Chickpea seed mass influences agronomical performance: a case for seed heteromorphism?

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Ariel Villalobos-Olivera, Roberto Campbell, Marcos Edel Martínez-Montero, and 5 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-2496069/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Cicer arietinum L. is a major food legume across the globe. However, the yield of legume crops appears to have reached a plateau in developing countries where yield is often impacted by poor crop establishment. Therefore, seed physiological characteristics (specific to the cultivars/landraces adapted to various regions of the world) and their impacts on plantlet establishment and performance should be investigated. This study determined the effect of seed size on germination, plant development and agronomic performance in Cuba. Biochemical parameters were also evaluated for up to 21 d of growth. The results showed that seeds of the largest mass (89 ± 3 mg, group 3) showed more rapid germination, emergence and plant growth than the other tested mass categories. This trend was sustained until plant maturity where group 3 seeds also generated the highest yields. Differences were also noted in the antioxidant profiles in developing plants with the highest levels of SOD and PER found in plants generated from seeds with the smallest mass (55 ± 3 mg per seed, group 1). The above findings raise the questions as to whether seeds of chickpea display heteromorphic behavior, however, further studies are required. Cicer arietinum L. field performance seed heteromorphism seed size Figures Figure 1 Introduction Chickpea ( Cicer arietinum L.) is an ancient, self-pollinated legume crop believed to have originated in south-eastern Turkey and the adjoining part of Syria (Van der Maessen 1972 ). It is an annual plant that belongs to the Fabaceae family within the Faboideae subfamily (Shagarodsky et al. 2001 ). Plants are generally about 50 cm in height, with white or violet flowers from which pods develop. Each pod produces two or three seeds at most. Chickpeas are widely consumed by humans, and used as an energy and protein source in animal feed (Duarte-Leal et al. 2016 ; Kaloki et al. 2019 ; Alam et al. 2020 ; Janghel et al. 2020 ; Varol et al. 2020 ). An associated advantage of chickpea cultivation is that it improves soil fertility. Most recent estimates are that more than 11 million tons enters the world market annually (Merga et al. 2019 ). Some chickpea components have shown, in preclinical and clinical studies, several health benefits, including antioxidant capacity, antifungal, antibacterial, analgesic, anticancer, anti-inflammatory and hypocholesterolemic properties (Kuhn 2020 ). There are numerous chickpea landraces available, for example, the International Crops Research Institute for Semi-Arid Tropics (ICRISAT) in India maintains in excess of 20 000 accessions sourced from different countries (Bhagyawant et al. 2018 ). Nevertheless, crop improvement is still being pursued with the major goals being to exploit the available genetic potential by developing lines with improved yield (and nutritional characteristics) or to minimize the adverse effects of diseases, insects, drought, heat and cold (Singh 1997 ; Chowdhury et al. 2002 ; Devasirvatham et al. 2012 ; Maqbool et al. 2017 ; Gaur et al. 2019 ). While there have been studies over the years on seed developmental characteristics and subsequent growth, which are relevant for breeding programs (Hayes et al. 1955 ), there appears to be a lack of reports on the hetero-morphism of the seeds and their influence on the germination and growth of chickpea plants. Seed heteromorphism is an adaptive trait in response to the spatio-temporal variability of the habitat in which plants develop (Venable et al. 1998 ). This involves the production of seeds with different morphologies and/or post-harvest behavior in different parts of the same plant (Imbert 2002 ; Lu et al. 2010 ). Differences between morphs are usually based on one or more of the following: color, size, morphology/anatomy, dispersion syndrome, latency, position in the pod or spike, and germination (Baskin and Baskin 1998 ). In the Fabaceae family, seed heteromorphism has been reported in Amphicarpaea bracteata (Trapp 1988 ), Lathyrus linifolius (Dello Jacovo et al. 2019 ) and Teramnus labialis (Acosta et al. 2020 ). Considering the growing importance of chickpea in addressing food security challenges in developing countries and as a substitute for meat-based protein in developed regions, it is important to characterize the heteromorphic nature of seeds. The effect of seed heteromorphism can have an influence on species establishment in the agronomical context and yield of plants in the field. This study evaluated the influence of three different seed morphs (55 ± 3 mg per seed (group 1); 72 ± 3 mg (group 2); 89 ± 3 mg (group 3)) on: germination, emergence, early growth of plantlets, and some biochemical indicators: chlorophylls, total protein, superoxide dismutase and peroxidase activities. We selected these compounds because they are related to a wide range of important biochemical and physiological pathways such as plant response to stress and photosynthesis (Gross et al. 2000 ; Moller 2001 ; Porra 2002 ; Yaginuma et al. 2002 ). These parameters were evaluated up to 21 days (d) of growth. Agronomical traits at 3 months after planting were also recorded. Materials And Methods Harvested chickpea seeds (cv. Nacional 29) were air-dried at room temperature to 6% moisture content (fresh mass basis) and then stored for 4 months at 4°C in the dark in hermetically sealed containers. Seeds with 6% moisture content were used in subsequent experiments, as recommended by ISTA ( 2005 ). Three seed sizes of chickpea were compared: 55 ± 3 mg per seed (group 1); 72 ± 3 mg (group 2); 89 ± 3 mg (group 3). Physical characteristics relating to germination (radicle longer than 5 mm), emergence, plant height, number of leaves, total leaf weight; stem length, diameter and weight and main root length, diameter and weight were recorded for up to 21 d. In addition, levels of leaf chlorophyll a and b (Porra 2002 ), total protein (Bradford 1976 ), superoxide dismutase (SOD) (McCord and Fridovich 1969 ) and peroxidase (PER) (Pascual et al. 1983 ) activities were also recorded. Biochemical determinations were assayed using three independent samples with 100 mg of leaf material per sample. Chlorophyll pigments were extracted with 750 µl methanol (100%). Samples were centrifuged (at 10 000 rpm and 4°C for 15 min), supernatants were collected and the absorbance was read at 652.4 nm (Porra 2002 ). Protein extraction was carried out with Tris-HCl 0.1 mol l − 1 buffer, pH = 8.5–8.8, 5 mmol l − 1 EDTA and 20 mmol l − 1 β-mercaptoethanol. The total protein content was determined according to Bradford ( 1976 ). To determine superoxide dismutase and guaiacol peroxidase-specific activities, samples were finely ground in liquid nitrogen. Extraction was performed with Tris–HCl buffer (0.35 mol l − 1 , pH 8.0), EDTA (20 mmol l − 1 ), cysteine (15 mmol l − 1 ), PVPP (50%) and PMSF (0.2 mmol l − 1 ). Samples were homogenized on ice with polytron apparatus (Ultra-turrax T25). The homogenate was filtered through a two-folded piece of gauze and centrifuged at 10,290 xg (Beckman J-21, Palo Alto, CA) for 30 min at 4°C. Superoxide dismutase activity was measured according to McCord and Fridovich ( 1969 ). The reaction mixtures included 80 µl of plant extract and 900 µl of potassium phosphate–KOH (50 mmol l − 1 , pH 7.6), EDTA (0.1 mmol l − 1 ); cytochrome C (0.01 mmol l − 1 ), xanthine (0.05 mmol l − 1 ) and 20 µl xanthine oxidase (EC 1.2.3.22; 0.03 units). Absorbance (550 nm) was measured every 15 s for 3 min. Averages of linear section absorbance were used and cytochrome c extinction coefficient (21.1 mmol − 1 l cm − 1 ) was used. Superoxide dismutase activity was defined as U g − 1 plant fresh mass [U = enzyme quantity hydrolyzing 1 µmol of superoxide per hour (37°C)]. Specific activity was calculated as the rate of superoxide dismutase activity relative to protein content. Guaiacol peroxidase activity was measured according to Pascual et al. ( 1983 ). The reaction mixtures included 100 µl of plant material extract, 1.0 ml Tris–HCl buffer (0.01 mol l − 1 , pH 7.0), 150 µl guaiacol (100 mmol l − 1 ) and 20 µl hydrogen peroxide. Absorbance (470 nm) was measured every 15 s for 3 min. Averages of linear section absorbance were used and guaiacol extinction coefficient (5,570 x 10 − 6 µmol − 1 l cm − 1 ) was used. Guaiacol peroxidase activity was defined as U g − 1 plant fresh mass [U = enzyme quantity hydrolyzing 1 µmol substrate per minute (37°C)]. Specific activity was calculated as the rate of guaiacol peroxidase activity relative to protein content. The following agronomical traits were also measured during three months of growth in the field, as recommended by the International Board for Plant Genetic Resources (FAO/IPGRI 1994 ): plant height, fresh and dry weight; number of branches; time until anthesis in 50% of plants; total number of pods; number of filled pods; number of grains per pod, number and weight of grains per plant; weight of 100 grains; duration of plant cycle, and yield. Germination was assessed by placing the three seed morphs on filter paper in Petri dishes (Ø: 10 cm moistened with 15 mL of distilled water and five replicates of 10 seeds per dish). To evaluate young plantlets up to 21 d of growth, seeds were planted into pots (500 mL volume, one seed per pot) containing Ferralytic-red soil and filter-cake-sugarcane ash (1:1, v:v). There were five replicates of 10 pots for each treatment (50 seeds per treatment). To study adult plants, 90 seeds of each seed morph were randomly selected and sown in a plant bed (with Ferralytic-red soil and filter-cake-sugarcane ashes) under field conditions. Seeds were planted 70 x 25 cm apart (3 replicates of 30 seeds each) in a randomized complete block design. The experiment was conducted at the Field Experimental Station of the Bioplant Centre, University of Ciego de Ávila, Cuba (21 52´48.6´´ N, 78 41´32.6´´ W; 53 meters above sea level; Sept 2020 – Jan 2021). Temperatures at 13:00 averaged 33 o C during the experiment; relative humidity reached 80%; and the Photosynthetic Photon Flux was 1 140 µmol s − 1 . Technical instructions as provided by the Cuban Ministry for Agriculture to cultivate chickpea were applied. Fertilizers were not supplied. Seeds were not inoculated with Rhizobium . Microjet irrigation was used to water the plants for 5 min every 8 h. Border plants, which had more space to grow, were not considered. All statistical analyses (One-Way ANOVA and Tukey, p = 0.05) were carried out using SPSS (Version 8.0 for Windows, SPSS Inc., New York). The overall coefficients of variation (OCV) were calculated as follows: (standard deviation/average) * 100. In this formula, we considered the average values of the three treatments compared (seed sizes) to calculate the standard deviation and average. Therefore, the higher the difference between the three materials compared, the higher the OCV (Lorenzo et al. 2015 ). The OCVs were classified as Low from 1.73 to 24.42%, Medium from 24.42 to 47.11% and High from 47.11 to 69.79%. The OCVs were only calculated for those indicators with statistically significant differences according to ANOVA and Tukey tests. Results And Discussion The current study investigated the growth and developmental responses of three seed morphs of chickpea based on seed mass. Tables 1 – 3 summarize the results for the phenotypical growth (Table 1 ) and biochemical attributes (Table 2 ) of plants from all three morphs at intervals up to 21 d and also agronomic characteristics following 3 months of growth in the field (Table 3 ). A total of 61 indicators were evaluated across the study with 33 related to germination, emergence and seedling establishment up to 21 d (Table 1 ), 13 for biochemical evaluations (Table 2 ) and an additional 