Effect of CaSO4 and NaCl on mineral content ofLeucaena leucocephala

The effects of varying CaSO₄ and NaCl levels on the nutrient content ofLeucaena leucocephala were established by examining the concentrations of Na, Ca, Cl, K and Mg in leucaena roots, stems and leaves. Leucaena was grown in nutrient solution at four levels of CaSO₄ (0.5, 1.0, 2.5 and 5.0 mM) and NaCl (1, 25, 50 and 100 mM), in randomized blocks with five replications. Leucaena excluded sodium from stems and leaves when NaCl concentration was 50 mM or less. Sodium uptake decreased as CaSO₄ concentration increased. Calcium uptake was affected by NaCl concentration when substrate CaSO₄ concentration was 0.5 mM. At this level, 100 mM NaCl caused a marked decrease in leaf calcium and a marked increase in leaf Cl. In all other treatments, Cl uptake was not affected by CaSO₄ concentration. Potassium uptake was strongly depressed as NaCl concentration increased at low Ca concentration, but this effect was offset at high Ca. Magnesium uptake decreased as CaSO₄ levels increased.     

Response of mesquite to nitrate and salinity in a simulated phreatic environment: Water use,dry matter and mineral nutrient accumulation

Mesquite plants (Prosopis glandulosa var. Torreyana) were grown in 2-m long columns 20 cm in diameter, and provided with a constant, stable ground water source 10 cm above the sealed base of the column. Ground water contained 0, 1 or 5 mM nitrate, or a mixed salt solution (1.4, 2.8, or 5.6 dS m^-1) with the ionic ratios of ground water found in a field stand of Prosopis at Harper's Well (2.8 dS m^-1). Water uptake in the highly salinized columns began to decrease relative to low salt columns when soil salinity probes 30 cm above the column base read approximately 28 dS m^-1, a potential threshold for mesquite salt tolerance. Prosopis growth increased with increasing nitrate, and decreased with increasing salinity. Water use efficiency was little affected by treatment, averaging approximately 2 g dry matter L^-1 water used. Most fine roots were recovered from a zone about 25 cm above the ground water surface where water content and aeration appeared to be optimal for root growth. Root-shoot ratio was little affected by nitrate, but increased slightly with increasing salinity. Plant tissue P concentrations tended to increase with increasing salinity and decrease with increasing N, approaching potentially deficient foliage concentrations at 5 mM nitrate. The whole-plant leaf samples increased in sodium concentration both with added salt and with added nitrate. Foliar manganese concentrations increased with increasing salt in the absence of nitrate. Concentrations of sodium in leaves were low (<10 g kg^-1), considering the high salt concentrations in the ground water. Prosopis appears to exclude sodium very effectively, especially from its younger leaves. Although Prosopis is highly salt tolerant, the degree to which it utilizes soil nitrate in place of biologically fixed N may lower its salinity tolerance and affect its nutrient relations in phreatic environments.     

Enhancement of Intact Bean Leaf Senescence by NaCl Salinity

Red kidney bean (Phaseolus vulgaris L.) plants were grown in nutrient solution and in nutrient solution plus four bars of added NaCl. Chlorophyll and protein decay occurred much more rapidly in intact leaves from plants subjected to four bars of added NaCl in the growth medium than in intact leaves from plants without added NaCl. Ribonucleic acid (RNA) content in intact leaves of salt treated plants was higher than in intact leaves from plants grown in nutrient solution alone. However, the tendency for RNA content variation in leaves during the experimental period was the same for both control and salt treated plants. The results support the idea that salinity enhances senescence and suggests that hormone imbalance plays an important role in this process.     

