Shahzad Jamaati-e-Somarin, Roghayyeh Zabihi-e-Mahmoodabad and Asgar Yari

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American-Eurasian J. Agric. & Environ. Sci., 9 (): 6-, 00 ISSN 88-6769 IDOSI Publications, 00 Response of Agronomical, Physiological, Apparent Recovery Nitrogen Use Efficiency and Yield of Potato Tuber (Solanum tuberosum L.), to Nitrogen and Plant Density Shahzad Jamaati-e-Somarin, Roghayyeh Zabihi-e-Mahmoodabad and Asgar Yari Young Researchers Club, Islamic Azad University, Ardabil Branch, Ardabil, Iran Abstract: In order to investigation the response of agronomical, physiological, apparent recovery nitrogen use efficiency and yield of potato tuber (Agria cultivar), to nitrogen and plant density, a field experiment was carried out in Ardabil, Iran. The experiment was factorial in the base of randomized complete block design with three replications. Factors included nitrogen fertilizer level (0, 80, 60 and 00 kg ha as urea) and plant density (5.5, 7.5 and plant m ). Results showed that increasing nitrogen level up to 80 kg ha led to more tuber yield. The highest yield, number of stolon, harvest index, number of tuber and dry weight of tuber were obtained with application of jointly 80 and60 kg ha nitrogen. Increasing plant density resulted in higher tuber yield, number of stolon, dry weight of tuber and total dry matter yield and decreased harvest index. Increasing nitrogen fertilizer rate up to 60 kg ha increased mean tuber weight and total dry matter yield. With increasing the nitrogen levels and plant densities agronomical nitrogen use efficiency, physiological nitrogen efficiency and nitrogen use efficiency were decreased and apparent recovery nitrogen efficiency was increased. The most nitrogen uptake in plant was observed at the 00 kg ha net nitrogen. So, to obtain the highest yield with the most nitrogen use efficiency and avoid of environmental pollutions, use of 80 kg ha nitrogen fertilizer with density of plant m is recommended. Key words: Nitrogen efficiency Nitrogen uptake Plant density Potato and Yield components INTRODUCTION crop N needs to match crop N requirements and mineral N supply throughout the growing season. Georgakis et al. Potato (Solanum tuberosum L.) is grown and eaten in [5] concluded that by increasing plant density, the tuber majority of countries more than any other crops and in yield was increased. Karafyllidis et al. [6] reported that the global economy, it is the fourth most important crop plant density strongly affected yield, both by number after the three cereals including maize, rice and wheat []. and by weight and more tubers and yield per square meter Plant density in potato affects some of important were expected in higher plant densities. Alvin et al. [7] plant traits such as total yield, tuber size distribution and reported that with increasing plant density, yield of potato tuber quality. So it might be argued that increase in increased. Efficiency has defined as crop production per plant density leads to decrease in mean tuber weight and unit input, which means nutritional efficiency consists in an increase in number of tuber and yield per unit area []. the amount of dry matter production per unit nutrient Increase in plant density decreases mean tuber size elements usage or uptake [8]. Haase et al. [9] found that probably because of plant nutrient elements reduction, tuber N uptake was significantly influenced by amounts increase in interspecies competition and large number of of nitrogen fertilizer. The most nutrient element efficiency tubers produced by high numbers of stems [3]. is attained, while its concentration reaches critical level, Marguerite et al. [4] showed that the mean maximum because without excessive elements, greatest yield can be increase in total tuber yield, generated by nitrogen gain. Rate of this efficiency decreases at abundance and fertilizer against the zero-n treatment, was 34.3% and luxury (poisonous) ranges because with increasing ranged from 0.5% to 54.7% and in regard to potato, the element concentration, yield remains constant or improvement of N efficiency should be also achieved by decreases [8]. Marguerite et al. [4] showed that the mean splitting N fertilizer applications and by monitoring the maximum increase in the total tuber yield, generated by N Corresponding Author: Shahzad Jamaati-e-Somarin, Young Researchers Club, Islamic Azad University, Ardabil Branch, Ardabil, Iran, Tel: +9894594490, E-mail: jamaati_36@yahoo.com. 6