13 for agronomical traits (Table 3 ). During the early stages of plant growth (Table 1 ), significant differences were found in the majority of parameters assessed (i.e. 31 out of 33) and in all cases, group 3 seeds (seeds with the highest mass) yielded superior plant growth characteristics compared with seeds from the other two groups. This was evident in Fig. 1 which demonstrated the faster growth of the heaviest seeds (group 3) followed by intermediate growth of group 2 seeds and the smallest plants were produced by the lightest seeds (group 1). Table 1 Germination, emergence and early growth of plantlets up to 21 d of growth. Morphological parameter Average ± SE OCV (%) * Classification of OCV ** Seed weight: 55 ± 3 mg (Group 1) Seed weight: 72 ± 3 mg (Group 2) Seed weight: 89 ± 3 mg (Group 3) Germination (%) 2 d 11.52 ± 0.41c 14.11 ± 0.37b 16.19 ± 0.44a 16.79 Low 4 d 86.33 ± 0.63c 89.82 ± 0.27b 92.67 ± 0.52a 3.55 Low 6 d 95.08 ± 0.48b 96.46 ± 0.61b 98.53 ± 0.43a 1.80 Low Emergence (%) 7 d 87.85 ± 0.4c 89.82 ± 0.27b 93.61 ± 0.43a 3.24 Low 14 d 93.38 ± 0.46b 94.26 ± 0.66b 96.55 ± 0.42a 1.73 Low 21 d 93.98 ± 0.50b 94.75 ± 0.70b 97.86 ± 0.26a 2.15 Low Plant height (cm) 7 d 3.220 ± 0.011b 3.268 ± 0.012b 3.365 ± 0.019a 2.25 Low 14 d 3.809 ± 0.014c 4.030 ± 0.010b 4.245 ± 0.016a 5.41 Low 21 d 5.103 ± 0.017c 5.291 ± 0.007b 5.515 ± 0.025a 3.89 Low Number of leaves 7 d 2.000 ± 0.000a 2.000 ± 0.000a 2.000 ± 0.000a 14 d 2.400 ± 0.163a 2.500 ± 0.167a 2.700 ± 0.153a 21 d 2.800 ± 0.133b 2.800 ± 0.133b 3.200 ± 0.133a 7.87 Low Total leaf weight (g) 7 d 0.128 ± 0.003c 0.155 ± 0.004b 0.215 ± 0.002a 26.83 Medium 14 d 0.202 ± 0.004c 0.234 ± 0.002b 0.273 ± 0.002a 15.05 Low 21 d 0.341 ± 0.005c 0.405 ± 0.006b 0.468 ± 0.004a 15.69 Low Stem length (cm) 7 d 1.115 ± 0.010c 1.140 ± 0.004b 1.200 ± 0.004a 3.79 Low 14 d 1.420 ± 0.012c 1.560 ± 0.010b 1.643 ± 0.009a 7.31 Low 21 d 2.119 ± 0.007c 2.198 ± 0.005b 2.280 ± 0.007a 3.66 Low Stem diameter (cm) 7 d 0.096 ± 0.001c 0.112 ± 0.005b 0.124 ± 0.002a 12.54 Low 14 d 0.185 ± 0.004c 0.230 ± 0.003b 0.267 ± 0.005a 18.06 Low 21 d 0.310 ± 0.005c 0.359 ± 0.007b 0.405 ± 0.007a 13.27 Low Stem weight (g) 7 d 0.136 ± 0.002c 0.151 ± 0.005b 0.175 ± 0.004a 12.77 Low 14 d 0.214 ± 0.003c 0.233 ± 0.004b 0.261 ± 0.003a 10.02 Low 21 d 0.341 ± 0.005c 0.397 ± 0.005b 0.440 ± 0.007a 12.64 Low Main root length (cm) 7 d 1.105 ± 0.003b 1.128 ± 0.009ab 1.165 ± 0.016a 2.67 Low 14 d 1.389 ± 0.010c 1.470 ± 0.005b 1.602 ± 0.010a 7.23 Low 21 d 1.984 ± 0.011c 2.093 ± 0.003b 2.235 ± 0.025a 5.98 Low Main root diameter (cm) 7 d 0.093 ± 0.001c 0.111 ± 0.002b 0.131 ± 0.002a 16.92 Low 14 d 0.179 ± 0.004c 0.219 ± 0.002b 0.254 ± 0.003a 17.27 Low 21 d 0.313 ± 0.006c 0.343 ± 0.006b 0.397 ± 0.006a 12.13 Low Main root weight (g) 7 d 0.130 ± 0.003c 0.151 ± 0.005b 0.179 ± 0.005a 16.03 Low 14 d 0.214 ± 0.003c 0.237 ± 0.005b 0.254 ± 0.003a 8.54 Low 21 d 0.321 ± 0.006c 0.375 ± 0.005b 0.435 ± 0.009a 15.13 Low Results within the same row, with the same letter are not statistically different (One-Way ANOVA, Tukey, p > 0.05). * Overall coefficient of variation = (Standard deviation/Average)*100. To calculate this coefficient, average values were considered. The higher the difference among the three treatments compared the higher the overall coefficient of variation. OCVs were only calculated for those indicators with statistically significant differences according to One-Way ANOVA and Tukey tests. ** Low from 1.73 to 24.42%, Medium from 24.42 to 47.11% and High from 47.11 to 69.79%. Table 2 Biochemical changes recorded up to 21 d of growth. Biochemical parameter Average ± SE OCV (%) * Classification of OCV ** Seed weight: 55 ± 3 mg (Group 1) Seed weight: 72 ± 3 mg (Group 2) Seed weight: 89 ± 3 mg (Group 3) Chlorophyll a (µg · g − 1 fw) 7 d 25.22 ± 0.42c 28.00 ± 0.35b 31.61 ± 0.77a 11.32 Low 14 d 32.42 ± 0.91b 34.64 ± 0.50b 37.46 ± 0.46a 7.26 Low 21 d 34.03 ± 0.53c 37.43 ± 0.49b 42.71 ± 0.69a 11.50 Low Chlorophyll b (µg · g − 1 fw) 7 d 20.85 ± 0.21c 24.21 ± 0.39b 26.09 ± 0.46a 11.20 Low 14 d 25.49 ± 0.28c 28.13 ± 0.31b 31.72 ± 0.52a 11.00 Low 21 d 28.07 ± 0.37c 30.01 ± 0.51b 36.80 ± 0.36a 14.49 Low Total content of proteins (mg · g − 1 fw) 7 d 8.39 ± 0.08c 10.31 ± 0.13b 12.15 ± 0.17a 18.28 Low 14 d 9.80 ± 0.20c 12.02 ± 0.14b 14.21 ± 0.21a 18.36 Low 21 d 11.26 ± 0.31c 13.75 ± 0.25b 18.23 ± 0.37a 24.52 Medium SOD specific activity (U mg − 1 proteins) 7 d 0.244 ± 0.008a 0.098 ± 0.001b 0.079 ± 0.001c 64.35 High 14 d 0.207 ± 0.005a 0.082 ± 0.001b 0.066 ± 0.002c 65.46 High 21 d 0.176 ± 0.006a 0.069 ± 0.002b 0.049 ± 0.001c 69.79 High PER specific activity (U mg − 1 proteins) 7 d 3.227 ± 0.085a 1.877 ± 0.054b 1.521 ± 0.035c 40.75 Medium 14 d 2.68 ± 0.07a 1.62 ± 0.04b 1.22 ± 0.04c 40.76 Medium 21 d 2.25 ± 0.06a 1.28 ± 0.03b 0.87 ± 0.03c 48.27 High Results within the same row, with the same letter are not statistically different (One-Way ANOVA, Tukey, p > 0.05). * Overall coefficient of variation = (Standard deviation/Average)*100. To calculate this coefficient, average values were considered. The higher the difference among the three treatments compared the higher the overall coefficient of variation. OCVs were only calculated for those indicators with statistically significant differences according to One-Way ANOVA and Tukey tests. ** Low from 1.73 to 24.42%, Medium from 24.42 to 47.11% and High from 47.11 to 69.79%. Table 3 Agronomical traits at 3 months of field growth. Morphological parameter Average ± SE OCV (%) * Classification of OCV ** Seed weight: 55 ± 3 mg (Group 1) Seed weight: 72 ± 3 mg (Group 2) Seed weight: 89 ± 3 mg (Group 3) Plant height (cm) 69.37 ± 0.71c 74.71 ± 0.70b 82.85 ± 0.67a 8.97 Low Number of branches 2.70 ± 0.11a 2.80 ± 0.09a 2.95 ± 0.09a Plant fresh weight (g) 35.08 ± 0.58c 39.25 ± 0.75b 45.32 ± 0.57a 12.91 Low Plant dry weight (g) 1.38 ± 0.01c 1.48 ± 0.01b 1.54 ± 0.01a 5.77 Low Time until anthesis of 50% plants (days) 64.00 ± 0.00a 62.00 ± 0.00b 60.00 ± 0.00c 3.23 Low Total number of pods per plant 50.90 ± 1.21c 58.30 ± 2.22b 64.30 ± 0.62a 11.61 Low Number of filled pods per plant 43.75 ± 1.14c 50.95 ± 2.25b 57.05 ± 0.56a 13.16 Low Number of grains per pod 3.65 ± 0.11a 3.70 ± 0.11a 3.75 ± 0.10a Number of grains per plant 159.45 ± 5.76c 187.65 ± 7.86b 213.65 ± 5.59a 14.50 Low Weight of all grains per plant (g) 48.93 ± 0.79c 55.99 ± 0.55b 61.13 ± 0.57a 11.07 Low Weight of 100 grains (g) 33.37 ± 0.50c 39.63 ± 0.64b 44.35 ± 0.63a 14.08 Low Duration of plant cycle (days) 114.00 ± 0.00a 112.00 ± 0.00b 110.00 ± 0.00c 1.79 Low Yield (t/ha) 1.49 ± 0.02c 1.68 ± 0.01b 1.81 ± 0.02a 9.46 Low Results within the same row, with the same letter are not statistically different (One-Way ANOVA, Tukey, p > 0.05). * Overall coefficient of variation = (Standard deviation/Average)*100. To calculate this coefficient, average values were considered. The higher the difference among the three treatments compared the higher the overall coefficient of variation. OCVs were only calculated for those indicators with statistically significant difference according to One-Way ANOVA and Tukey tests. ** Low from 1.73 to 24.42%, Medium from 24.42 to 47.11% and High from 47.11 to 69.79%. Biochemical changes during plant growth and development are outlined in Table 2 . Significant differences were noted for all biochemical indicators assessed. In this context, plants regenerated from seeds of the highest mass had the highest chlorophyll a and b and protein percentage while seeds of the lightest weight morphotype (group 1) produced the highest levels of SOD and PER. When agronomic traits were assessed after 3 months of growth in the field (Table 3 ), the trend observed after 21 d of growth was consistent as plants produced from the heaviest seeds (group 3) maintained faster growth rates compared with the other two seed classes. The relatively slower growth rate of the lightest seeds (group 1) resulted in these plants taking the longest time to reach 50% anthesis in each plot, leading to an extended plant life cycle. The OCVs were calculated as in indicator of meaningful relationships. During the initial stage of germination and plant growth, only one medium OCV was observed where the total leaf weight from plants generated from the heaviest seeds were significantly higher than the other two seed morphs. For the biochemical indicators recorded during plant development (up to 21 d), seven medium and high OCVs were noted. Medium OCVs were recorded for total protein content at 21 d and PER activity at 7 and 14 d. High OCVs were calculated for SOD activity at all time periods and PER at 21 d. The number of species producing heteromorphic seeds is constantly increasing (Lu et al. 2010 ; Leverett and Jolls 2014 ; Bhatt and Santo 2016 ; Hughes 2018 ; Acosta et al. 2020 ). For legumes that show seed heteromorphism, testa size or color may be important traits for predicting seed quality and germination potential (Ochuodho and Modi 2013 ; Dello Jacovo et al. 2019 ). Given the wide range of benefits that Leguminosae species provide to agronomical systems, the morphological indicators associated with heteromorphic seed vigor are particularly important (Acharya et al. 2006 ; Acosta et al. 2020 ). A significant result from the early growth stages (up to 21 d) was that chickpea plants obtained from the group 3 seeds (89 ± 3 mg per seed) displayed a greater mass of leaves than plants from the other two groups after 7 days of growth (Table 1 , medium OCV). This observation could be related to the fact that seeds of a larger size possess greater accumulated reserves which can be used during germination and emergence of plants (leading to more rapid growth and development than seeds with a smaller mass, as noted in the present study). These results are in accordance with theoretical models predicting that large seeds are more likely to exhibit early germination than small seeds (Venable and Brown 1988 ; Rees 1994 ; Cintra 1997 ). Similar results have been reported for other species where larger seeds showed better germination than smaller seeds (Kidson and Westoby 2000 ; Bhatt and Santo 2016 ; Bhatt et al. 2016 ). Variation in seed size is recognized as a type of seed heteromorphism. Mandák and Pyšek ( 2005 ) reported that seeds of Atriplex sagittata with heteromorphic behavior showed variable germination rates with larger seeds germinating faster than smaller seeds. These authors propose that the evolutionary impact of this phenomenon would be dominance of larger seeds as they out-compete smaller seeds in plant populations. Leverett and Jolls ( 2014 ) further proposed that variable mass seed heteromorphism might be advantageous by increasing the number of favorable microsites for germination in species that grow under challenging conditions. Seed heteromorphism is regarded as a bet hedging strategy which describes an evolutionary adaptation to promote survival (at the expense of overall fitness) of a species under variable conditions (Gianella et al. 2021 ). The high OCVs observed between morphs at the biochemical level (for SOD and PER) (Table 2 ) may be related to differences in the physiological state of plants. The results found in chickpeas differ from those observed in Suaeda aralocaspica , where the descendants of heteromorphic seeds did not present significant differences in the accumulation of osmolites, activities of antioxidant enzymes, phosphoenolpyruvate carboxylase and the corresponding gene expression patterns (Xu et al. 2011 ; Gul et al. 2013 ). Berwal and Ram ( 2018 ) have also postulated that plants that retain high levels of antioxidants such as SOD typically display improved tolerance to abiotic stress, and in this regard, SOD has been proposed to be a stable biomarker for abiotic stress tolerance. This warrants further investigation, particularly in the context of the proposed potential seed heteromorphism in chickpea. The differences observed in the reproductive phase (Table 3 ) are related to the better development of plants during the vegetative cycle, associated with a greater availability of assimilate necessary for the reproductive