CROP NUTRITION IN HYDROPONICS P. Adams Horticulture Research International Littlehampton, West Sussex United Kingdom BNL? 6LP Additional index words Tomato, cucumber, light, temperature, humidity, aeration, salinity Abstract Concern about pollution of the environment by waste nutrients has lead to increasing interest in recirculation of the nutrient solution. In order to optimise the uptake and minimise the wastage of nutrients, inputs must match uptake and good quality water must be used. Nutrient uptake is generally proportional to the concentration of nutrients around the roots but, in flowing solutions, plants are able to make good growth over a wide range of concentrations. Environmental factors can also have a profound effect on nutrition. With tomato and cucumber, the uptake of water, N and K were highly correlated with solar radiation and air temperature while P uptake increased with root temperature. High humidity decreased the Ca and K contents of tomato leaves but increased the Ca content of the fruit. Increasing the salinity by adding NaCl to the solution decreased the K content of tomatoes without adversely affecting the quality. High salinity always decreased the Ca content of tomatoes irrespective of the salts used; blossom-end rot was induced more easily by high levels of major nutrients than by NaCl. 1,_Introduction Management of nutrition for vegetables grown hydroponically depends on whether the system used is open with drainage like rockwool or closed without drainage like NFT. Recirculation of the nutrient solution is now accepted as an important means of minimising pollution of the environment with waste nutrients, notably N and P, as the use of such systems can avoid regular discharges of nutrient solutions into the soil. In order to achieve their full cropping potential, however, more precise information is needed about the nutrient requirements of the plants than is the case when the excess solution is allowed to run to waste. For example, nutrient input must equal plant uptake as there is no leaching, otherwise some nutrients will accumulate while others are depleted. Nutrient uptake is generally proportional to the concentration of nutrients in the solution near the roots; this may be depleted locally in static solutions but can be maintained virtually constant when the solution is flowing. Environmental factors must also be considered as they can have a marked effect on the crop by increasing or reducing nutrient uptake, and by changing the distribution of nutrients within the plant. Acta Horticulturae 323, 1992 Soil & Soilless Media under Protected Cultivation 289In view of the current interest in recycling the nutrient solution, it seems opportune to give an account of the more important aspects of plant nutrition in hydroponics, with the emphasis on the responses obtained at our Institute using recirculating solutions. In this review, therefore, examples have been chosen to illustrate responses to both nutrient levels and environmental factors such as light, air and root temperature, humidity, and the aeration, pH and salinity of the nutrient solution. The value of alternative bases for assessing changes in nutrient status as a result of the applied treatments is also demonstrated. Materials and Methods The experimental plants were grown in NFT systems, except where noted otherwise. Tomato and cucumber plants were propagated in rockwool cubes before being placed either in the NFT channels or on the rockwool slabs. Fully adequate levels of essential nutrients other than those under test were supplied to all crops. The glasshouse atmosphere was enriched with carbon dioxide to 1000 vpm until late April only, when frequent venting rendered it impracticable. Leaf samples consisted of the fifth leaf below the head of the plant. Complete leaves were used for tomato, unless specified differently, but only the laminae were used for cucumber. Values for plant uptake were calculated on the assumption that all nutrients removed from the solution had been absorbed by the plants. All crops were grown under semi-commercial conditions, using local tap water, The pH was controlled at 5.8 for tomato and 6.2 for cucumber by addition of dilute nitric