Am-Euras. J. Agric. & Environ. Sci., 9 (): 6-, 00 fertilization against to zero-n treatment, was 34.3% and Amount of N absorbed by plant calculated as shown ranged from 0.5% to 54.7% and in regard to potato, the by Hashemidezfooli et al. [8] (Element absorption = improvement of N efficiency should be also achieved by element concentration dry matter). N uptake by plant splitting N fertilizer applications. Karafyllidis et al. [6] was measured of sum of its rates in aerial parts and tubers. reported that plant density strongly affected yield, Also, tuber nitrogen content was calculated by Kjeldahl both by number and by weight and more tubers and Method. Agronomical, apparent and physiological yield per square meter were expected in higher plant nitrogen efficiencies were calculated from densities. Georgakis et al. [5] concluded that by Hashemidezfooli et al. [8]: increasing plant density, the tuber yield was increased. The aim of this study was to evaluate effects of the NAE = [(Y f Y f0)/n f] nitrogen levels and plant densities on the nitrogen PHNE = [(TDM f TDM f0)/ (NU f NU f0)] efficiency components (agronomical, physiological, ARNE% = [(NU f NU f0)/n f] 00 apparent recovery and nitrogen use efficiency), the NUE = (PHNE ARNE) = kg kg amount of nitrogen uptake by plant, yield and yield components in order to determination of the most suitable NAE: Agronomical N efficiency (kg ha ), Y f: Tuber density and nitrogen level to get favorite yield with the yield per plot with N application (kg ha ), Y f0: Tuber yield highest nitrogen use efficiency in Ardabil region, Iran. per plot without N application (kg ha ), N f: N rates (kg ha ), PHNE: Physiological nitrogen efficiency MATERIALS AND METHODS (kg ha ), TDM f: Total plant dry matter with N application (kg ha ), TDM f0: Total plant dry matter without N In order to investigation the response of agronomical, application (kg ha ), NU f: N uptake by plant with N physiological, apparent recovery nitrogen use efficiency application (kg ha ), NU f0: N uptake by plant without N and yield of potato tuber (Agria cultivar), to nitrogen and application (kg ha ), ARNE%: Apparent recovery plant density, a field experiment was carried out in nitrogen efficiency percent, NUE: Nitrogen use efficiency Ardabil, Iran in 008. The experiment was factorial in the (kg ha ). base of randomized complete block design with three In order to normalize data distribution, yield and replications. Factors included nitrogen fertilizer level efficiency data analysis of variance was done by SAS (0, 80, 60 and 00 kg ha as urea) and plant density Software, means of variances were compared through (5.5, 7.5 and plant m ). Nitrogen was given as urea Duncan's multiple range test and graphs were drawn by form at stages (namely, planting date and date of Excel software. By the way, ANOVA results were not earthing up). According to soil analysis results, total shown. nitrogen content was 0.56% and soil texture was sandy-loam. Rows were spaced 60 cm to each other and RESULTS plots contained 6 rows each 3 meters. In order to prevent nitrogen effects in adjacent plots (border effects),.5 m Tuber Yield and Dry Weight: The effect of plant borders were made. Tubers of 60-70 grams were sown density and nitrogen level was significant (P<0.0) on 3 May 008. Sowing depth was -3 cm. The last on tuber yield. Results showed that increasing harvest was assigned for the yield. To promote storage nitrogen rates up to favorable point increased tuber capability, ten days before harvest, aerial parts were yield per unit area. The highest value of tuber yield was removed [3]. Sampling was done from m plot area and obtained at 80 and 60 N kg h, while the lowest one then tubers were transferred to the laboratory. Before any belonged to control. With increasing nitrogen application, measurements, tubers were washed along with roots and the number of stolons including tuber, number of tubers stolons. Different plant tissues were dried separately for and consequently yield were increased. Also, increase in 48 hours at 75 C and weighed. Amount of harvest index density led to significant increase in tuber yield so that, was calculated as follows: the most and the least tuber yield was obtained at plant m and at 5.5 and 7.5 plant m, respectively Harvest index = (Economical yield/biological yield) 00 (Table ). The effect of plant density and nitrogen level on Also, total dry matter yield was calculated from sum tuber dry weight was significant (P<0.0). Based on the of dry matter of aerial and sub-ground parts. results, tuber dry weight was increased up to favorable 7