stage. The superior vegetative development of group 3 seeds resulted in a greater leaf area available for the capture of solar radiation for photosynthesis, which optimizes flowering and fruiting in most plant species including legumes (Shibles and Weber 1966 ; Matías and Matías 1995 ; Sakowska et al. 2018 ). The present study reports differences in germination and plant growth in three chickpea morphs based on seed mass. A developmental lag was evident starting from germination up to grain production in the field with the seeds of the largest mass displaying superior growth and production characteristics. These results could be indicative of seed heteromorphic behavior, however, this need to be confirmed in other chickpea materials. Declarations Ethics approval and consent to participate : Informed consent was obtained from all individual participants included in the study. Additional informed consent was obtained from all individual participants for whom identifying information is included in this article. Consent for publication: All authors have read and approved the final manuscript. Availability of data and material: Not applicable. Competing interests: Authors do not have any competing interests. Funding: Not applicable. Authors contributions : AVO, RC, MEMM, DG, YA, BEZB, EH and JCL designed the research; AVO and RC conducted the experiment; AVO, MEMM, DG, YA, BEZB, EH and JCL analyzed the data and wrote the paper; and JCL had primary responsibility for the final content. Acknowledgements: This research was supported by the Bioplant Centre, University of Ciego de Ávila (Cuba); Universidad de Concepción (Chile); Universidad Estatal del Sur de Manabí (Ecuador); and Agricultural Research Council (South Africa). References Acharya SN, Thomas JE, Basu SK (2006) Fenugreek: an “old world” crop for the “new world”. 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Gaur PM, Samineni S, Thudi M, Tripathi S, Sajja SB, Jayalakshmi V, Mannur DM, Vijayakumar AG, Ganga Rao NV, Ojiewo C (2019) Integrated breeding approaches for improving drought and heat adaptation in chickpea ( Cicer arietinum L.). Plant Breed 138:389-400 Gianella M, Bradford K, Guzzon F (2021) Ecological (epi)genetic and physiological aspects of bet-hedging in angiosperms. Plant Reprod 34:21-36 Gross NT, Hultenby K, Mengarelli S (2000) Lipid peroxidation by alveolar macrophages challenged with Cryptococcus neoformans , Candida albicans or Aspergillus fumigatus . Med Mycol 38:443–449 Gul B, Ansari R, Flowers TJ, Khan MA (2013) Germination strategies of halophyte seeds under salinity. Environ Exp Bot 92:4-18 Hayes HK, Immer FR, Smith DC (1955) Methods of Plant Breeding McGraw-Hill Publishing Co Ltd, London. Hughes PW (2018) Minimal-risk seed heteromorphism: proportions of seed morphs for optimal risk-averse heteromorphic strategies. Front Plant Sci https://doiorg/103389/fpls201801412 9:1412 Imbert E (2002) Ecological consequences and ontogeny of seed heteromorphism. Perspect Plant Ecol Evol Syst 5:13-36 ISTA (2005) International Rules for Seed Testing International Seed Testing Association, Bassersdorf, Switzerland. Janghel D, Kumar K, Sunil R, Chhabra A (2020) Genetic diversity analysis, characterization and evaluation of elite chickpea ( Cicer arietinum L.) genotypes. Int J Curr Microbiol App Sci DOI:1020546/ijcmas2020901023 9:199-209 Kaloki P, Trethowan R, Tan DK (2019) Genetic and environmental influences on chickpea water‐use efficiency. J Agron Crop Sci https://doiorg/101111/JAC12338 205:470-476 Kidson R, Westoby M (2000) Seed mass and seedling dimensions in relation to seedling establishment. Oecologia 125:11-17 Kuhn ME (2020) Meet the next generation of plant-based meat. Food Technol 74:24-34 Leverett LD, Jolls CL (2014) Cryptic seed heteromorphism in Packera tomentosa (Asteraceae): differences in mass and germination. Plant Sp Biol 29:169-180 Lorenzo JC, Yabor L, Medina N, Quintana N, Wells V (2015) Coefficient of variation can identify the most important effects of experimental treatments. Not Bot Horti Agrobo Cluj-Nap https://doiorg/1015835/NBHA4319881 43:287-291 Lu J, Tan D, Baskin JM, Baskin CC (2010) Fruit and seed heteromorphism in the cold desert annual ephemeral Diptychocarpus strictus (Brassicaceae) and possible adaptive significance. Ann Bot 105:999-1014 Mandák B, Pyšek P (2005) How does seed heteromorphism influence the life history stages of Atriplex sagittata (Chenopodiaceae)? Flora 200:516-526 Maqbool MA, Aslam M, Ali H (2017) Breeding for improved drought tolerance in chickpea ( Cicer arietinum L.). Plant Breed 136:300-318 Matías C, Matías Y (1995) Efecto de los soportes en la producción de semillas Teramnus labialis , cv. semilla clara. Past Forr 18:7 McCord J, Fridovich I (1969) Superoxide dismutase: an enzymic function for erythrocuprein. J Inorg Biochem 244:6049-6055 Merga B, Haji J, Yildiz F (2019) Economic importance of chickpea: production, value, and world trade. Cog Food Agric 5:1, DOI: 10.1080/23311932.2019.1615718 Moller IM (2001) Plant mitochondria and oxidative stress: electron transport, NADPH turnover, and metabolism of reactive oxygen species. Annual Rev Plant Physiol Plant Mol Biol 52:561–591 Ochuodho J, Modi A (2013) Association of seed coat colour with germination of three wild mustard species with agronomic potential. Afr J Agric Res 8:4354-4359 Pascual M, Pereda C, Pérez R (1983) Inverse correlation between estrogen receptor content and peroxidase activity in human mammary tumors. Neoplasma 30:611-613 Porra R (2002) The chequered history of the development and use of simultaneous equations for the accurate determination of chlorophylls a and b. Photosynth Res 73:149-156 Rees M (1994) Delayed germination of seeds: A look at the effects of adult longevity, the timing of reproduction, and population age/stage structure. Am Nat 144:43- 64 Sakowska K, Alberti G, Genesio L, Peressotti A, Delle Vedove G, Gianelle D, Colombo R, Rodeghiero M, Panigada C, Juszczak R (2018) Leaf and canopy photosynthesis of a chlorophyll deficient soybean mutant. Plant Cell Environ 41:1427-1437 Shagarodsky T, Chiang ML, López Y (2001) Evaluación de cultivares de garbanzo ( Cicer arietinum L.) en Cuba. Agron Mesoam DOI:1015517/AMV12I117298 12:95-98 Shibles R, Weber C (1966) Interception of solar radiation and dry matter production by various soybean planting patterns 1. Crop Sci 6:55-59 Singh KB (1997) Chickpea ( Cicer arietinum L.). Field Crops Res 53:161-170 Trapp EJ (1988) Dispersal of heteromorphic seeds in Amphicarpaea bracteata (Fabaceae). Am J Bot 75:1535-1539 Van der Maessen L (1972) Cicer L., a monograph of the genus, with special reference to the chickpea (Cicer arietinum L.), its ecology and cultivation Wageningen University and Research. Varol IS, Kardes YM, Irik HA, Kirnak H, Kaplan M (2020) Supplementary irrigations at different physiological growth stages of chickpea ( Cicer arietinum L.) change grain nutritional composition. Food Chem DOI:101016/jfoodchem2019125402 303:125402 Venable DL, Brown JS (1988) The selective interactions of dispersal, dormancy, and seed size as adaptations for reducing risk in variable environments. Am Nat 131:360-384 Venable DL, Dyreson E, Pinero D, Becerra JX (1998) Seed morphometrics and adaptive geographic differentiation. Evolution 52:344-354 Xu H, Lu Y, Tong S, Song F (2011) Lipid peroxidation, antioxidant enzyme activity and osmotic adjustment changes in husk leaves of maize in black soils region of Northeast China. African J Agric Res 6:3098-3102 Yaginuma S, Shiraishi T, Ohya H, Igarashi K (2002) Polyphenol increases in cucumber seedlings exposed to strong visible light limited water. Biosci Biotech Biochem 66:65-72 Additional Declarations No competing interests reported. Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-2496069","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":169311534,"identity":"021d7b1d-b642-42dd-84f7-8f1ca911b0f5","order_by":0,"name":"Ariel Villalobos-Olivera","email":"","orcid":"","institution":"University of Ciego de Ávila","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Ariel","middleName":"","lastName":"Villalobos-Olivera","suffix":""},{"id":169311536,"identity":"d6090b54-5946-4bd1-afdd-50f7cd64f146","order_by":1,"name":"Roberto Campbell","email":"","orcid":"","institution":"University of Ciego de Ávila","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Roberto","middleName":"","lastName":"Campbell","suffix":""},{"id":169311537,"identity":"bf41d3e9-0269-4f62-b3b5-19d8eec27e4a","order_by":2,"name":"Marcos Edel Martínez-Montero","email":"","orcid":"","institution":"University of Ciego de Ávila","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Marcos","middleName":"Edel","lastName":"Martínez-Montero","suffix":""},{"id":169311538,"identity":"99db0665-3833-4505-a863-16ea64f0b6c4","order_by":3,"name":"Daviel Gómez","email":"","orcid":"","institution":"Universidad de Concepción","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Daviel","middleName":"","lastName":"Gómez","suffix":""},{"id":169311539,"identity":"90ef1f41-dfd1-4a01-89f0-9af3b3551d78","order_by":4,"name":"Yanier Acosta","email":"","orcid":"","institution":"University of Ciego de Ávila","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Yanier","middleName":"","lastName":"Acosta","suffix":""},{"id":169311540,"identity":"b497ed8b-cd32-451c-9aa0-2a529cf2764d","order_by":5,"name":"Byron E. Zevallos-Bravo","email":"","orcid":"","institution":"Universidad Estatal del Sur de Manabí (UNESUM)","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Byron","middleName":"E.","lastName":"Zevallos-Bravo","suffix":""},{"id":169311541,"identity":"e9e39984-5b32-47bf-9322-16209d2bba42","order_by":6,"name":"Elliosha Hajari","email":"","orcid":"","institution":"Agricultural Research Council-Tropical and Subtropical Crops","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Elliosha","middleName":"","lastName":"Hajari","suffix":""},{"id":169311542,"identity":"22888b4d-4aa5-44dd-9340-c7260ba5fe48","order_by":7,"name":"José Carlos Lorenzo","email":"data:image/png;base64,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","orcid":"","institution":"University of Ciego de Ávila","correspondingAuthor":true,"submittingAuthor":false,"prefix":"","firstName":"José","middleName":"Carlos","lastName":"Lorenzo","suffix":""}],"badges":[],"createdAt":"2023-01-19 14:59:25","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-2496069/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-2496069/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":31970253,"identity":"a4b0f03d-cc23-4fe9-a281-85e0a6054933","added_by":"auto","created_at":"2023-01-23 23:26:21","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":71503,"visible":true,"origin":"","legend":"\u003cp\u003eChickpea phenotypes at 40 d of growth under controlled conditions.\u003c/p\u003e","description":"","filename":"1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-2496069/v1/e2fb0660bf920b88bac02604.jpg"},{"id":38251614,"identity":"bfc2ca4c-3d98-4877-8c3c-9b592304eff0","added_by":"auto","created_at":"2023-06-08 20:29:35","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":529032,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-2496069/v1/db62ae56-9b04-4876-93ae-af2581d11d51.