance to frost damages of May Crest peach variety and their susceptibility on Phytophthora citrophthora. ­ Scientia Hortic., 103: 421­ Adv. Hort. Sci., 2022 36(4): 303­314 314 428. ZHANG X., KONG W., WANG W., ZHANG J., LIU L., WANG W., LIU Y., WANG X., ZHANG H., DENG Q., 2020 ­ Genetic diversity analysis of 34 fig varieties (Ficus carica L.) based on ISSR molecular marker. ­ Genet. Resour. Crop Evol., 67: 913­921.