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SAN J O S E NEIKER - Instituto Vasco de Investigaci6n y Desarrollo Agrario, Apartado 46, 01080 Vitoria- Gasteiz, Spain Accepted for publication: 26 October 2000 Additional keywords: Sola

Potato Research 44 (2001) - Comparison of hydroponic and aeroponic cultivation systems for the production of potato minitubers E R I T T E R , B A N G U L O , P R I G A , C H E R R A N , J R E L L O S O and M SAN JOSE NEIKER - Instituto Vasco de Investigaci6n y Desarrollo Agrario, Apartado 46, 01080 VitoriaGasteiz, Spain Accepted for publication: 26 October 2000 Additional keywords: Solalmm mberosum L., in vitro culture, micropopagation, seed potato, repeated harvesting Summary Two different cultivation systems, aeroponics and hydroponics in greenhouse beds, were compared for the production of potato minitubers Plants in the aeroponic system showed increased vegetative growth, delayed tuber formation and an extended vegetative cycle of about seven months after transplanting Therefore in 1999, two production cycles were obtained with the hydroponic system, but only one with the aeroponic system However, compared with total production in hydroponics, the tuber yield per plant in the aeroponic system was almost 70% higher and tuber number more than 2.5 fold higher Average tuber weight was reduced by 33% in the aeroponic system Advantages and possible problems with the aeroponic system for minituber production are discussed Introduction Seed potato production is mostly based on in vitro plantlets or microtubers, and on the subsequent production of minitubers as first ex vitro generation (Ranalli, 1997) Minitubers can be produced after acclimatisation from plantlets which are planted at high densities in the greenhouse in beds (Wiersema et al., 1987) or in containers (Jones, 1988) using different substrate mixtures, or even in hydroponic culture (Muro et al., 1997) L o m m e n (1995) presented alternative production techniques for minitubers using very high plant densities and non-destructive, repeated harvesting of minitubers by lifting plants carefully from the soil mixture and replanting them after harvest These techniques allowed minitubers of ideal size to be produced, the number of tubers could be increased considerably, while total yield was reduced In modern horticulture, different soil-less production techniques such as Nutrient Film Techniques (NFT, Cooper, 1979) and aeroponics (Peterson et al., 1968) have been developed The technique of aeroponic culture is an optional device of the soilless culture methods in growth-controlled environments such as greenhouses This method consists of enclosing the root system in a dark chamber and supplying a solution of water and mineral nutrients with a mist device This technique has been applied successfully for the production of different horticultural species including Potato Research 44 (2001) 127 E RITTER, B ANGULO, P RIGA, C HERRAN, J RELLOSO AND M SAN JOSE lettuce (Cho et al., 1996; Gysi & v o n Allmen, 1997; He & Lee, 1998), tomato (Biddinger et al., 1998), cucumber (Park et al., 1997) and ornamental plants such as chrysanthemum (Molitor et al., 1999) or poinsettia (Scoggins & Mills, 1998) Despite increasing interest in soil-less culture methods in commercial horticultural production, little information is available for potatoes Earlier work has shown good results with NFT for potato tuber production (Wheeler et al., 1990; Wan et al., 1994) However, tuber initiation was poorer in nutrient solution without solid media than in porous media (e.g perlite or vermiculite) The tuberization inhibition of stolons immersed in a solution could be the consequence of the lack of mechanical resistance (Vreugdenhil & Struik, 1989) Recently, an aeroponic system for seed potato production was established successfully in Korea (Kang, 1996a,b; Kim et al., 1997, 1999) under tropical and subtropical conditions In view of the favourable results obtained by Lommen (1995) with respect to repeated harvesting and considering the characteristics of aeroponic culture, the combination of both techniques seems to be particularly useful for minituber production in potato In this paper we describe a method for the production of minitubers using aeroponics, and present first comparative results with respect to traditional production schemes in greenhouse beds using hydroponics Materials and methods Plandet preparation Plantlets of the local cultivar Nagore were produced in vitro according to Espinoza et al (1984) from replicated nodes on standard MurashigeSkoog medium (Murashige & Skoog, 1962) After the last multiplication step, nodes of the plantlets were cultivated under non-sterile conditions on filter paper