Am-Euras. J. Agric. & Environ. Sci., 9 (): 6-, 00 Table : Effects of plant density and nitrogen level on measured traits Number of Number of Mean tuber Total plant dry Dry weight of Tuber Harvest Treatments stolon m tuber m weight (g) matter (kg m ) tuber (g m ) yield (g m ) index (%) Nitrogen fertilizer values (kg ha ) 0.0 7.5c 63.86b 3.9b 0.9c 498.79b 04.6b 68.79b 80.0 94.56a 93.35a 30.ab.7b 669.95a 994.a 70.83a 60.0 96.83a 00.9a 33.67a.9a 78.8a 374.6a 7.5a 00.0 86.98b 80.3ab 4.85b.b 498.3b 457.0b 67.49b Plant densities (plant m ) 5.5 66.94c 77.b 30.55a 0.9c 55.35b 346.3b 7.4a 7.5 83.8b 8.6ab 7.36ab.05b 580.3b 473.8b 69.07b.0.4a 95.0a 6.ab.4a 74.45a 36.6a 68.4b Numbers with the same letters in each column, have no significant differences to each other amounts of applied nitrogen per unit area. Also, it was increased. Of course, it was expected that increasing observed that increasing plant density significantly plant density and decrease in the inter plant distances increased tuber dry weight. The highest and lowest tuber would lead to decrease in the number of stolons but, dry weight were gained at plant m and jointly at because of increase in applied nitrogen amounts, this 5.5 and 7.5 plant m, respectively (Table ). appears to be rational. As it is obvious in the results section, this trait follows the same trend in two other Mean Tuber Weight and Total Plant Dry Matter: The densities. With regard to the fact that produced tubers effect of nitrogen and plant density (P<0.0) was per plant are the functions of the number of stolons so, significant on mean tuber weight. With increasing it is expectable that in higher densities, the number of nitrogen up to definite point, tuber weight was increased produced tubers be more, as well. This is, with partial so that, the highest value was obtained with application difference, similar to the increment of the number of of 60 kg N ha. Meanwhile, control or 00 kg N ha stolons. In fact, these traits were increased with rates had significant effect on mean tuber weight. increasing applied nitrogen value and density, separately. Also, the lowest mean tuber weight was achieved at Furthermore, the interaction effect of nitrogen fertilizer 7.5 and plant m and the highest one was achieved at plant density, had significant impact on them so that, 5.5 plant m (Table ). density of plant m along with the application of 80 kg Total plant dry matter was affected by plant density N ha and also, 60 kg N ha with the mentioned and nitrogen, significantly (P<0.0). Results presented density, had the highest number of stolons of.55 and in Table showed that application of nitrogen up to 5.03, respectively (Table ). According to the results, 60 kg ha resulted in an increase in total plant dry in both treatments, the highest density along with the matter,while excess rates decreased in this trait. Plant nitrogen levels of the 80 or 60 kg N ha increased this density had very positive effect. As it was seen in trait as they increased the number of stolons. In other majority of traits, increasing nitrogen rate up to a distinct words, high nitrogen levels could overcome the inter rate increased total plant dry matter yield and afterwards, plant competitions caused by the high densities limiting decreased. the environmental resources for plants. Range of these changes in the number of tuber per unit area was higher Number of Stolon and Tuber: Using 80 and 60 kg N ha, than the number of stolons so that, interaction effect of the number of stolon m significantly was more than all plant densities nitrogen fertilizers could show the control or 00 N kg ha. The highest value of number of significant differences. However, values of mentioned stolon was obtained at the rate of 60 kg N ha of about treatments which recorded the highest number of stolons, 96.83. This trend is obvious for the number of