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Chickpea seed mass influences agronomical performance: a case for seed heteromorphism?","fulltext":[{"header":"Introduction","content":"\u003cp\u003eChickpea (\u003cem\u003eCicer arietinum\u003c/em\u003e L.) is an ancient, self-pollinated legume crop believed to have originated in south-eastern Turkey and the adjoining part of Syria (Van der Maessen \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e1972\u003c/span\u003e). It is an annual plant that belongs to the Fabaceae family within the Faboideae subfamily (Shagarodsky et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). Plants are generally about 50 cm in height, with white or violet flowers from which pods develop. Each pod produces two or three seeds at most. Chickpeas are widely consumed by humans, and used as an energy and protein source in animal feed (Duarte-Leal et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Kaloki et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Alam et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Janghel et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Varol et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). An associated advantage of chickpea cultivation is that it improves soil fertility. Most recent estimates are that more than 11\u0026nbsp;million tons enters the world market annually (Merga et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Some chickpea components have shown, in preclinical and clinical studies, several health benefits, including antioxidant capacity, antifungal, antibacterial, analgesic, anticancer, anti-inflammatory and hypocholesterolemic properties (Kuhn \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThere are numerous chickpea landraces available, for example, the International Crops Research Institute for Semi-Arid Tropics (ICRISAT) in India maintains in excess of 20 000 accessions sourced from different countries (Bhagyawant et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Nevertheless, crop improvement is still being pursued with the major goals being to exploit the available genetic potential by developing lines with improved yield (and nutritional characteristics) or to minimize the adverse effects of diseases, insects, drought, heat and cold (Singh \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Chowdhury et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Devasirvatham et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Maqbool et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Gaur et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). While there have been studies over the years on seed developmental characteristics and subsequent growth, which are relevant for breeding programs (Hayes et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e1955\u003c/span\u003e), there appears to be a lack of reports on the hetero-morphism of the seeds and their influence on the germination and growth of chickpea plants.\u003c/p\u003e \u003cp\u003eSeed heteromorphism is an adaptive trait in response to the spatio-temporal variability of the habitat in which plants develop (Venable et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e1998\u003c/span\u003e). This involves the production of seeds with different morphologies and/or post-harvest behavior in different parts of the same plant (Imbert \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Lu et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Differences between morphs are usually based on one or more of the following: color, size, morphology/anatomy, dispersion syndrome, latency, position in the pod or spike, and germination (Baskin and Baskin \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1998\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn the Fabaceae family, seed heteromorphism has been reported in \u003cem\u003eAmphicarpaea bracteata\u003c/em\u003e (Trapp \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e1988\u003c/span\u003e), \u003cem\u003eLathyrus linifolius\u003c/em\u003e (Dello Jacovo et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) and \u003cem\u003eTeramnus labialis\u003c/em\u003e (Acosta et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Considering the growing importance of chickpea in addressing food security challenges in developing countries and as a substitute for meat-based protein in developed regions, it is important to characterize the heteromorphic nature of seeds. The effect of seed heteromorphism can have an influence on species establishment in the agronomical context and yield of plants in the field. This study evaluated the influence of three different seed morphs (55\u0026thinsp;\u0026plusmn;\u0026thinsp;3 mg per seed (group 1); 72\u0026thinsp;\u0026plusmn;\u0026thinsp;3 mg (group 2); 89\u0026thinsp;\u0026plusmn;\u0026thinsp;3 mg (group 3)) on: germination, emergence, early growth of plantlets, and some biochemical indicators: chlorophylls, total protein, superoxide dismutase and peroxidase activities. We selected these compounds because they are related to a wide range of important biochemical and physiological\u003c/p\u003e \u003cp\u003epathways such as plant response to stress and photosynthesis (Gross et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Moller \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Porra \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Yaginuma et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). These parameters were evaluated up to 21 days (d) of growth. Agronomical traits at 3 months after planting were also recorded.\u003c/p\u003e"},{"header":"Materials And Methods","content":"\u003cp\u003eHarvested chickpea seeds (cv. Nacional 29) were air-dried at room temperature to 6% moisture content (fresh mass basis) and then stored for 4 months at 4\u0026deg;C in the dark in hermetically sealed containers. Seeds with 6% moisture content were used in subsequent experiments, as recommended by ISTA (\u003cspan class=\"CitationRef\"\u003e2005\u003c/span\u003e).\u003c/p\u003e\n\u003cp\u003eThree seed sizes of chickpea were compared: 55\u0026thinsp;\u0026plusmn;\u0026thinsp;3 mg per seed (group 1); 72\u0026thinsp;\u0026plusmn;\u0026thinsp;3 mg (group 2); 89\u0026thinsp;\u0026plusmn;\u0026thinsp;3 mg (group 3). Physical characteristics relating to germination (radicle longer than 5 mm), emergence, plant height, number of leaves, total leaf weight; stem length, diameter and weight and main root length, diameter and weight were recorded for up to 21 d. In addition, levels of leaf chlorophyll a and b (Porra \u003cspan class=\"CitationRef\"\u003e2002\u003c/span\u003e), total protein (Bradford \u003cspan class=\"CitationRef\"\u003e1976\u003c/span\u003e), superoxide dismutase (SOD) (McCord and Fridovich \u003cspan class=\"CitationRef\"\u003e1969\u003c/span\u003e) and peroxidase (PER) (Pascual et al. \u003cspan class=\"CitationRef\"\u003e1983\u003c/span\u003e) activities were also recorded. Biochemical determinations were assayed using three independent samples with 100 mg of leaf material per sample.\u003c/p\u003e\n\u003cp\u003eChlorophyll pigments were extracted with 750 \u0026micro;l methanol (100%). Samples were centrifuged (at 10 000 rpm and 4\u0026deg;C for 15 min), supernatants were collected and the absorbance was read at 652.4 nm (Porra \u003cspan class=\"CitationRef\"\u003e2002\u003c/span\u003e). Protein extraction was carried out with Tris-HCl 0.1 mol l\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e buffer, pH\u0026thinsp;=\u0026thinsp;8.5\u0026ndash;8.8, 5 mmol l\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e EDTA and 20 mmol l\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e \u0026beta;-mercaptoethanol. The total protein content was determined according to Bradford (\u003cspan class=\"CitationRef\"\u003e1976\u003c/span\u003e).\u003c/p\u003e\n\u003cp\u003eTo determine superoxide dismutase and guaiacol peroxidase-specific activities, samples were finely ground in liquid nitrogen. Extraction was performed\u0026nbsp;\u003cspan style=\"text-align: inherit;\"\u003ewith Tris\u0026ndash;HCl buffer (0.35 mol l\u003c/span\u003e\u003csup style=\"text-align: inherit;\"\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003cspan style=\"text-align: inherit;\"\u003e, pH 8.0), EDTA (20 mmol l\u003c/span\u003e\u003csup style=\"text-align: inherit;\"\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003cspan style=\"text-align: inherit;\"\u003e), cysteine (15 mmol l\u003c/span\u003e\u003csup style=\"text-align: inherit;\"\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003cspan style=\"text-align: inherit;\"\u003e), PVPP (50%) and PMSF (0.2 mmol l\u003c/span\u003e\u003csup style=\"text-align: inherit;\"\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003cspan style=\"text-align: inherit;\"\u003e). Samples were homogenized on ice\u0026nbsp;\u003c/span\u003e\u003cspan style=\"text-align: inherit;\"\u003ewith polytron apparatus (Ultra-turrax T25). The homogenate was filtered through a two-folded piece of gauze and centrifuged at 10,290 xg (Beckman J-21, Palo Alto, CA) for\u0026nbsp;\u003c/span\u003e\u003cspan style=\"text-align: inherit;\"\u003e30 min at 4\u0026deg;C. Superoxide dismutase activity was measured according to McCord and Fridovich (\u003c/span\u003e\u003cspan class=\"CitationRef\" style=\"text-align: inherit;\"\u003e1969\u003c/span\u003e\u003cspan style=\"text-align: inherit;\"\u003e). The reaction mixtures included 80 \u0026micro;l of plant extract and\u0026nbsp;\u003c/span\u003e\u003cspan style=\"text-align: inherit;\"\u003e900 \u0026micro;l of potassium phosphate\u0026ndash;KOH (50 mmol l\u003c/span\u003e\u003csup style=\"text-align: inherit;\"\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003cspan style=\"text-align: inherit;\"\u003e, pH 7.6), EDTA (0.1 mmol l\u003c/span\u003e\u003csup style=\"text-align: inherit;\"\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003cspan style=\"text-align: inherit;\"\u003e); cytochrome C (0.01 mmol l\u003c/span\u003e\u003csup style=\"text-align: inherit;\"\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003cspan style=\"text-align: inherit;\"\u003e), xanthine (0.05 mmol l\u003c/span\u003e\u003csup style=\"text-align: inherit;\"\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003cspan style=\"text-align: inherit;\"\u003e) and 20 \u0026micro;l xanthine oxidase (EC 1.2.3.22; 0.03 units). Absorbance (550 nm) was measured every 15 s for 3 min. Averages of linear section absorbance were used and cytochrome c extinction coefficient (21.1 mmol\u003c/span\u003e\u003csup style=\"text-align: inherit;\"\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003cspan style=\"text-align: inherit;\"\u003e l cm\u003c/span\u003e\u003csup style=\"text-align: inherit;\"\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003cspan style=\"text-align: inherit;\"\u003e) was used. Superoxide dismutase activity was defined as U g\u003c/span\u003e\u003csup style=\"text-align: inherit;\"\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e\u003cspan style=\"text-align: inherit;\"\u003e plant fresh mass [U\u0026thinsp;=\u0026thinsp;enzyme quantity hydrolyzing 1 \u0026micro;mol of superoxide\u0026nbsp;\u003c/span\u003e\u003cspan style=\"text-align: inherit;\"\u003eper hour (37\u0026deg;C)]. Specific activity was calculated as the rate of superoxide dismutase activity relative to protein content.\u003c/span\u003e\u003c/p\u003e\n\u003cp\u003eGuaiacol peroxidase activity was measured according to Pascual et al. (\u003cspan class=\"CitationRef\"\u003e1983\u003c/span\u003e). The reaction mixtures included 100 \u0026micro;l of plant material extract, 1.0 ml Tris\u0026ndash;HCl buffer (0.01 mol l\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e, pH 7.0), 150 \u0026micro;l guaiacol (100 mmol l\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) and 20 \u0026micro;l hydrogen peroxide. Absorbance (470 nm) was measured every 15 s for 3 min. Averages of linear section absorbance were used and guaiacol extinction coefficient (5,570 x 10\u003csup\u003e\u0026minus;\u0026thinsp;6\u003c/sup\u003e \u0026micro;mol\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e l cm\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e) was used. Guaiacol peroxidase activity was defined as U g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e plant fresh mass [U\u0026thinsp;=\u0026thinsp;enzyme quantity hydrolyzing 1 \u0026micro;mol substrate per minute (37\u0026deg;C)]. Specific activity was calculated as the rate of guaiacol peroxidase activity relative to protein content.\u003c/p\u003e\n\u003cp\u003eThe following agronomical traits were also measured during three months of growth in the field, as recommended by the International Board for Plant Genetic Resources (FAO/IPGRI \u003cspan class=\"CitationRef\"\u003e1994\u003c/span\u003e): plant height, fresh and dry weight; number of branches; time until anthesis in 50% of plants; total number of pods; number of filled pods; number of grains per pod, number and weight of grains per plant; weight of 100 grains; duration of plant cycle, and yield.