acid. Other experimental details can be found in the publications cited. Resi and Discuss n 3.1. Responses to nutrient concentratior 3.1.1. Major nutrients Plant growth and the yield and quality of the fruit may be affected adversely by inadequate or excessive levels of nutrients. Studies in NFT showed that, in most cases, the responses to nutrient concentration were smaller than expected. Low concentrations of N (10-80 mg/£) that were maintained continuously without depletion had little effect on the growth, yield or quality of tomatoes (Massey and Winsor, 1980). However, the use of NH,-N to assist with pH control affected fruit quality and the uptake of divalent cations. With 40% of the N supplied as NH,-N, almost 20% of the fruit had blossom-end rot (BER) and the Ca and Mg contents of the leaves were reduced by 52 and 31% respectively. The corresponding values with 10% NH,-N were 6% (BER), 19% and 14%. There was little response of cucumber to P over the range 3-81 mg/2. A very high concentration (243 mg/2) reduced the yield, particularly during the second four weeks of harvesting, and reduced the final yield by 11% as compared with that obtained with 3-27 mg P/2 (Massey, Adams and Winsor, 1986). With tomato, increasing the concentration of K in the solution caused a marked reduction in the proportion of tomato fruit with ripening disorders, but had little effect on yield (table 1). The concentration of K in the leaf increased by 63% over the range of K levels tested whereas the total amount of K per fruit (mg) increased by 230104%; the latter appears to be closely related to the increase in the rate of uptake (100%). Studies of nutrient uptake by fruiting tomato plants gave daily absorption rates (mg/plant) of about 115 N, 25 P and 290 K (Adams, 1986). The uptake by cucumber plants was rather higher (543 N, 89 P and 759 K), due to the faster growth rate (Massey, unpublished data). As the acidity of cucumber fruit is very low, the K requirement is considerably less than that of comato, and the K/N ratio is correspondingly lower (Adams, unpublished data). With cucumber, a simultaneous reduction in the applied concentrations of N, P and K from 175, 40 and 300 mg/Z to 60, 15 and 100 respectively had no effect on yield or fruit quality, but the corresponding uptakes decreased by 9, 20 and 14% (Massey and Winsor, unpublished data). Decreasing the Ca and Mg concentrations from 250 and 50 mg/f to 60 and 5 mg/f respectively reduced the final yield after 12 weeks of harvesting by 11 and 16% respectively. The early yields were reduced by 8-9%, but the effect of Mg deficiency became increasingly severe; in the last 4 weeks of harvesting, the reductions due to low Ca and Mg were 17 and 31% respectively (Massey, Adams and Winsor, 1986). The corresponding uptakes of Ca and Mg were reduced by 61 and 80% respectively (Massey, unpublished data). With tomato, the yield of marketable fruit from trusses 1-3 increased with the concentration of Ca in solution over the range 10-70 mg/2 (figure 1) while the incidence of BER decreased from 87% at 10 mg/2 to 9% at 70 mg/2. The data suggest a minimum of 100 mg/£ for healthy crops. These data support the hypothesis that plants can absorb sufficient nutrients for normal growth from flowing solutions at relatively low concentrations (Clement et al., 1974). This may be due to a compensating increase in the rate of uptake, as with P (Clarkson, 1981), in addition to the effect of the flow of solution, which prevents local depletion around the roots. Thus, normal growth is possible when low nutrient concentrations are maintained continuously and never allowed to deplete. There is, nevertheless, a definite relationship between total uptake and nutrient concentration. For example, the uptake of N by cucumbers was closely related to the concentration of N in solution (Schacht and Schenk, 1990) whilst the uptake of K by tomato was linearly related to loge mg K/£ (r-0.99; Adams and Grimmett, 1986). 