soaked with the same medium but without agar and sugar in order to induce root formation Rooted nodes were transferred after to days to seedling trays containing a mixture of 60% white and 40% black turf with added fertilizers as substrate Plantlets were acclimatised for one week under the humid conditions of a plastic cover and regenerated for three weeks to small potato plants of to cm height This plant material constituted the starting material for the assays Two cultivation systems (hydroponics in greenhouse beds and aeroponics) were assayed and compared Hydroponic system Two production cycles were obtained during 1999 near Vitoria in Northern Spain One was from March to June 29 and the second from July 15 to November 10 Temperature was maintained between 18 and 22 ~ by a cooling/ heating system The plants grew under natural light conditions without additional light supply The small potato plants were planted without washing the roots into a greenhouse bed containing perlite (Agroperl) Substrate depth was approximately 30 cm The plant density was 10x10 cm Nutrient solution (pH 6.5, EC=mS cm -t, NO3-, SO42-, H2PO 4", CI-, K +, Ca 2+, Mg 2+, NH4 +, Na + and micronutrients) were supplied when necessary using drop irrigation Fungicide and insecticide treatments were applied when appropriate After 16 weeks, the plants had nearly completed their vegetative cycle, haulms were destroyed by a total herbicide (Paraquat) and tubers 128 Potato Research 44 (2001) HYDROPONICS A N D AEROPONICS FOR M I N I T U B E R P R O D U C T I O N were harvested one week later Four representative areas of m (100 plants) were selected within the greenhouse bed and harvested separately T u b e r numbers, and tuber yield were recorded for each replicate Shortly after harvest a new set of small potato plants were planted in the same greenhouse bed The plants were cultivated under identical conditions and harvested as described before Aeroponics Aeroponic culture was performed in the same greenhouse in six closed containers (60 cm height, 45 cm width, 85 cm depth), which had a removable front panel for control and harvesting Small acclimatised plantlets of cv Nagore were planted on May by fixing them on top of the boxes with the help of a gum grid Perlite was used to seal the grid In this way the foliage of the small plants grew in the light while roots, stolons and tubers developed in total darkness in the containers The plant density was 10xl0 cm with a total of 36 plants per container Water and nutritive elements were supplied with the same nutrition solution as in the hydroponic culture For this purpose four fog nozzles (4 l/h) located at the bottom of the containers sprayed nutrient solution every 10 for sec into the inner part of the box in order to keep the roots wet Residual nutrition solution was recycled Fig shows the aeroponic device with plants Since the plants reached a considerable height during their development it was necessary to stake them using ribb.ons The same phytosanitary treatments were applied as in the hydroponic system Tubers were harvested repeatedly and in total eight times (Table 1) The harvest criteria were to remove all tubers which were larger than 15 to 20 m m in length At the last harvest all tubers larger than m m were considered The first harvest was on July when the first tubers of sufficient size were available The following harvest intervals were around one month and were later shortened due to increased production and tuber development (Table 1) The n u m b e r of tubers and total yield were recorded each time in each box After each harvest the plants were re-fixed in the gum grid approximately to cm lower, in order to allow new stolon and subsequent tuber formation Data processing and statistical analyses Analyses of variance were performed using SAS, Proc G L M (SAS Institute, 1989) Results The potato plants in the greenhouse bed reached a final height of 90 to 110 cm, while those of the aeroponic system reached a total height of 150 to 180 cm Increased vegetative growth of plants in the aeroponic system was also observed for the root system and for stolon length (data not shown) Also as expected, plants in the bed completed their biological cycle approximately four months after transplanting, while the aeroponic plants continued to grow and form new shoots from lateral buds Occasionally secondary growth was observed which sometimes led to the formation of new stolons with additional tubers Plants in aeroponic culture finished their cycle approximately seven months after transplanting Potato Research 44 (2001) 129 E RITTER, B ANGULO, P RIGA, C HERR,s J RELLOSO AND M SAN JOSE Fig Aeroponic system: A) empty aeroponic containers with fog nozzles and