\u003c/p\u003e\n\u003cp\u003eGermination was assessed by placing the three seed morphs on filter paper in Petri dishes (\u0026Oslash;: 10 cm moistened with 15 mL of distilled water and five replicates of 10 seeds per dish). To evaluate young plantlets up to 21 d of growth, seeds were planted into pots (500 mL volume, one seed per pot) containing Ferralytic-red soil and filter-cake-sugarcane ash (1:1, v:v). There were five replicates of 10 pots for each treatment (50 seeds per treatment). To study adult plants, 90 seeds of each seed morph were randomly selected and sown in a plant bed (with Ferralytic-red soil and filter-cake-sugarcane ashes) under field conditions. Seeds were planted 70 x 25 cm apart (3 replicates of 30 seeds each) in a randomized complete block design. The experiment was conducted at the Field Experimental Station of the Bioplant Centre, University of Ciego de \u0026Aacute;vila, Cuba (21 52\u0026acute;48.6\u0026acute;\u0026acute; N, 78 41\u0026acute;32.6\u0026acute;\u0026acute; W; 53 meters above sea level; Sept 2020 \u0026ndash; Jan 2021). Temperatures at 13:00 averaged 33\u003csup\u003eo\u003c/sup\u003eC during the experiment; relative humidity reached 80%; and the Photosynthetic Photon Flux was 1 140 \u0026micro;mol s\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e. Technical instructions as provided by the Cuban Ministry for Agriculture to cultivate chickpea were applied. Fertilizers were not supplied. Seeds were not inoculated with \u003cem\u003eRhizobium\u003c/em\u003e. Microjet irrigation was used to water the plants for 5 min every 8 h. Border plants, which had more space to grow, were not considered.\u003c/p\u003e\n\u003cp\u003eAll statistical analyses (One-Way ANOVA and Tukey, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.05) were carried out using SPSS (Version 8.0 for Windows, SPSS Inc., New York). The overall coefficients of variation (OCV) were calculated as follows: (standard deviation/average) * 100. In this formula, we considered the average values of the three treatments compared (seed sizes) to calculate the standard deviation and average. Therefore, the higher the difference between the three materials compared, the higher the OCV (Lorenzo et al. \u003cspan class=\"CitationRef\"\u003e2015\u003c/span\u003e). The OCVs were classified as \u003cem\u003eLow\u003c/em\u003e from 1.73 to 24.42%, \u003cem\u003eMedium\u003c/em\u003e from 24.42 to 47.11% and \u003cem\u003eHigh\u003c/em\u003e from 47.11 to 69.79%. The OCVs were only calculated for those indicators with statistically significant differences according to ANOVA and Tukey tests.\u003c/p\u003e"},{"header":"Results And Discussion","content":"\u003cp\u003eThe current study investigated the growth and developmental responses of three seed morphs of chickpea based on seed mass. Tables\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e\u0026ndash;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e summarize the results for the phenotypical growth (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) and biochemical attributes (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) of plants from all three morphs at intervals up to 21 d and also agronomic characteristics following 3 months of growth in the field (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). A total of 61 indicators were evaluated across the study with 33 related to germination, emergence and seedling establishment up to 21 d (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e), 13 for biochemical evaluations (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) and an additional 13 for agronomical traits (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). During the early stages of plant growth (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e), significant differences were found in the majority of parameters assessed (i.e. 31 out of 33) and in all cases, group 3 seeds (seeds with the highest mass) yielded superior plant growth characteristics compared with seeds from the other two groups. This was evident in Fig.\u0026nbsp;1 which demonstrated the faster growth of the heaviest seeds (group 3) followed by intermediate growth of group 2 seeds and the smallest plants were produced by the lightest seeds (group 1).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eGermination, emergence and early growth of plantlets up to 21 d of growth.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eMorphological parameter\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colspan=\"3\" nameend=\"c5\" namest=\"c3\"\u003e \u003cp\u003eAverage\u0026thinsp;\u0026plusmn;\u0026thinsp;SE\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eOCV (%)\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eClassification of OCV\u003csup\u003e**\u003c/sup\u003e\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSeed weight: 55\u0026thinsp;\u0026plusmn;\u0026thinsp;3 mg (Group 1)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSeed weight: 72\u0026thinsp;\u0026plusmn;\u0026thinsp;3 mg (Group 2)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSeed weight: 89\u0026thinsp;\u0026plusmn;\u0026thinsp;3 mg (Group 3)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eGermination (%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e11.52\u0026thinsp;\u0026plusmn;\u0026thinsp;0.41c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e14.11\u0026thinsp;\u0026plusmn;\u0026thinsp;0.37b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e16.19\u0026thinsp;\u0026plusmn;\u0026thinsp;0.44a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e16.79\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e4 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e86.33\u0026thinsp;\u0026plusmn;\u0026thinsp;0.63c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e89.82\u0026thinsp;\u0026plusmn;\u0026thinsp;0.27b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e92.67\u0026thinsp;\u0026plusmn;\u0026thinsp;0.52a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e3.55\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e6 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e95.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.48b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e96.46\u0026thinsp;\u0026plusmn;\u0026thinsp;0.61b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e98.53\u0026thinsp;\u0026plusmn;\u0026thinsp;0.43a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.80\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eEmergence (%)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e7 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e87.85\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e89.82\u0026thinsp;\u0026plusmn;\u0026thinsp;0.27b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e93.61\u0026thinsp;\u0026plusmn;\u0026thinsp;0.43a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e3.24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e14 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e93.38\u0026thinsp;\u0026plusmn;\u0026thinsp;0.46b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e94.26\u0026thinsp;\u0026plusmn;\u0026thinsp;0.66b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e96.55\u0026thinsp;\u0026plusmn;\u0026thinsp;0.42a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.73\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e21 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e93.98\u0026thinsp;\u0026plusmn;\u0026thinsp;0.50b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e94.75\u0026thinsp;\u0026plusmn;\u0026thinsp;0.70b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e97.86\u0026thinsp;\u0026plusmn;\u0026thinsp;0.26a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2.15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003ePlant height (cm)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e7 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3.220\u0026thinsp;\u0026plusmn;\u0026thinsp;0.011b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3.268\u0026thinsp;\u0026plusmn;\u0026thinsp;0.012b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3.365\u0026thinsp;\u0026plusmn;\u0026thinsp;0.019a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e14 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3.809\u0026thinsp;\u0026plusmn;\u0026thinsp;0.014c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e4.030\u0026thinsp;\u0026plusmn;\u0026thinsp;0.010b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e4.245\u0026thinsp;\u0026plusmn;\u0026thinsp;0.016a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e5.41\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e21 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e5.103\u0026thinsp;\u0026plusmn;\u0026thinsp;0.017c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e5.291\u0026thinsp;\u0026plusmn;\u0026thinsp;0.007b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e5.515\u0026thinsp;\u0026plusmn;\u0026thinsp;0.025a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e3.89\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eNumber of leaves\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e7 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.000\u0026thinsp;\u0026plusmn;\u0026thinsp;0.000a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.000\u0026thinsp;\u0026plusmn;\u0026thinsp;0.000a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.000\u0026thinsp;\u0026plusmn;\u0026thinsp;0.000a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e14 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.400\u0026thinsp;\u0026plusmn;\u0026thinsp;0.163a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.500\u0026thinsp;\u0026plusmn;\u0026thinsp;0.167a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.700\u0026thinsp;\u0026plusmn;\u0026thinsp;0.153a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e21 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.800\u0026thinsp;\u0026plusmn;\u0026thinsp;0.133b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.800\u0026thinsp;\u0026plusmn;\u0026thinsp;0.133b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3.200\u0026thinsp;\u0026plusmn;\u0026thinsp;0.133a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e7.87\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eTotal leaf weight (g)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e7 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.128\u0026thinsp;\u0026plusmn;\u0026thinsp;0.003c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.155\u0026thinsp;\u0026plusmn;\u0026thinsp;0.004b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.215\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e26.83\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eMedium\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e14 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.202\u0026thinsp;\u0026plusmn;\u0026thinsp;0.004c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.234\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.273\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e15.05\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e21 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.341\u0026thinsp;\u0026plusmn;\u0026thinsp;0.005c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.405\u0026thinsp;\u0026plusmn;\u0026thinsp;0.006b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.468\u0026thinsp;\u0026plusmn;\u0026thinsp;0.004a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e15.69\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eStem length (cm)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e7 