3.1.2. Micronutrients Essential micronutrients are widely distributed in soil but in hydroponics they must all be supplied at suitable concentrations. The growth, earliness of flowering and early yield of tomatoes were improved by high levels of Fe but there was no effect on the final yield (table 2). The response to B concentrations was also small. The early yield was reduced at 0.1 mg B/£ due to a slight decrease in fruit number, The final yield was lowest at 5 mg/Z (table 3); this concentration was obviously excessive and symptoms of toxicity appeared after 5 weeks of growth in that treatment. Plants grown with only 0.05 mg B/2 developed very severe symptoms of B deficiency and the growing points died after 4 weeks of harvesting (yield, 0.85 kg/plant). However, they made a remarkable recovery when supplied with 0.4 mg/f, and 4 weeks later had become indistinguishable from plants receiving 0.4 mg/2 throughout growth. 291There was little response to Mn, Cu or Zn within the ranges 0.2-1.25 mg Mn/£, 0.5-5 mg Cu/2 and 0.5-10 mg Zn/2. Symptoms of Fe deficiency were induced after several months growth with 3.13 mg Mn/£, With young plants, severe symptoms of Fe deficiency had developed after 3 weeks at 25 mg Cu/2, a concentration that seriously inhibited root growth. Thus, 7 weeks after sowing, the dry weight of the aerial parts of the plants was reduced by 63% and that of the roots by 90% compared with those grown at 0.5 mg Cu/Z, Plant growth was also restricted after 6 weeks at 25 mg Zn/Z, and the symptoms of Zn toxicity had become severe. The dry weight of the tops was reduced by 50% but the weight of root had increased by 38% (Massey, Adams and Winsor, 1986). Unconfirmed observations suggest that Cu and Zn concentrations below 0.2 mg/f are too low for healthy growth in NFT. These data reveal a considerable tolerance of tomato to a relatively wide range of concentrations of most micronutrients, but illustrate the danger of allowing micronutrients to accumulate in the solution, particularly B. 3.2.1, Responses to the light and temperature Light is essential for photosynthesis, which provides material for growth, whereas air temperature controls the rate of growth. Nutrient uptake therefore changes with both the environment and plant size. Tomatoes grown under the same conditions with 1 or 9 trusses in flower absorbed similar amounts of N but the older plants, which carried a heavy load of fruit, required twice as much K: this increased the K/N ratio from 1.24/1 to 2.57/1 (Adams, 1986). Assessments of uptake by young tomato plants (3rd truss in flower) were made every 90 min throughout the day in February, when the glasshouse did not need venting. Water, N and K uptake were highly correlated with solar radiation and air temperature but less well related to solution temperature. The reverse was true for P uptake (table 4). Similar results were obtained at higher light intensities in summer, except that N and K were less well correlated with air temperature (r-0.80-0.82) and solution temperature (r-0.58-0.61; Adams and Massey, 1984). Presumably, these differences were largely due to venting, which results in a poorer relationship between light and air temperature. The increases in uptake rates found during the early part of a growing season include responses to simultaneous increases in growth and solar radiation (table 5); the differences in the rates for the young and mature plants were therefore much greater than those mentioned earlier. Good correlations between solar radiation and N uptake by cucumber were found by Schacht and Schenk (1990), who also calculated different uptake rates for different growth stages. Although the response to root temperature is generally smaller than to light and air temperature, changes in root temperature do have a considerable effect on plant growth and nutrition. For example, the nutrient contents of tomato leaves increased with root temperature between 14-22°C (table 6). Excluding the very low temperature (14°C), the greatest responses were with P and Mg. Recalculation of the data to account for leaf size did not change this pattern of response but increased the differences due to changes in temperature considerably (table 7). Thus the observed magnitude of the response often depends on the basis on which the results were calculated. 