Effects of CaSO4 and NaCl on growth and nitrogen fixation ofLeucaena leucocephala

Effects and interactions of varying CaSO₄ and NaCl levels on growth and nitrogen fixation ofLeucaena leucocephala K8 were examined. Leucaena was grown in nutrient solution at four levels of CaSO₄ (0.5, 1.0, 2.5 and 5.0 mM) and NaCl (1, 25, 50 and 100 mM) in randomized blocks with five replications. While NaCl significantly reduced plant growth, additions of CaSO₄ increased plant height, leaf number, and biomass of salt treated plants. For the nonsaline treatments, high CaSO₄ levels slightly depressed growth, which contradicts suggestions that Leucaena has a high calcium requirement. A significant calcium/sodium interaction was not seen for nodule number or weight. Nodule number was significantly depressed by 100 mM NaCl and nodule weight of the salt stressed plants significantly increased as CaSO₄ concentration increased from 0.5 to 2.5 mM. Effects of NaCl and CaSO₄ on nitrogen content of plant parts were inconclusive. The promotion of Leucaena salinity tolerance by addition of CaSO₄ may be attributed to the effect of calcium in maintaing the selective permeability of membranes.     

Salinity-calcium interactions on growth and ionic concentration of Citrus plants

Two-year-old Navel orange scions (Citrus sinensis (L.) Osbeck) budded to either Cleopatra mandarin (C. reticulata) and Troyer citrange (C. sinensis × P. trifoliata) rootstocks were used in this experiment. Cleopatra manda in rootstock was considered more tolerant to salinity than Troyer citrange, and this property was attributed to a greater capacity to exclude chloride ions.Plants were grown under glasshouse conditions and supplied with nutrient solution containing either no or 45 mM NaCl. Calcium concentration was increased from 3 to 30 mM. Sodium, potassium, calcium and chloride concentrations in plant organs were analyzed after 90 days of treatment.Supplemental Ca was found to mitigate the adverse effects of salinity on plant growth, defoliation or leaf injury.Chemical analysis indicated that in plants grafted on Troyer citrange Ca restricted uptake and subsequent translocation of Na to the leaves and increased K concentration in both roots and leaves. However, in Cleopatra mandarin-grafted plants increasing Ca levels seemed to reduce transport of Na from roots to leaves, and Na accumulation in roots was associated with reduced concentration of K in this rootstock.Organ chloride analysis showed that Cl accumulation in leaves of plants grafted on both rootstocks was reduced when external Ca concentration increased, whereas Cl concentration in roots remained constant or increased. The data of distribution of Cl in plants showed that a high external Ca level increased Cl accumulation in the basal stem and roots, and reduced the transport of Cl from roots to leaves.     

Effects of silicon on enzyme activity and sodium,potassium and calcium concentration in barley under salt stress

Two contrasting barley (Hordeum vulgare L.) cultivars: Kepin No.7 (salt sensitive), and Jian 4 (salt tolerant) were grown in a hydroponics system containing 120 mol m^-3 NaCl only and 120 mol m^-3 NaCl with 1.0 mol m^-3 Si (as potassium silicate). Compared with the plants treated with salt alone, superoxide dismutase (SOD) activity in plant leaves and H⁺-ATPase activity in plant roots increased, and malondialdehyde (MDA) concentration in plant leaves decreased significantly for both cultivars when treated with salt and Si. The addition of Si was also found to reduce sodium but increase potassium concentrations in shoots and roots of salt-stressed barley. Sodium uptake and transport into shoots from roots was greatly inhibited by added Si under salt stress conditions. However, Si addition exhibited little effect on calcium concentrations in shoots of salt-stressed barley. Thus, Si-enhanced salt tolerance is attributed to selective uptake and transport of potassium and sodium by plants. The results of the present study suggest that Si is involved in the metabolic or physiological changes in plants. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effects of aluminium on growth and nutrient uptake of small Picea abies and Pinus sylvestris plants