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.115\u0026thinsp;\u0026plusmn;\u0026thinsp;0.010c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.140\u0026thinsp;\u0026plusmn;\u0026thinsp;0.004b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.200\u0026thinsp;\u0026plusmn;\u0026thinsp;0.004a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e3.79\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e14 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.420\u0026thinsp;\u0026plusmn;\u0026thinsp;0.012c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.560\u0026thinsp;\u0026plusmn;\u0026thinsp;0.010b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.643\u0026thinsp;\u0026plusmn;\u0026thinsp;0.009a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e7.31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e21 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.119\u0026thinsp;\u0026plusmn;\u0026thinsp;0.007c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.198\u0026thinsp;\u0026plusmn;\u0026thinsp;0.005b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.280\u0026thinsp;\u0026plusmn;\u0026thinsp;0.007a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e3.66\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eStem diameter (cm)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e7 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.096\u0026thinsp;\u0026plusmn;\u0026thinsp;0.001c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.112\u0026thinsp;\u0026plusmn;\u0026thinsp;0.005b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.124\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e12.54\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e14 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.185\u0026thinsp;\u0026plusmn;\u0026thinsp;0.004c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.230\u0026thinsp;\u0026plusmn;\u0026thinsp;0.003b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.267\u0026thinsp;\u0026plusmn;\u0026thinsp;0.005a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e18.06\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e21 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.310\u0026thinsp;\u0026plusmn;\u0026thinsp;0.005c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.359\u0026thinsp;\u0026plusmn;\u0026thinsp;0.007b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.405\u0026thinsp;\u0026plusmn;\u0026thinsp;0.007a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e13.27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eStem weight (g)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e7 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.136\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.151\u0026thinsp;\u0026plusmn;\u0026thinsp;0.005b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.175\u0026thinsp;\u0026plusmn;\u0026thinsp;0.004a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e12.77\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e14 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.214\u0026thinsp;\u0026plusmn;\u0026thinsp;0.003c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.233\u0026thinsp;\u0026plusmn;\u0026thinsp;0.004b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.261\u0026thinsp;\u0026plusmn;\u0026thinsp;0.003a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e10.02\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e21 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.341\u0026thinsp;\u0026plusmn;\u0026thinsp;0.005c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.397\u0026thinsp;\u0026plusmn;\u0026thinsp;0.005b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.440\u0026thinsp;\u0026plusmn;\u0026thinsp;0.007a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e12.64\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eMain root length (cm)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e7 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.105\u0026thinsp;\u0026plusmn;\u0026thinsp;0.003b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.128\u0026thinsp;\u0026plusmn;\u0026thinsp;0.009ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.165\u0026thinsp;\u0026plusmn;\u0026thinsp;0.016a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2.67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e14 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.389\u0026thinsp;\u0026plusmn;\u0026thinsp;0.010c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.470\u0026thinsp;\u0026plusmn;\u0026thinsp;0.005b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.602\u0026thinsp;\u0026plusmn;\u0026thinsp;0.010a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e7.23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e21 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.984\u0026thinsp;\u0026plusmn;\u0026thinsp;0.011c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.093\u0026thinsp;\u0026plusmn;\u0026thinsp;0.003b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.235\u0026thinsp;\u0026plusmn;\u0026thinsp;0.025a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e5.98\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eMain root diameter (cm)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e7 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.093\u0026thinsp;\u0026plusmn;\u0026thinsp;0.001c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.111\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.131\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e16.92\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e14 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.179\u0026thinsp;\u0026plusmn;\u0026thinsp;0.004c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.219\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.254\u0026thinsp;\u0026plusmn;\u0026thinsp;0.003a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e17.27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e21 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.313\u0026thinsp;\u0026plusmn;\u0026thinsp;0.006c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.343\u0026thinsp;\u0026plusmn;\u0026thinsp;0.006b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.397\u0026thinsp;\u0026plusmn;\u0026thinsp;0.006a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e12.13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eMain root weight (g)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e7 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.130\u0026thinsp;\u0026plusmn;\u0026thinsp;0.003c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.151\u0026thinsp;\u0026plusmn;\u0026thinsp;0.005b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.179\u0026thinsp;\u0026plusmn;\u0026thinsp;0.005a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e16.03\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e14 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.214\u0026thinsp;\u0026plusmn;\u0026thinsp;0.003c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.237\u0026thinsp;\u0026plusmn;\u0026thinsp;0.005b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.254\u0026thinsp;\u0026plusmn;\u0026thinsp;0.003a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e8.54\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e21 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.321\u0026thinsp;\u0026plusmn;\u0026thinsp;0.006c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.375\u0026thinsp;\u0026plusmn;\u0026thinsp;0.005b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.435\u0026thinsp;\u0026plusmn;\u0026thinsp;0.009a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e15.13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"7\" nameend=\"c7\" namest=\"c1\"\u003e \u003cp\u003eResults within the same row, with the same \u003cem\u003eletter\u003c/em\u003e are not statistically different (One-Way ANOVA, Tukey, p\u0026thinsp;\u0026gt;\u0026thinsp;0.05).\u003c/p\u003e \u003cp\u003e\u003csup\u003e*\u003c/sup\u003e Overall coefficient of variation = (Standard deviation/Average)*100. To calculate this coefficient, average values were considered. The higher the difference among the three treatments compared the higher the overall coefficient of variation. OCVs were only calculated for those indicators with statistically significant differences according to One-Way ANOVA and Tukey tests.\u003c/p\u003e \u003cp\u003e\u003csup\u003e**\u003c/sup\u003e \u003cem\u003eLow\u003c/em\u003e from 1.73 to 24.42%, \u003cem\u003eMedium\u003c/em\u003e from 24.42 to 47.11% and \u003cem\u003eHigh\u003c/em\u003e from 47.11 to 69.79%.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eBiochemical changes recorded up to 21 d of growth.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eBiochemical parameter\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colspan=\"3\" nameend=\"c5\" namest=\"c3\"\u003e \u003cp\u003eAverage\u0026thinsp;\u0026plusmn;\u0026thinsp;SE\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eOCV (%)\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eClassification of OCV\u003csup\u003e**\u003c/sup\u003e\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSeed weight: 55\u0026thinsp;\u0026plusmn;\u0026thinsp;3 mg (Group 1)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSeed weight: 72\u0026thinsp;\u0026plusmn;\u0026thinsp;3 mg (Group 2)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSeed weight: 89\u0026thinsp;\u0026plusmn;\u0026thinsp;3 mg (Group 3)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eChlorophyll a (\u0026micro;g \u0026middot; g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e fw)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e7 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e25.22\u0026thinsp;\u0026plusmn;\u0026thinsp;0.42c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e28.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.35b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e31.61\u0026thinsp;\u0026plusmn;\u0026thinsp;0.77a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e11.32\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e14 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e32.42\u0026thinsp;\u0026plusmn;\u0026thinsp;0.91b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e34.64\u0026thinsp;\u0026plusmn;\u0026thinsp;0.50b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e37.46\u0026thinsp;\u0026plusmn;\u0026thinsp;0.46a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e7.26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e21 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e34.03\u0026thinsp;\u0026plusmn;\u0026thinsp;0.53c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e37.43\u0026thinsp;\u0026plusmn;\u0026thinsp;0.49b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e42.71\u0026thinsp;\u0026plusmn;\u0026thinsp;0.69a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e11.50\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eChlorophyll b (\u0026micro;g \u0026middot; g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e fw)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e7 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e20.85\u0026thinsp;\u0026plusmn;\u0026thinsp;0.21c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e24.21\u0026thinsp;\u0026plusmn;\u0026thinsp;0.39b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e26.09\u0026thinsp;\u0026plusmn;\u0026thinsp;0.46a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e11.20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e14 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e25.49\u0026thinsp;\u0026plusmn;\u0026thinsp;0.28c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e28.13\u0026thinsp;\u0026plusmn;\u0026thinsp;0.31b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e31.72\u0026thinsp;\u0026plusmn;\u0026thinsp;0.52a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e11.00\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e21 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e28.07\u0026thinsp;\u0026plusmn;\u0026thinsp;0.37c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e30.01\u0026thinsp;\u0026plusmn;\u0026thinsp;0.51b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e36.80\u0026thinsp;\u0026plusmn;\u0026thinsp;0.36a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e14.49\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eTotal content of proteins (mg \u0026middot; g\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e fw)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e7 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e8.39\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e10.31\u0026thinsp;\u0026plusmn;\u0026thinsp;0.13b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e12.15\u0026thinsp;\u0026plusmn;\u0026thinsp;0.17a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e18.