292Increasing the root temperature from 18-26°C increased the uptake of water by 23% and the uptakes of N, P, K, Ca and Mg by 8, 33, 12, 16 and 14% respectively. The K/N ratio, however, remained unchanged at 1.72/1 (Adams, 1989). These data demonstrate how much plant nutrient requirements can change with the environment and growth stage. Some understanding of these responses is therefore necessary when anticipating or interpreting changes in nutrient uptake during cropping. At the concentrations of nutrients used for these crops, the uptake of N and K appear to be controlled by transpiration (passive uptake) whereas the uptake of P is controlled actively by the roots. 3.2.2. Responses to humidity In recent years, energy saving in glasshouses has resulted in higher humidities, particularly during the cold weather. The increase in humidity reduces transpiration, and hence Ca movement into the leaves, and may also inhibit growth. High humidity reduced the dry weight of tomato leaves but did not affect the %Ca or %Mg in them (table 8). However, when the data were recalculated on the basis of complete leaves, i.e., taking into account the differences in dry weight, the accumulation of Ca and Mg were both reduced by about 30% at the highest humidity (table 8). The terminal leaflet gave a more sensitive indication of the response to high humidity than the complete leaf (Adams, 1991a). For example, high humidity (>90%) decreased the %Ca in the leaflets by 42% and the Ca content (mg/leaflet) by 68%, compared with 33% and 48% respectively in the complete leaves. High humidity also reduced the %4P and %Mg in the terminal leaflets by 17-20%. The decrease in the K content of both leaves and leaflets was greater, and of similar magnitude to that found with Ca, e.g., 39% for XK and 67% for mg K/leaflet (table 9). The corresponding changes in the complete leaves were much smaller. The reduction in the nutrient of the leaves by high humidity increased with the time of exposure to that environment. One beneficial effect of high humidity was an increase in the Ca content of the fruit. For example, after 6 weeks of constant high humidity (93% RH) the concentration of Ca(%) in ripe tomato fruit was 47% greater than in those grown at constant low humidity (68% RH). This response increased with the concentration of Ca in the nutrient solution (Adams and Holder, 1992). These data demonstrate how effectively high humidity can reduce the movement of nutrients into leaves, particularly to those parts that are furthest from the main stem. Also that leaves and fruit respond differently to changes in humidity; high humidities reduce Ca movement into the rapidly expanding leaves, which results in more Ca moving into the fruit. Low humidities have the opposite effect. 3.2.3. Responses to pH and aeration The main effect of solution pH in the range 5-8 is on nutrient solubility(i.e., availability) whereas aeration affects nutrient uptake more directly. The growth of tomato plants is not greatly affected by pH between pH 5 and 7, but declines rapidly at higher or lower pH values (figure 2; Arnon and Johnson, 1942). The amount of acid requixed to maintain the 293roots at a constant pH was highly correlated with solution temperature (r=0.95; Gislered and Adams, 1983). Tomato plants grown in deep water culture without aeration absorbed 30% less K and 26-28% less NO3-N, P, Ca and Mg than those properly aerated (Arnon and Hoagland, 1940). Now the solubility of oxygen decreases as the temperature rises but there is a simultaneous increase in the rate of respiration of the roots and microflora. Thus, the available supply of oxygen decreases as the demand for it increases. Good aeration is therefore of crucial importance for healthy growth in recirculating solutions. Studies with cucumbers showed that if the solution was poorly aerated in the catchment tank before being circulated through the channels, the decrease in oxygen content (3.5 mg/£) could be considerably larger than that simply due to increased solution temperature(1.9 mg/2; Gislered and Adams, 1983). An example with less serious depletion (2.2 mg/2) is shown in figure 3. The concentration of dissolved oxygen was not significantly correlated with solar radiation or air temperature during a warm day, but was highly correlated with solution temperature (Gislered and Adams, 1983). This was because