tuber m made the highest number of tubers per unit area, as well. so that, the rate of 60 kg N ha produced the number of 00.9 tubers m (Table ). In contrast, the highest plant Harvest Index: Plant density and nitrogen level had density had the highest number of stolon and other two significant effect (P<0.0) on harvest index. With levels were placed afterwards. The impact of the first and increasing plant density and nitrogen level, harvest the third densities showed that the third one produced index was decreased so that, density of 5.5 plant m approximately two times more stolons than the first recorded the highest harvest index value,while densities (.4 stolons m ). Same to this, plant density has been of 7.5 and plant m jointly recorded the lowest. 8

Am-Euras. J. Agric. & Environ. Sci., 9 (): 6-, 00 Table Interaction effects of plant density and nitrogen level on measured traits Number of Number of Mean tuber Total plant dry Dry weight of Tuber Harvest Interaction effects stolon m tuber m weight (g) matter (kg m ) tuber (g m ) yield (g m ) index (%) Control 5.5 plant m 57. h 58.8 b 34. ab 0.67 f 37.48 f 650.00e 7.04 abc Control 7.5 plant m 67.87 fg 75. ab 7.95 b 0.84 e 445.7 f 65.7e 67.4 bc Control plant m 9.58 c 58.8 b 3.4 b. c 679.6 abcd 77.abcd 65.64 c 80 kg ha 5.5 plant m 7.77 ef 86.7 ab 38.00 ab 0.97 de 649.5 bcd 39.6abc 74.8 ab 80 kg ha 7.5 plant m 90.37 c 80. ab 3.7 b 0.99 d 58.47 de 478.7bcde 70.87 abc 80 kg ha plant m.55 a.88 a 36.53 ab.4 b 778.4 ab 33.ab 67.49 bc 60 kg ha 5.5 plant m 7.6 e 86.6 ab 47.43 a.0 d 6.43 cd 3004.8abc 76.6 a 60 kg ha 7.5 plant m 9.87 c 03.55ab 8.93 b.3 c 744.3 abc 906.9abcd 69.67 abc 60 kg ha plant m 5.03 a 3.3 a 3.9 b.6 a 88.78 a 36.00a 68.63 bc 00 kg ha 5.5 plant m 66.7 g 60.4 b 34.67 ab 0.97 de 459. ef 047.de 70.4 abc 00 kg ha 7.5 plant m 84. d 84.56 ab 8.05 b. cd 549. de 349.5cde 69.7 abc 00 kg ha plant m 0.55 b 95.73 ab 3.03 b.39 b 693.8 abcd 974.4abc 67.55 bc Numbers with the same letters in each column, have no significant differences to each other Table 3: Effects of plant density and nitrogen levels on measured traits Nitrogen uptake Agronomical nitrogen Apparent recovery Physiological nitrogen Nitrogen use Treatments by plant (g m ) use efficiency (kg kg ) nitrogen efficiency (%) efficiency (kg kg ) efficiency (kg kg ) Nitrogen fertilizer doses (kg ha ) 0.0 753.8c - - - - 80.0 308.b.9a 69.8a 0.56a 6.8a 60.0 507.b 7.87b 47.08b 0.54a 3.89ab 00.0 759.0a.6c 50.5ab 0.8b 3.3b Plant density (plant m ) 5.5 85.c 98.95a 38.5b 0.68a 5.8a 7.5 34.4b 7.37ab 53.94ab 0.39b 8.96a.0 90.45a 43.37b 74.7a 0.3b 9.78a Numbers with the same letters in each column, have no significant differences to each other Also, application of 80 or 60 kg N ha resulted in the Apparent Recovery Nitrogen Efficiency: It was found highest value of harvest index, while the lowest was that nitrogen levels (P<0.05) and plant density (P<0.0) recorded with the control treatment or 00 kg N ha had the significant effects on apparent recovery nitrogen (Table ). Application of 80 or 60 kg N/ha_resulted in an efficiency. As shown in Table 3, at level of 80 N kg ha, increase in the harvest index, also it was increased by ARNE was 69.8 %. Also, application of 00 N kg ha using plants m. According to Table, application of recorded 50.5 % which were significant. In regard to 60 kg N ha 5.5 plants m led to the highest value in plant density, the most efficiency concerned to density the harvest index. of plant m about 74.7 % furthermore in density of 7.5 plant m this value was 53.94 % which both were Agronomical Nitrogen Use Efficiency: Results revealed significant. significant effects of the plant density (P<0.05) and nitrogen levels (P<0.0) on agronomical nitrogen use Physiological Nitrogen Efficiency: Effect of plant density efficiency. Increasing nitrogen rates and plant density, and nitrogen levels (P<0.05) on physiological nitrogen NAE was reduced, however, the highest (.9 kg ha ) efficiency was significant. In this case, with increasing and lowest (.6 kg ha ) efficiency values were recorded plant density and nitrogen levels, physiological nitrogen at 80 and 00 N kg ha, respectively. Also, data in efficiency was decreased. In the other words, the plant Table 3 indicated that NAE recorded in the, the highest density of 5.5 plant m recorded the highest PHNE, (98.95 kg ha ) and lowest (43.37 kg ha ) under 5.5 and while at 7.5 and plant m recorded the lowest plant m. efficiency. Application of 80 N kg ha had the highest 9