Summary The effects of aluminium concentrations between 0.2 and 30 mM at pH 3.8 ±0.2 on small plants of Norway spruce [(Picea abies (L.) Karst], Scots pine (Pinus sylvestris L.), and Scots pine infected with the ectomycorrhizal fungus Suillus bovinus (L. ex Fr.) O. Kuntze were investigated. The plants were grown at maximum relative growth rate (R_G % day^–1) with free access but very low external concentrations of nutrients. Steady-state conditions with respect to relative growth rate (R_G) and internal nutrient concentrations were achieved before addition of aluminium, which was added as AlCl₃ and/or Al(NO₃)₃. There were reductions in rg at aluminium concentrations of 0.3 mM in spruce, 6 mM in pine and 10 mM in ectomycorrhizal pine, i. e. at aluminium concentrations considerably higher than those normally occurring in the top layer of the mineral soil where most fine roots are found. Nutrient uptake rate per unit root growth rate was calculated for different nutrient elements. The uptake rate of calcium and magnesium was reduced at aluminium concentrations of 0.2 mM (spruce), 1 mM (pine) and 3 mM (ectomycorrhizal pine), without influencing R_g. The results question the validity of the hypothesis of aluminium toxicity to forest tree species at low external concentrations.     

Importance of seed Zn content for wheat growth on Zn-deficient soil

Eggplants (Solanum melongena L. cv. Bonica) were grown in a glasshouse during summer under natural light with one unbranched shoot or one shoot with 3 to 4 branches and with or without fruit in quartz sand buffered and not buffered with 0.5% CaCO₃ (w : v), respectively. Nutrient solutions supplied contained nitrate or ammonium as the sole nitrogen source. Compared with nutrient solutions containing nitrate (10 mM), solutions containing ammonium (10 mM) caused a decrease in net photosynthesis of eggplants during early stages of vegetative growth when grown in quartz sand not buffered with CaCO₃. The decrease was not observed before leaves showed interveinal chlorosis. In contrast, net photosynthesis after bloom at first increased more rapidly in eggplants supplied with ammonium than with nitrate nitrogen. However, even in this case, net photosynthesis decreased four weeks later when ammonium nutrition was continued. The decrease was accompanied by epinasty and interveinal chlorosis on the lower leaves and later by severe wilting, leaf drop, stem lesions, and hampered growth of stems, roots, and fruits. These symptoms appeared later on plants not bearing fruits than on plants bearing fruits. If nutrient solutions containing increasing concentrations of ammonium (0.5–30 mM) were supplied after the time of first fruit ripening, shoot growth and set of later flowers and fruits were promoted. In contrast, vegetative growth and reproduction was only slightly affected by increasing the concentration of nitrate in the nutrient solutions. In quartz sand buffered with CaCO₃ ammonium nutrition caused deleterious effects only under low light conditions (shade) and on young plants during rapid fruit growth. If eggplants were supplied with ammonium nitrogen before bloom, vegetative growth was promoted, and set of flowers and fruit occurred earlier than on plants supplied with nitrate. Furthermore, the number of flowers and fruit yield increased. These effects of ammonium nutrition were more pronounced when plants were grown with branched shoots than with unbranched shoots. The results indicate that vegetative and reproductive growth of eggplants may be manipulated without causing injury to the plants by supplying ammonium nitrogen as long as the age of the plants, carbohydrate reserves of the roots, quantity of ammonium nitrogen supplied, and pH of the growth medium are favourable. T W Rufty Section editor     

Zinc-binding peptides as a function of Zn and S in soybeans

Soybean (Glycine max [L.] Merr. cv. Ransom) plants were grown for two weeks in nutrient solutions at 0.8 and 40 M Zn and 0.02 and 20 mM S. Zinc toxicity appeared to induce synthesis of low molecular weight Zn-binding peptides in soybean roots and leaves. This binding of Zn-ions seemed to increase in leaves when the S concentration was high in the nutrient solution.     

The influence of ammonia in nutrient solution on growth and metabolism of cucumber plants

Summary Shoot yield of cucumber plants grown 18 days in nutrient solution with 0.06 mM NH₃ was decreased. Root yield was diminished at 0.09 mM NH₃ The ammonia treatment caused heavy chlorosis increasing with age of leaves. This chlorosis was not due to any nutrient deficiency. Ammonia also influenced the morphology of roots. They were clearly shorter caused by a much smaller size of root cells.The decrease of yield was linked to a reduction of assimilation occurring not only after a long influence of ammonia lasting 14 days, but also within one hour after starting the NH₃ treatment. The decline of assimilation was probably caused by a higher resistance of stomata against CO₂ influx in leaf tissue as can be concluded from the observation that transpiration was decreased in the same way as assimilation.The effect of ammonia in nutrient solution could also be due to the occurrence of higher NH₃ concentrations in leaf tissue, because both, pH of plant press sap as well as NH₄ concentration of plant tissue, were increased.Furthermore, it is shown that the nitrate content of plant tissue was diminished by ammonia whereas ammonium and amide content were raised. Regulation of nitrate uptake of plants by means of ammonium and amide content of tissue is discussed.     