the solution temperature increased more slowly than air temperature, remained at a maximum for some time after air temperature began to decline, and decreased slowly during the evening. Growth was adversely affected when the oxygen content of the solution fell below 3 mg/Z. Daily additions of glucose to the recirculating solution, which provided more substrate for the micro flora, reduced the oxygen content of the solution to less than 1 mg/Z, particularly during the warmest part of the day (Gislered and Kempton, 1983). Oxygen depletion by tomato was found to be less marked than with cucumber, and the percent saturation of the solution was always higher (table 10). The lower oxygen levels found with cucutber reflect the larger root mat; the dry weight was twice that of tomato roots per unit length of channel. The larger root mat would not only require more oxygen but, by leakage, would also supply more substrate for the microflora. The response to added glucose supports this interpretation. These data suggest that maintenance of good aeration in the root zone depends on minimising the substrate available to the microflora, which are very effective competitors for oxygen. 3.2.4, Responses to salinity Increased salinity reduces water uptake and restricts plant growth. Thus, salinities of 6-8 mS cm? can be used to control the vegetative vigour of tomato plants under poor light conditions. High salinities are generally avoided with more sensitive crops such as cucumber and sweet pepper. Increasing the salinity of the basic nutrient solution from 2.2 to 3.8 mS cm by adding either NaCl or major nutrients had no effect on the yield and quality of cucumbers (Massey and Winsor, unpublished data). There was little effect of this increase in salinity with added NaCl on the uptake of N and P, but that of K uptake was reduced by 16%, and the K/N ratio declined accordingly (table 11). At the same salinity, achieved by addition of extra major nutrients, the uptake of N,P and K increased by 32, 32 and 37% respectively. Reducing the salinity from 2.2 to 1.2 mS cm! by the use of demineralised water had no effect on yield and little effect on nutrient uptake. 294Salinities of 5.5 mS cm and above reduced the dry weight of all parts of cucumber plants and increased the proportion of dry matter partitioned to the fruit (table 12). The Ca content of the leaves was also reduced by the highest salinity. At 5.5 mS em™?, the uptake of water was decreased by 30% compared with that at 3 mS cm; the uptakes of N, P and K were reduced by 52, 21 and 38% respectively. The corresponding reductions at 8 mS cm? were 49, 73, 53 and 71%. Tomato yields were reduced by increasing the salinity from 3 to 8 mS cm? with added NaCl. At 8 mS cm}, the yield during the first four weeks of harvesting was reduced by 25% from 1.86 kg/plant at 3 mS cm? to 1.40 kg/plant, and by 43% from 3.04 to 1.72 kg/plant during harvesting weeks 13-16, The corresponding reductions at 5.5 mS cm? were 15, 20 and 16% (Adams and Ho., 1989, and unpublished data). The uptake of water and of most nutrients was reduced considerably at 8 mS cm™ but only slightly at 5.5 mS em, The uptake of K, however, was seriously decreased at that salinity due to the competing effect of the Na ions; this resulted in a marked decline in the K/N ratio (table 13). Similar responses were obtained with tomatoes grown in rockwool (Adams, 1991b). The unpublished data for the K content of the fruit illustrate the response to increasing the salinity with either major nutrients or NaCl. The concentration of K in the fruit juices increased with the salinity, except at 8 mS cm? with added NaCl (figure 4a). However, the concentration of K in the dry matter increased with extra major nutrients but decreased with increasing additions of NaCl (figure 4b). This decline was linearly related to salinity when the K content was expressed as mg K per fruit (figure 5). The total Ca content of these fruit was depressed more by the extra major nutrients than by NaCl, despite the higher concentration of Ca in the nutrient solution (e.g. 530 mg/2 compared to 267 mg/2 with NaCl, for plants grown at 8 mS cmt), The corresponding values for K and Na in solution. were 1750 and 425 