connecting tubes, B) acclimatised plantlets growing in aeroponic system, C) fully developed potato plants, D) minitubers obtained Results of the measured yield parameters are shown in Table A total of 4.9 tubers per plant was obtained in both cycles with the hydroponic plants Total yield was approximately 65.3 g per plant and the corresponding average tuber weight was 13.3 g This weight corresponds to a grade of 40-45 mm in cv Nagore which has long tubers with a form index of 1.8 to 2.2 (length/width) In the first production cycle a higher yield with slightly reduced tuber numbers but considerably larger tubers were obtained Although plants of the hydroponic system were planted two months earlier, tuberization and tuber development were delayed in the aeroponic system as can be seen in Table 1, with respect to the first three harvesting dates The development of aerial parts of the plants was initially favoured Production between consecutive harvests increased during development and was reduced as the plants senesced After a slow initial phase of tuber formation, fewer but larger tubers were produced A large peak in total production due to an increasing number of smaller sized tubers was observed after five months of development (6 th harvest date) A total tuber yield 130 Potato Research 44 (2001) HYDROPONICS A N D AEROPONICS FOR MINITUBER P R O D U C T I O N Table Yield parameters per plant obtained with different minituber production systems Hydroponic system Cycle I; March to June 29 Cycle II;July 15 to Nov 10 Total TN/P Y/P ATW 2.3 2.6 4.9 36.2 29.l 65.3 15.8 11.2 13,3 TN/P Y/P ATW 0.2 0.4 1.1 1.3 1.5 3.7 1.4 2.8 12.4 0.7 2.5 17.8 19.3 19.2 37.6 6.0 6.7 109.9 4.6 6.5 15.5 14.7 13.2 10.2 4.2 2.4 8.9 Aeroponic system (Cycle from May to December 3) Harvesting date July August 3 September 4 October October22 November5 November 19 December Total Pr>F 0.0002 0.002 0.0001 TN/P= average number of tubers per plant, Y/P=yield per plant (g), ATW=average tuber weight (g), Pr>F=probability for F-value from Analyses of variance of 109.9 g was obtained per plant which was composed on average of 12.4 tubers with an average tuber weight of 8.9 g (grade 30-35 mm) C o m p a r e d with each single harvest in the hydroponic system, its number of tubers and total yield was three to four times higher In order to compare the two systems, we considered the whole year as a productive period Therefore, tuber numbers and yields were summed over both production cycles in the hydroponic system and compared with total tuber number and yield of the aeroponic system by means of analyses of variance The influence of the cultivation system was highly significant for all analysed yield parameters (Table 1) Compared with the total production in hydroponics, the tuber yield per plant in the aeroponic system was almost 70% higher and tuber number was over 2.5 fold higher However, average tuber weight was 33% less in the aeroponic system Previous work showed that the post-harvest behaviour of tubers seem to be similar in both production systems (unpublished data) Tubers grown in aeroponics showed an increased n u m b e r of opened lenticels, due to high humidity conditions in the system However, when stored under appropriate conditions, no differences were found with respect to storage (% rotten tubers; velocity and intensity of sprouting) When planted in the greenhouse or field, no apparent differences in development and tuber yield was observed compared with hydroponic seed tubers (results not shown) Potato Research 44 (2001) 131 E RITTER, B ANGULO, P RIGA, C HERRAN, J RELLOSO AND M SAN JOSE Discussion Soil-less cultivation systems frequently used in horticulture present several advantages compared with soil production Hydroponics lead to higher production (Resh, 1978; FAO, 1990) Gysi & v o n Allmen (1997) found higher yield for tomatoes grown in NFT and aeroponics than grown in soil In previous experiments the hydroponic system proved more convenient than the traditional propagation system using peat/ sand mixtures and mineral fertilisers (unpublished data) These findings confirmed those of Muro et al (1997), who obtained increased yield and tuber numbers with hydroponics Moreover, hydroponics gave higher quality and were from one week to one month earlier (Morard, 1995), avoided several soil pathogens or unsuitable physical and chemical soil properties, and optimised crop production by the enhancement of water and mineral availibility in the root environment However, aeroponic techniques optimise root aeration which is certainly the major factor leading to a yield increase compared with classical hydroponic systems (Softer & Burger, 1988) Cho et al (1996) observed in cherry tomatoes a growth and yield