Am-Euras. J. Agric. & Environ. Sci., 9 (): 6-, 00 value of PHNE, whereas, there was no significant tuber dry weight per unit area whereas, increase in difference between 80 and 60 N kg ha. At 00 kg density, increased the competition between and within Nha, the lowest value of PHNE was obtained (Table 3). the plants and hence, leads to decreased availability of nutrients for each plant and consequently, results in Nitrogen Use Efficiency: Application of nitrogen rates decline of mean tuber weight [6]. Applied nitrogen less had significant effect on nitrogen use efficiency (P<0.05), affects number of tuber but more affects tuber size and but there was no significant effect due to plant density. increases it so, directly increases mean tuber weight but The highest NUE (6.8 kg ha ) value was obtained at in condition of excess rates of nitrogen, mean tuber the 80 N kg ha, while the lowest (3.3 kg ha ) weight is decreased []. Alvin et al. [7] showed that with (Table 3)at the 00 kg ha. increasing plant density, plant dry matter was decreased in each plant but increased per unit area. It can be said Nitrogen Uptake by Plant: Effects of plant densities that all factors affecting tuber yield, affect total plant and nitrogen levels significantly (P<0.0) affected N dry matter, as well [8]. Based on the results, with uptake by plant. The highest (759.0 g m ) and lowest increasing plant density, harvest index was decreased (753.8 g m ) N uptake values were observed at 00 and vice versa. Since, both total plant dry matter and dry N kg ha and control treatments, respectively. For plant weight of tuber were increased but this increase for density, the highest (90.45 g m ) and lowest (85. total plant dry matter was more than tuber dry weight, this g m ) values were obtained at plant m and 5.5 plant trait was decreased. Also, Cox and Cherny [] found m, respectively (Table 3). that with increasing density, harvest index is decreased. Arnout [3] revealed that high rates of applied nitrogen DISCUSSION can increase tuber dry weight. This finding approves our results on tuber dry weight and hence, we see that Increase nitrogen application amounts up to a harvest index is increased with increasing nitrogen rates. definite point, increased stolons including tuber but On the contrary, Osaki et al. [] stated that effect of beyond that, reversely decreased them. In other studies, nitrogen on biological yield is higher than economical it has been shown that number of stolon in suitable one. In addition, it is obvious from the results that nitrogen levels, was significantly more than other levels. increase in dry weight of tuber at the highest rate of Also, it has been approved that with increasing nitrogen nitrogen, is less than total plant dry matter. So, it can be amounts, number of tuber was increased, It is clear that the reason of this fact that with increasing nitrogen up to with increasing plant number, numbers of stems grown 00 kg ha, harvest index was decreased. from the planted tuber and consequently, number of With increasing nitrogen application from 80 kg ha, produced tubers per stem, are increased. Thus, agronomical nitrogen use efficiency and nitrogen use increase plant density increase tubers production [3]. efficiency were decreased. Since in this level of Over-application of nitrogen may result in decrease in fertilization, the most yields were gained and thereafter, yield. This may attributable to the fact that in such this increase was educational, so, it could be the reason conditions, vegetative growth of the aerial parts can be for decrease of these efficiencies. Hartemink et al. [4] increased and hence, prevented transferring of reported that nitrogen use efficiency increases with photosynthetically matters into the storage parts increased N application for sweet potato. For apparent (Tubers). Marguerite et al. [4] and Alam et al. [0] recovery nitrogen efficiency, it could be said that since revealed that tuber yield per unit area was increased highest tuber yield have been gained jointly at 80 and with increasing nitrogen fertilizer up to suitable level. 