mg K/2 and 45 and 1200 mg Na/2. This supports the notion that the uptake of Ca is depressed more seriously by K than by Na. The distribution of Ca in tomato fruit is uneven; the concentration in the dry matter is highest near the calyx and lower at the distal (blossom) end. Thus, analysis of a disc from that end of the fruit may be a better indicator of Ca status, and hence susceptibility to blossom- end rot, than analysis of the complete fruit. For example, increasing the salinity from 3-8 mS cm! reduced the %Ca in the distal dise by 52% (from 0.094 to 0.045% Ca); the corresponding values for the rest of the fruit were 31% (0.115 and 0.079% Ca; Adams and Ho, 1989). Studies of tomatoes grown at an exceptionally high salinity (17 mS cm) indicated that very high concentrations of Na in the nutrient solution (4500 mg/l) do not induce BER when there is no extra water stress due to high temperature, provided that the solution contains an adequate amount of Ca (230 mg/2 in this case). Plants grown at the same salinity, achieved by adding Nos-N, K and Ca to increase the salinity of the basic solution from 3 to 17 mS cm™, produced a high proportion of fruit with BER during April and May (table 14); with this treatment, the average Ca concentration in solution throughout March, April and May was 1388 mg/f while that of K was 2429 mg/2. This also supports the notion that the antagonism between K and Ca is greater than that between Na and Ca. These data illustrate responses to salinities that were maintained at constant values in NFT; this was possible because of good water quality. Use of water containing appreciable concentrations of Na and Cl will 295inevitably result in a continuous rise in the salinity of the recirculating solution. In these circumstances, the efficiency of water usage will be greatly reduced by the frequent need to replace part of the solution with some fresh solution of low NaCl content. In this way, or by means of a small but continuous leak, it is possible to maintain the salinity within an acceptable range of variation. However, disposal of the unwanted solution may be difficult without causing pollution to the environment. 4, Conelud: rks Complete leaves or fruit are not necessarily the best choice for analysis when assessing their nutrient status in relation to localised nutrient deficiencies. In some cases a part of the leaf or fruit may be more appropriate. Changes in the growing environment affect plant growth as well as nutrient uptake. Therefore the evaluation of a response to environmental changes is incomplete unless it includes the effect of any change in dry matter content. Thus, recalculation of analytical data to give the weight of nutrients per leaf or fruit often reveals responses not apparent in the original analyses. Plants can grow well at lower nutrient concentrations in flowing solutions than in static ones. With tomato, however, low levels of K do not favour good fruit quality. Furthermore, low nutrient concentrations result in lower salinities which may be undesirable for fruit development, for instance. A favourable salinity could be achieved by addition of NaCl; this would result in a proportional decrease in the K content of the fruit, but would not affect the flavour or quality adversely. In practice, the concentrations of major nutrients used for tomato in NET are very similar to those used in rockwool, with a K/N ratio of at least 1.5/1, e.g., 180 N, 30 P, 325 K, 75 Mg and 200 Ca (mg/2). For cucumber, lower concentrations such 100 N, 20 P, 140 K, 50 Mg and 150 Ca mg/2 suffice. Recirculating solutions can be used successfully to reduce environmental pollution provided that good quality water is available and nutrient additions match uptake by the plants. Then the solution can be retained throughout cropping. By allowing gross depletion of the nutrient concentrations during the final few days of a crop, the disposal of nutrients becomes minimal. The optimisation of factors such as the temperature and humidity of the aerial environment, and aeration, salinity and temperature in the root zone is essential if the full potential of hydroponic culture for plant growth, fruit yield and quality is to be achieved. References Adams, P., 1986. Mineral nutrition. In: The Tomato Crop, eds. Atherton, J.G. and Rudich, J. Chapman and Hall, London: 281-334. Adams, P., 1988. Some effects of environment on the calcium status of tomato leaves. CEC Workshop, The effects of high humidity on plant growth in energy saving greenhouses. Report EUR 11261, Brussels: 61-70. Adams, P., 1989. Some effects of root temperature on the growth and nutrient uptake of tomatoes in NFT. Proc. 7th Int. Cong. Soilless Culture, Flevohof, 1988, ISOSC, Wageningen:73-82. 