increase in aeroponic compared with classical hydroponics In the present study, increased vegetative growth of potato plants and delayed tuberization was observed in aeroponic systems compared with hydroponics, which might be due to the unlimited nitrogen supply in our aeroponic system Kang et al, (1996a) presented similar observations when the N supply was increased in the nutrient solution of aeroponically-grown potato plants Furthermore, Vreugdenhil & Struik (1989) found that tuber initiation needs the cessation of stolon growth, which is linked to ethylene synthesis A delay of tuberization has been observed when the stolon environment did not provide mechanical stress (Lugt et al., 1964) This is so in aeroponic cultures, where roots grow in air without mechanical resistance Moreover, the increased stolon length obtained in the present study supports this hypothesis Previous observations showed that absolute darkness is necessary for tuber formation, otherwise with a minimum of light stolon tips developed small bleached leaves and no tuber formation occurred However, hand pulling down the plants after each harvest was necessary to increase the formation of new stolons with tubers The same behaviour was also observed in the repeated harvest assays of Lommen (1995) who replanted the plants deeper after a previous harvest The number of tubers and yield in our hydroponic system was favourable and within the expected range Compared with the results obtained by Muro et al (1997), production in terms of tuber number, yield and average tuber weight was higher, possibly because a higher plant density was used in our study Nevertheless, with the aeroponic system for minituber production with repeated harvesting we were able to increase yield and especially tuber number although there were two productive cycles per year with hydroponics (Table 1) This can be attributed to an improved availability of nutrients, mainly calcium, that is required in stolon tips for tuber initiation (Balamani et al., 1986), and due to the removal of the dominant large tubers, which allow initiation of new tubers as well as the development of existing tubers (Lommen, 1995) Moreover, tuber number is positively correlated with leaf area and number of 132 Potato Research 44 (2001) H Y D R O P O N I C S A N D A E R O P O N I C S FOR MINITUBER P R O D U C T I O N young leaves (Kahn et al., 1983), and this is so in aeroponic culture, where potato plants showed greater vegetative growth Using repeated harvesting systems Lommen (1995) obtained up to 3500 tubers of small size (>5 mm) using high plant densities of up to 800 plants/m However, our local agroclimatical conditions require larger minitubers to obtain an acceptable yield when planted in the heavy soils of the local fields This is particularly important under dry weather conditions if irrigation is not available Furthermore, compared with Lommen (1995), total tuber yield could be increased in our aeroponic system, since root damage due to lifting and replanting can be avoided Similar post-harvest behaviour of tubers from both production systems is consistent with the results of Kim et al (1999), who also did not find differences in growth and tuberization behaviour of seed tubers obtained from aeroponics compared with normal seed tubers The harvest technique in aeroponics is convenient and clean, and repeated harvesting offers the possibility of obtaining tubers of a desired size However, many aspects of this technique have to be investigated in order to optimise the system These include, for example, studies on appropriate nutrient solutions, plant densities and number of harvests and harvest intervals, as well as all possible interactions between production factors With respect to harvesting intervals Kim et al (1997) found that highest yields were obtained with shorter intervals of 10 days Furthermore, Kim et al (1999) improved production by increasing light intensity and by CO enrichment Other aspects which need to be considered include possibilities of automatization Labour and cost intensive processes in our system represented, for example, the staking of plants, manual harvesting and pulling down of the plants after each harvest Furthermore, in order to introduce the aeroponic system for large-scale minituber production an economic evaluation is necessary because methods of soil-less culture, and particularly aeroponics, present some disadvantages for industrial production: lack of water and chemical buffering capacities that must be compensated by security systems (alarms, pumps), high infrastructure costs, high technology and a specialised organisation of growers The results presented here lead to the conclusion that aeroponics can be an