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e14 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e9.80\u0026thinsp;\u0026plusmn;\u0026thinsp;0.20c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e12.02\u0026thinsp;\u0026plusmn;\u0026thinsp;0.14b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e14.21\u0026thinsp;\u0026plusmn;\u0026thinsp;0.21a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e18.36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e21 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e11.26\u0026thinsp;\u0026plusmn;\u0026thinsp;0.31c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e13.75\u0026thinsp;\u0026plusmn;\u0026thinsp;0.25b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e18.23\u0026thinsp;\u0026plusmn;\u0026thinsp;0.37a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e24.52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eMedium\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eSOD specific activity (U mg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e proteins)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e7 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.244\u0026thinsp;\u0026plusmn;\u0026thinsp;0.008a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.098\u0026thinsp;\u0026plusmn;\u0026thinsp;0.001b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.079\u0026thinsp;\u0026plusmn;\u0026thinsp;0.001c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e64.35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eHigh\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e14 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.207\u0026thinsp;\u0026plusmn;\u0026thinsp;0.005a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.082\u0026thinsp;\u0026plusmn;\u0026thinsp;0.001b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.066\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e65.46\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eHigh\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e21 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.176\u0026thinsp;\u0026plusmn;\u0026thinsp;0.006a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.069\u0026thinsp;\u0026plusmn;\u0026thinsp;0.002b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.049\u0026thinsp;\u0026plusmn;\u0026thinsp;0.001c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e69.79\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eHigh\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003ePER specific activity (U mg\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e proteins)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e7 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3.227\u0026thinsp;\u0026plusmn;\u0026thinsp;0.085a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.877\u0026thinsp;\u0026plusmn;\u0026thinsp;0.054b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.521\u0026thinsp;\u0026plusmn;\u0026thinsp;0.035c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e40.75\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eMedium\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e14 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.68\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.62\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.22\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e40.76\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eMedium\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e21 d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.25\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.28\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.87\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e48.27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u003cem\u003eHigh\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"7\" nameend=\"c7\" namest=\"c1\"\u003e \u003cp\u003eResults within the same row, with the same \u003cem\u003eletter\u003c/em\u003e are not statistically different (One-Way ANOVA, Tukey, p\u0026thinsp;\u0026gt;\u0026thinsp;0.05).\u003c/p\u003e \u003cp\u003e\u003csup\u003e*\u003c/sup\u003e Overall coefficient of variation = (Standard deviation/Average)*100. To calculate this coefficient, average values were considered. The higher the difference among the three treatments compared the higher the overall coefficient of variation. OCVs were only calculated for those indicators with statistically significant differences according to One-Way ANOVA and Tukey tests.\u003c/p\u003e \u003cp\u003e\u003csup\u003e**\u003c/sup\u003e \u003cem\u003eLow\u003c/em\u003e from 1.73 to 24.42%, \u003cem\u003eMedium\u003c/em\u003e from 24.42 to 47.11% and \u003cem\u003eHigh\u003c/em\u003e from 47.11 to 69.79%.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eAgronomical traits at 3 months of field growth.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eMorphological parameter\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"3\" nameend=\"c4\" namest=\"c2\"\u003e \u003cp\u003eAverage\u0026thinsp;\u0026plusmn;\u0026thinsp;SE\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eOCV (%)\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eClassification of OCV\u003csup\u003e**\u003c/sup\u003e\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSeed weight: 55\u0026thinsp;\u0026plusmn;\u0026thinsp;3 mg (Group 1)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSeed weight: 72\u0026thinsp;\u0026plusmn;\u0026thinsp;3 mg (Group 2)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSeed weight: 89\u0026thinsp;\u0026plusmn;\u0026thinsp;3 mg (Group 3)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePlant height (cm)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e69.37\u0026thinsp;\u0026plusmn;\u0026thinsp;0.71c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e74.71\u0026thinsp;\u0026plusmn;\u0026thinsp;0.70b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e82.85\u0026thinsp;\u0026plusmn;\u0026thinsp;0.67a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e8.97\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eNumber of branches\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2.70\u0026thinsp;\u0026plusmn;\u0026thinsp;0.11a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.80\u0026thinsp;\u0026plusmn;\u0026thinsp;0.09a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.95\u0026thinsp;\u0026plusmn;\u0026thinsp;0.09a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePlant fresh weight (g)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e35.08\u0026thinsp;\u0026plusmn;\u0026thinsp;0.58c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e39.25\u0026thinsp;\u0026plusmn;\u0026thinsp;0.75b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e45.32\u0026thinsp;\u0026plusmn;\u0026thinsp;0.57a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e12.91\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePlant dry weight (g)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.38\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.48\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.54\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e5.77\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTime until anthesis of 50% plants (days)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e64.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e62.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e60.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3.23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTotal number of pods per plant\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e50.90\u0026thinsp;\u0026plusmn;\u0026thinsp;1.21c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e58.30\u0026thinsp;\u0026plusmn;\u0026thinsp;2.22b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e64.30\u0026thinsp;\u0026plusmn;\u0026thinsp;0.62a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e11.61\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eNumber of filled pods per plant\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e43.75\u0026thinsp;\u0026plusmn;\u0026thinsp;1.14c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e50.95\u0026thinsp;\u0026plusmn;\u0026thinsp;2.25b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e57.05\u0026thinsp;\u0026plusmn;\u0026thinsp;0.56a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e13.16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eNumber of grains per pod\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e3.65\u0026thinsp;\u0026plusmn;\u0026thinsp;0.11a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3.70\u0026thinsp;\u0026plusmn;\u0026thinsp;0.11a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3.75\u0026thinsp;\u0026plusmn;\u0026thinsp;0.10a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eNumber of grains per plant\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e159.45\u0026thinsp;\u0026plusmn;\u0026thinsp;5.76c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e187.65\u0026thinsp;\u0026plusmn;\u0026thinsp;7.86b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e213.65\u0026thinsp;\u0026plusmn;\u0026thinsp;5.59a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e14.50\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWeight of all grains per plant (g)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e48.93\u0026thinsp;\u0026plusmn;\u0026thinsp;0.79c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e55.99\u0026thinsp;\u0026plusmn;\u0026thinsp;0.55b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e61.13\u0026thinsp;\u0026plusmn;\u0026thinsp;0.57a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e11.07\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWeight of 100 grains (g)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e33.37\u0026thinsp;\u0026plusmn;\u0026thinsp;0.50c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e39.63\u0026thinsp;\u0026plusmn;\u0026thinsp;0.64b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e44.35\u0026thinsp;\u0026plusmn;\u0026thinsp;0.63a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e14.08\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDuration of plant cycle (days)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e114.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e112.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e110.00\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.79\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eYield (t/ha)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.49\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.68\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.81\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e9.46\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u003cem\u003eLow\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"6\" nameend=\"c6\" namest=\"c1\"\u003e \u003cp\u003eResults within the same row, with the same \u003cem\u003eletter\u003c/em\u003e are not statistically different (One-Way ANOVA, Tukey, p\u0026thinsp;\u0026gt;\u0026thinsp;0.05).\u003c/p\u003e \u003cp\u003e\u003csup\u003e*\u003c/sup\u003e Overall coefficient of variation = (Standard deviation/Average)*100. To calculate this coefficient, average values were considered. The higher the difference among the three treatments compared the higher the overall coefficient of variation. OCVs were only calculated for those indicators with statistically significant difference according to One-Way ANOVA and Tukey tests.