236Adams, P., 1991a. Effect of diurnal fluctuations in humidity on the accumulation of nutrients in the leaves of tomato (Lycopersicon esculentum). J.Hort.Sci. 66:545-550. Adams, P., 1991b. Effects of increasing the salinity of the nutrient solution with major nutrients or sodium chloride on the yield, quality and composition of tomatoes grown in rockwool. J.Hort.Sci. 66:201-207. Adams, P., and Grimmett, M.M., 1986. Some responses of tomatoes to the concentration of potassium in recirculating nutrient solutions. Acts Hort. 178:29-35. Adams, P., and Ho. L.c., 1989. Effects of constant and fluctuating salinity on the yield, quality and calcium status of tomatoes. J.Hort Sei, 64:725-732. Adams, P., and Holder, R., 1992. Effects of humidity, Ca and salinity on the accumulation of dry matter and Ca by the leaves and fruit of tomato (Lycopersicon esculentum). J.Hort.Sci, 67:137-142. Adams, P., and Massey, D.M., 1984, Nutrient uptake by tomatoes from recirculating solutions. Proc. 6th Int. Cong. Soilless Culture, Lunteren, 1984, ISOSC, Wageningen:71-79. Arnon, D.I., and Hoagland, D.R., 1940. Crop production in artificial culture solutions and in soils with special reference to factors influencing yield and adsorption of inorganic nutrients. Soil Sci. 50:463-483, Arnon, D.I., and Johnson, C.M., 1942. Influence of hydrogen ion concentration on the growth of higher plants under controlled conditions. Pl. Physiol. 17:525-539. Clarkson, D.T., 1981. Phosphate transport in phosphate-stressed tomato plants. Rep. Letcombe Lab., 1980:58-60. Clement, C.R., Hopper, M.J., Canaway, R.J., and Jones, L.H.P., 1974. A system for measuring the uptake of ions by plants from flowing solutions of controlled composition. J.Exp.Bot. 25:81-99. Gislered, H.R., and Adams, P., 1983. Diurnal variations in the oxygen content and acid requirement of recirculating nutrient solutions and in the uptake of water and potassium by cucumber and tomato plants. Scientia Hort. 21:311-321. Gislered, H.R., and Kempton, R.J., 1983. The oxygen content of flowing nutrient solutions used for cucumber and tomato culture. Scientia Hort. 20:23-33. Massey, D.M., Adams, P., and Winsor, G.W., 1986. Five years’ work on NFT. Grower 105, BGLA Supplement, 23 January:90,93,94,97 and 99. Massey, D.M., and Winsor, G.W., 1980. Some responses of tomatoes to nitrogen in recirculating solutions. Acta Hort. 98:127-137. Schacht, J., and Schenk, M.K., 1990. Control of nitrogen supply of cucumber (Cucumis sativus) grown in soilless culture. In: Plant Nutrition, Physiology and Applications, ed. Beusichem, van, M.L. Kluwer, Dordrecht :753-758. 297Table 1 - Effect of K concentration on the yield, quality, K content and rate of K uptake of tomato plants (After Adams and Grimmett, 1986, with some recalculated data). K Yield Ripening K in leaf K per fruit Daily uptake (mg/£) (kg/plant) disorders (%) (%) (ng) (mg/plent) * After 14 weeks of harvesting. Table 2 - Effect of Fe concentration on the growth, earliness of flowering and yield of tomatoes (After Massey, Adams and Winsor, 1986). Plant height at 42 d (cm) 68 72 75 74 Early flowers open* 0.3 0.6 1.0 1.5 Open flowers and set fruitt 7 d later 4.5 5.0 6.7 71 Yield in weeks 1-5 (kg/plant) 136) 152 ol 172 9.83 10.40 10.40 10.00 Yield in weeks 1-28 (kg/plant) * Truss 2. * As FeEDTA. Table 3 - Effect of B concentration on the yield and size of tomatoes (After Massey, Adams and Winsor, 1986). B in solution (mg/2) Harvesting ------- weeks ol 0% Early yield (kg/plant) 1-4 2.06 2.49 2,29 2.36 2.20 Final yield (kg/plant) 1-27 ie ie ie ie Weight/fruit (g) 1-27 70 66 65 66 62 298Table 4 - Correlations between tomato plant uptake and environmental variables in late February (truss 3 in flower; Adams, unpublished data). Solar radiation 0.91 0.91 0.90 0.48 Air temperature 0.95 0.97 0.96 0.58 Root temperature 0.88 0.81 0.85 0.89 Table 5 - Combined effects of plant age and light intensity on water and nutrient uptake by tomato plants between mid-February and early April (mi or mg/plant; Adams, unpublished data). Truss in Solar flower radiation Water N P K K/N (My mv? dt) 240 49 Sor 1.16 1. 