60 kg ha nitrogen, so, it might concluded that values According to the Alvin et al. [7], in high densities, of N uptake at this level (80 kg ha ) had been much more number of tuber and yield of potato is increased. Same to than next level, per unit fertilizer. There fore, the most the tuber yield, tuber dry weight is affected by rates of efficiency have occurred at this rate and since, utmost nitrogen application and is increased with increasing N uptake have been made at 00 kg ha, so, this nitrogen [, 0]. This increase is probably due to increase efficiency have been improved, as well. Darwish et al. [5] in number of stolon, number of tuber and tuber yield. found that decreasing N application of 5 kg ha, With regard to the fact that with increasing plant density, (against 50, 375 and 500 kg N ha ) significantly led to the number of produced tuber is increased, it can be said higher N recovery. In this study, we found that higher that increase in number of tuber may result in increase in nitrogen levels resulted in lowest physiological nitrogen 0

Am-Euras. J. Agric. & Environ. Sci., 9 (): 6-, 00 efficiency. In general, it could be said that with increasing 7. Alvin, J.B., P.D. Mitchell, M.E. Copas and nitrogen fertilizer and plant density, agronomical, M.J. Drilias, 007. Evaluation of the Effect of Density physiological and nitrogen use efficiencies were on Potato Yield and Tuber Size Distribution. Crop decreased and apparent recovery nitrogen efficiency Sci., 47: 46-47. was increased. As shown in this study, nitrogen 8. Hashemidezfooli, A., A. Koochaki and M. Banayan application over the 60 kg ha, total plant dry matter, Avval, 998. Crop plant improvement (translation). number of tuber and mean tuber weight were decreased. Jehad Daneshgahi Mashad press. Mashad. Iran. Likely, this is attributable to intra competition of different Edition Number: 3. ISBN 964-603-05-3. plant parts to achieve environmental resources such 9. Haase, T., C. Schuler and J. Heb, 007. The effect of as water, light and nutrient elements and effect of different N and K sources on tuber nutrient excessive nitrogen to stimulate growth of plant aerial uptake, total and graded yield of potatoes parts and directly, reducing photosynthetic matters (Solanum tuberosum L.) for processing. Euro. J. transfer to the tubers. Agronomy, 6: 87-97. 0. Alam, M.N., M.S. Jahan, M.K. Ali, M.A. Ashraf REFERENCES and M.K. Islam, 007. Effect of vermicompost and chemical fertilizers on growth, yield and yield. Stephen, D.J., 999. Multiple signaling pathways components of potato in barind soils of Bangladesh. control tuber induction in potato. Plant Physiol., J. Appl. Sci. Res., 3(): 879-888. 9: -8.. Koochaki, A. and G.H. Sarmadnia, 00. Physiology. Osaki, M., H. Ueda, T. Shinano, H. Matsui and of crop plants (translation). Jehad Daneshgahi T. Tadano, 995. Accumulation of carbon and Mashhad press, Mashhad. Iran. nitrogen compounds in sweet potato plants grown. Cox, W.J. and D.J.R. Cherny, 00. Row spacing, under deficiency of N, P, or K nutrients. Soil Science plant density and nitrogen effects on corn silage. and Plant Nutrition, 4(3): 557-566. Agron. J., 93: 597-60. 3. Khajehpour, M., 006. Production of industrial 3. Arnout, V.D., 00. Yield and Growth Components of plants. Jehad-e-Daneshgahi Isfahan press, Isfahan. Potato and Wheat under Organic Nitrogen Iran. Management. Agron. J., 93: 370-385. 4. Marguerite, O., G. Jean-Pierre and L. Jean-Francois, 4. Hartemink, A.E., M. Johnston, J.N. O Sullivan and 006. Threshold Value for Chlorophyll Meter as S. Poloma, 000. Nitrogen use efficiency of taro and Decision Tool for Nitrogen Management of Potato. sweet potato in the humid lowlands of Papua New Agron. J., 98: 496-506. Guinea. Agric. Ecos. Environ., 79: 7-80. 5. Georgakis, D.N., D.I. Karafyllidis, N.I. Stavropoulos, 5. Darwish, T.M., T.W. Atallah, S. Hajhasan and E.X. Nianiou and I.A. Vezyroglou, 997. Effect of A. Haidar, 006. Nitrogen and water use efficiency of planting density and size of potato seed-minitubers fertigated processing potato. Agric. Water on the size of the produced potato seed tubers. Acta Management, 85: 95-04. Hort. (ISHS), 46: 935-94. 6. Karafyllidis, D.I., D.N. Georgakis, N.I. Stavropoulos, E.X. Nianiou and I.A. Vezyroglou, 997. Effect of planting density and size of potato seed-minitubers on their yielding capacity. Acta Hort. (ISHS), 46: 943-950.