appropriate system for producing potato minitubers under temperate climatical conditions as it is in tropical areas (Kang et al., 1996a) References Balamani, V., K Veluthambi & B.W Poovaiah, 1986 Effect of calcium on tuberization in potato ( Solanum tuberosum L.) Plant Physiology 80: 856-858 Biddinger, E.J., C.M Liu, R.J Joly & K.G Raghothama, 1998 Physiological and molecular responses of aeroponically grown tomato plants to phosphorus deficiency Journal of the American Society for Horticult, ral Science 123: 330-333 Cho, Y.D., S.G Kang, Y.D Kim, G.H Shin & K.T Kim, 1996 Effects of culture systems on growth and yield of cherry tomatoes in hydroponics RDA Journal of Agricultural Science 38: 563-567 Cooper, A., 1979 The ABC of NFT Grower Books, London, 181 pp FAO, 1990 Soilless culture for horticultural crop produccion Plant Production and Protection paper n ~ 101, Rome, 188 pp Potato Research 44 (2001) 133 E RITTER, B ANGULO, P RIGA, C HERR/~N, J RELLOSO AND M SAN JOSE Espinoza, N.O., R Estrada, P Tovar, I.E Bryan & J Dodds, 1984 Tissue culture micropropagation, conservation and export of potato germplasm Specialized Technology Document CIP, Lima Gysi, C & F von Allmen, 1997 Balance of water and nutrients in tomatoes grown on soilless systems Agralforschung 4:1 (supplement) He, J & S.K Lee, 1998 Growth and photosynthetic responses of three aerponically grown lettuce cultivars (Lactuca sativa L.) to different rootzone temperatures and growth irradiances under tropical aerial conditions Journal of Horticultural Science and Biotechnology 73: 173-180 Jones, E.D., 1988 A current assesment of in vitro culture and other rapid multiplication methods in North America and Europe American Potato Journal 65: 209-220 Kahn, B.A., E.E Ewing & A.H Senesac, 1983 Effects of leaf age, leaf area, and other factors on tuberization of cuttings from induced potato (Solanum tuberosum L.) shoots Canadian Journal of Botany 61:3193-3201 Kang, J.G., S.Y Yang & S.Y Kim, 1996a Effects of nitrogen levels on the plant growth, tuberization and quality of potatoes grown in aeroponics Journal of the Korean Society for Horticultural Science 37:761-766 Kang, J.G., S.Y Kim, H.J Kim, Y.H Om & J.K Kim, 1996b Growth and tuberization of potato (Solarium tuberosum L.) cultivars in aeroponic, deep flow technique and nutrient film technique culture systems Journal of the Korean Society for Horticultural Science 37: 24-27 Kim, K.T., S.B Kim, S.B Ko & Y.B Park, 1997 Effects of minituber picking intervals on the yield and tuber weight of potato grown in aeroponics RDA Journal of Horticulture Science 39: 65-69 Kim, H.S., E.M Lee, M.A Lee, I.S Woo, C.S Moon, Y.B Lee & S.Y Kim, 1999 Production of high quality potato plantlets by autotrophic culture for aeroponic systems Journal of the Korean Society for Horticultural Science 123: 330-333 Lommen, W.J.M., 1995 Basic studies on the production and performance of potato minitubers Doctoral thesis, Wageningen Agricultural University, Wageningen, The Netherlands, 181 pp Lugt, C., K.B.A Bodlaender & G Goodijk, 1964 Observation on the induction of secondgrowth in potato tubers European Potato Journal 4: 219-227 Molitor, H.D., M Fischer & A.P Popadopoulos, 1999 Effect of several parameters on the growth of 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453 Resh, H., 1978 Hydroponic food production Woodbridge Press Publishing Company, Santa Barbara, California, USA, 287 pp SAS Institute Inc., 1989 SAS/STAT Users Guide, Version 6, Fourth edition, Vol SAS Institute Inc., Cary, N.C., 846 pp Scoggins, H.L & H.A Mills, 1998 Poinsettia growth, tissue nutrient concentration, and nutrient up take as influenced by nitrogen form and stage of growth Journal of Plant Nutrition 21: 191-198 Soffer, H & D.W Burger, 1988 Effects of dissolved oxygen concentration in aero-hydroponics on the formation and growth of adventitious roots Journal of the American Society for Horticuhural Science 113: 218-221 134 Potato Research 44 (2001) HYDROPONICS AND AEROPONICS FOR MINITUBER PRODUCTION Vreugdenhil, D & P.C Struik, 1989 An integrated view of the hormonal regulation of tuber formation in potato (Solantml tuberosttm) Physiologia Plantarttm 75: 525-531 Wan, W.Y., W Cao & T.W Tibbitts, 1994 Tuber initiation in hydroponically grown potatoes by alteration of solution pH HortScience 29: 621-623 Wheeler, R.M., C.L Mackowiak, J.C Sager, W.M Knott & C.R Hinkle, 1990 Potato growth and yield using nutrient film technique (NFT), American Potato Jottrna167: 177-I87, Wiersema, S.G., R Cabello, P Tovar & J.H Dodds, 1987 Rapid seed multiplication by planting into beds micro tubers and in vitro plants Potato Research 30:117-120 Potato Research 44 (2001) 135

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