15 4 4.3 468 89 24 144 1.62 7 5.9 727 130 30 317 2.44 Table 6 - Effect of root temperature on the nutrient concentrations in tomato leaf laminae after four weeks of treatment (mid- February; after Adams, 1989). Nutrient content (%) Root temperature (°C) N Pp K Ca Mg, 96 0.61 1.16 1.37 0.27 14 L 18 2.11 0.57 1.63 1.75 0.35 22 2.72 0.83 1.86 1.91 0.43 26 2.30 om 1.75 1.99 0.42 % Increase between 18-26° 9 25 7 14 20 299Table 7 - Effect of root temperature on the dry weight and total nutrient contents of the tomato leaf laminae shown in Table 6. temperature (°C) weight (g) N P K Ca Mg 16 1.07 a1 6 12 15 3 18 1.20 25 720 2G 22 1.59 a3) is 6g 26 1.79 6 is Gi eB % Increase between 18-269 49 64 86 = 5542100 Table 8 - Effect of humidity on the dry weight and Ca and Mg contents of complete tomato leaves after four weeks of treatment (Adams, 1988, with some recalculated data). Vapour pressure % RH Dry weight mg Ca mg Mg deficit (kPa) at 20°C = /leaf (g) %Ca /leaf “Mg /leaf 0.65 68 1.47 0.88 12.9 0.23 3.38 0.43 78 1.29 0.87 9 11.2 0.24 3.10 0.24 88 1.29 0:62 103 0.23 2.88 0.15 93 1.02 0.88 9.0 0.24 2.44 % Decrease 31 0 30 4 28 between 0.65-0.15 Table 9 - Effect of humidity on the concentration and total content of K in complete tomato leaves and in the terminal leaflet only after four weeks of treatment (mid-February; after Adams, 1991a). Terminal leaflet 300Table 10 - Oxygen status (% saturation) of NFT solutions in relation to crop and time of day (After Gislered and Adams, 1983), Time of day (h) 06.30 17.00 23.00 Table 11 - Effect of salinity and salinity source on the uptake of nutrients by cucumber plants six weeks after sowing (Massey and Winsor, unpublished data). Demineralised water 1.2 0.62 5.3 1.39 Tap water (control) 2.2 O76 = 51 15, Tap water plus extra major nutrients 3.8 Tis 70 120 Tap water plus NaCl 3.6 0.71 4.3 1.19 Table 12 - Effect of salinity on the distribution of dry matter and Ca in cucumber plants harvested seven weeks after sowing in August. The salinity was increased by addition of NaCl (Ho and Adams; full data to be published elsewhere). Dry weight (g/plant) % Ca in leaves EG Fruit 3 84 6 56 38 5.4 10.6 5.5 62 4 52 44 5.3 10.5 8 35 2 4. 53 3.9 8.0 301°Table 13 - Effect of salinity on the total uptake of water and nutrients by tomato plants during four weeks in April (2 or g/plant; Adams and Ho, unpublished data) : Table 14 - Effect of salinity source on the proportion (%) of fruit harvested with blossom-end rot. All plants were grown at a constant salinity of 17 mS cm? of which essential nutrients contributed 3 mS cm (Adams and Ho, unpublished data). 302Maotabe iad (kota) — ° 100 200 300 Cain soltion (mai) Figure 1 - Relation between the marketable yield of tomatoes from trusses 1-3 and the Ca content of the recirculating solution (After Massey, Adams and Winsor, 1986). 100 oN - Shoots Bo 7 - 0 2 Roots : — eA ° ‘ 5 6 7 8 9 1H of auton sottion Figure 2 - Effect of solution pH on the growth of tomato plants (After Arnon and Johnson, 1942). 3030 30 saturation onvcentation Solution temperate (0) Dissowed 0, (mg ") ol S&S 0 eee | 06.00 08.00 10.00 1200 1400 16.00 18.00 20.00 22.00 24.00 ie o by Figure 3 - Changes in the oxygen content and temperature of the NFT solution during a bright day, with the corresponding values for water saturated with oxygen (cucumber; Gislered and Adams, unpublished data). 240 i K (mgt 60 120 4 6 8 10 2 EC of rutsent gluion (mS em) Figure 4 - Effect of increasing the salinity with NaCl on the total K content (mg) of tomato fruit (Adams, unpublished data). 304content (maq/00m ice} 50 K (mp'@ ary mater, EC of nutiontsokaton (ms em") Figure 5 - Effect of increasing salinity with major nutrients (@) or NaCl (M@) on (a) the K content of the expressed juices from tomato fruit and (b) the K content of the dry matter (Adams, unpublished data). 305