\u003c/p\u003e \u003cp\u003e\u003csup\u003e**\u003c/sup\u003e \u003cem\u003eLow\u003c/em\u003e from 1.73 to 24.42%, \u003cem\u003eMedium\u003c/em\u003e from 24.42 to 47.11% and \u003cem\u003eHigh\u003c/em\u003e from 47.11 to 69.79%.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eBiochemical changes during plant growth and development are outlined in Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e. Significant differences were noted for all biochemical indicators assessed. In this context, plants regenerated from seeds of the highest mass had the highest chlorophyll a and b and protein percentage while seeds of the lightest weight morphotype (group 1) produced the highest levels of SOD and PER. When agronomic traits were assessed after 3 months of growth in the field (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e), the trend observed after 21 d of growth was consistent as plants produced from the heaviest seeds (group 3) maintained faster growth rates compared with the other two seed classes. The relatively slower growth rate of the lightest seeds (group 1) resulted in these plants taking the longest time to reach 50% anthesis in each plot, leading to an extended plant life cycle.\u003c/p\u003e \u003cp\u003eThe OCVs were calculated as in indicator of meaningful relationships. During the initial stage of germination and plant growth, only one medium OCV was observed where the total leaf weight from plants generated from the heaviest seeds were significantly higher than the other two seed morphs. For the biochemical indicators recorded during plant development (up to 21 d), seven medium and high OCVs were noted. Medium OCVs were recorded for total protein content at 21 d and PER activity at 7 and 14 d. High OCVs were calculated for SOD activity at all time periods and PER at 21 d.\u003c/p\u003e \u003cp\u003eThe number of species producing heteromorphic seeds is constantly increasing (Lu et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Leverett and Jolls \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Bhatt and Santo \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Hughes \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Acosta et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). For legumes that show seed heteromorphism, testa size or color may be important traits for predicting seed quality and germination potential (Ochuodho and Modi \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Dello Jacovo et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Given the wide range of benefits that Leguminosae species provide to agronomical systems, the morphological indicators associated with heteromorphic seed vigor are particularly important (Acharya et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Acosta et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eA significant result from the early growth stages (up to 21 d) was that chickpea plants obtained from the group 3 seeds (89\u0026thinsp;\u0026plusmn;\u0026thinsp;3 mg per seed) displayed a greater mass of leaves than plants from the other two groups after 7 days of growth (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, medium OCV). This observation could be related to the fact that seeds of a larger size possess greater accumulated reserves which can be used during germination and emergence of plants (leading to more rapid growth and development than seeds with a smaller mass, as noted in the present study). These results are in accordance with theoretical models predicting that large seeds are more likely to exhibit early germination than small seeds (Venable and Brown \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e1988\u003c/span\u003e; Rees \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; Cintra \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e1997\u003c/span\u003e). Similar results have been reported for other species where larger seeds showed better germination than smaller seeds (Kidson and Westoby \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Bhatt and Santo \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Bhatt et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2016\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eVariation in seed size is recognized as a type of seed heteromorphism. Mand\u0026aacute;k and Pyšek (\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2005\u003c/span\u003e) reported that seeds of \u003cem\u003eAtriplex sagittata\u003c/em\u003e with heteromorphic behavior showed variable germination rates with larger seeds germinating faster than smaller seeds. These authors propose that the evolutionary impact of this phenomenon would be dominance of larger seeds as they out-compete smaller seeds in plant populations. Leverett and Jolls (\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2014\u003c/span\u003e) further proposed that variable mass seed heteromorphism might be advantageous by increasing the number of favorable microsites for germination in species that grow under challenging conditions. Seed heteromorphism is regarded as a bet hedging strategy which describes an evolutionary adaptation to promote survival (at the expense of overall fitness) of a species under variable conditions (Gianella et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe high OCVs observed between morphs at the biochemical level (for SOD and PER) (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) may be related to differences in the physiological state of plants. The results found in chickpeas differ from those observed in \u003cem\u003eSuaeda aralocaspica\u003c/em\u003e, where the descendants of heteromorphic seeds did not present significant differences in the accumulation of osmolites, activities of antioxidant enzymes, phosphoenolpyruvate carboxylase and the corresponding gene expression patterns (Xu et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Gul et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Berwal and Ram (\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) have also postulated that plants that retain high levels of antioxidants such as SOD typically display improved tolerance to abiotic stress, and in this regard, SOD has been proposed to be a stable biomarker for abiotic stress tolerance. This warrants further investigation, particularly in the context of the proposed potential seed heteromorphism in chickpea.\u003c/p\u003e \u003cp\u003eThe differences observed in the reproductive phase (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e) are related to the better development of plants during the vegetative cycle, associated with a greater availability of assimilate necessary for the reproductive stage. The superior vegetative development of group 3 seeds resulted in a greater leaf area available for the capture of solar radiation for photosynthesis, which optimizes flowering and fruiting in most plant species including legumes (Shibles and Weber \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e1966\u003c/span\u003e; Mat\u0026iacute;as and Mat\u0026iacute;as \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Sakowska et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe present study reports differences in germination and plant growth in three chickpea morphs based on seed mass. A developmental lag was evident starting from germination up to grain production in the field with the seeds of the largest mass displaying superior growth and production characteristics. These results could be indicative of seed heteromorphic behavior, however, this need to be confirmed in other chickpea materials.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eEthics approval and consent to participate\u003c/strong\u003e: Informed consent was obtained from all individual participants included in the study. Additional informed consent was obtained from all individual participants for whom identifying information is included in this article.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent for publication:\u003c/strong\u003e All authors have read and approved the final manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAvailability of data and material:\u003c/strong\u003e Not applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests:\u003c/strong\u003e Authors do not have any competing interests.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding:\u003c/strong\u003e Not applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthors contributions\u003c/strong\u003e:\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eAVO, RC, MEMM, DG, YA, BEZB, EH\u0026nbsp;and JCL\u003csup\u003e\u0026nbsp;\u003c/sup\u003edesigned the research;\u0026nbsp;AVO and RC\u0026nbsp;conducted the experiment;\u0026nbsp;AVO, MEMM, DG, YA, BEZB, EH and JCL\u003csup\u003e\u0026nbsp;\u003c/sup\u003eanalyzed the data and wrote the paper; and\u0026nbsp;JCL\u003csup\u003e\u0026nbsp;\u003c/sup\u003ehad primary responsibility for the final content.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAcknowledgements:\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis research was supported by the Bioplant Centre, University of Ciego de \u0026Aacute;vila (Cuba); Universidad de Concepci\u0026oacute;n (Chile); Universidad Estatal del Sur de Manab\u0026iacute; (Ecuador); and Agricultural Research Council (South Africa).\u0026nbsp;\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n \u003cli\u003eAcharya SN, Thomas JE, Basu SK (2006) Fenugreek: an \u0026ldquo;old world\u0026rdquo; crop for the \u0026ldquo;new world\u0026rdquo;. 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Biosci Biotech Biochem 66:65-72\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Cicer arietinum L., field performance, seed heteromorphism, seed size","lastPublishedDoi":"10.21203/rs.3.rs-2496069/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-2496069/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003e \u003cem\u003eCicer arietinum\u003c/em\u003e L. is a major food legume across the globe. However, the yield of legume crops appears to have reached a plateau in developing countries where yield is often impacted by poor crop establishment. Therefore, seed physiological characteristics (specific to the cultivars/landraces adapted to various regions of the world) and their impacts on plantlet establishment and performance should be investigated. This study determined the effect of seed size on germination, plant development and agronomic performance in Cuba. Biochemical parameters were also evaluated for up to 21 d of growth. The results showed that seeds of the largest mass (89\u0026thinsp;\u0026plusmn;\u0026thinsp;3 mg, group 3) showed more rapid germination, emergence and plant growth than the other tested mass categories. This trend was sustained until plant maturity where group 3 seeds also generated the highest yields. Differences were also noted in the antioxidant profiles in developing plants with the highest levels of SOD and PER found in plants generated from seeds with the smallest mass (55\u0026thinsp;\u0026plusmn;\u0026thinsp;3 mg per seed, group 1). The above findings raise the questions as to whether seeds of chickpea display heteromorphic behavior, however, further studies are required.\u003c/p\u003e","manuscriptTitle":"Chickpea seed mass influences agronomical performance: a case for seed heteromorphism?","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2023-01-23 23:26:16","doi":"10.21203/rs.3.rs-2496069/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"399ea973-4ba9-4fdd-8e54-c45987800881","owner":[],"postedDate":"January 23rd, 2023","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2023-06-08T20:29:23+00:00","versionOfRecord":[],"versionCreatedAt":"2023-01-23 23:26:16","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-2496069","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-2496069","identity":"rs-2496069","version":["v1"]},"buildId":"7rjqhiLT3MXkJMwkYKINL","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}