Changes in the concentration of phenolic compounds and exudation induced by phosphate deficiency in bean plants (Phaseolus vulgaris L.)

The effect of prolonged phosphate starvation of bean plants (Phaseolus vulgaris L.) on the concentration of phenolics and their exudation by roots was studied. Plants cultured on phosphate-deficient media maintained a steady concentration of total phenolics in the leaves, whereas in the leaves of plants grown on complete nutrient media the phenolic concentration decreased. After 18 days of culture, higher total phenolics and anthocyanin concentrations in phosphate-deficient leaves compared with control leaves were observed. The divergent trends in total phenolic concentrations between phosphate-deficient and control leaves corresponded to the changes in the activity of L-phenylalanine ammonia-lyase. In the roots, the concentration of total phenolics was lower in phosphate-deficient plants compared with control plants. However, after 18 days of culture of bean plants, the amount of exuded phenolics from phosphate-deficient roots was 5-times higher than that from the roots of control plants. The activity of L-phenylalanine ammonia-lyase was twice as high in the roots of phosphate-starved plants. Comparable rates in the exudation of phenolics by bean roots observed after 18 days of culture on nitrogen-deficient or phosphate-deficient medium may suggest a similar system of signal transduction for phenolics release. The results are discussed in relation to the possible functions of phenolics in nutrient uptake and as chemical signals in root-soil microbe interactions to enhance the plant adaptation to particular environmental conditions.     

A comparative study of the tolerance of salt marsh plants to manganese

Summary Salicornia europaea, Puccinellia maritima, Triglochin maritima, Aster tripolium, Plantago maritima, Armeria maritima, Juncus gerardii andFestuca rubra, collected as seed from a salt marsh at Portaferry, County Down, were grown on saline (340 mM NaCl) and non saline nutrient solutions at five concentrations of manganese sulphate (0.025–10.0 mM). After an eight week growing period, shoot and root yields and the concentrations of sodium, potassium, calcium and manganese in the shoots were determined.Except forS. europaea the saline treatments had a strongly limiting effect on plant growth. Each of the species investigated showed a degree of tolerance to high concentrations of manganese which was similar to that of calcifuge species and plants characteristic of waterlogged sand dune slack communities, but which was very much greater than that ofArrhenatherum elatius a species usually excluded from acidic soils. There was little evidence to support the hypothesis that tolerance of high manganese concentrations was correlated with the position of the experimental plants in the salt marsh ecotone or that the manganese nutrition of halophytic and glycophytic marsh species differs. Whilst manganese uptake increased proportionally with solution manganese concentration, there were few other major effects of manganese on the balance of shoot cation concentrations in the plants investigated. Both antagonistic and synergistic effects of sodium on manganese uptake were recorded for different species.     

Uptake and translocation of zinc by intact plants

Summary Certain cations were added to nutrient solution cultures, in which intact bean plants were being grown, in an attempt to establish whether they influenced either entry of radiozinc (Zn*) into these plants or translocation of this element from the roots to the tops, or both. The Zn*-level in the culture solution was maintained at 5µM and the concentrations of the added cations were varied from zero through 60µM in the highest treatments.When copper was present as the added cation at the Zn*-level (5µM), Zn*-uptake was severely reduced but internal translocation was not especially altered.When zinc was present as the added cation it exerted the expected competition on Zn*-uptake and in addition exerted an unexpected, suppressing effect on Zn* being translocated to the tops.Manganese, as an added cation, has an effect on Zn*-uptake only at high concentrations (30µM or more) but it did not alter the internal Zn* distribution.Cadmium behaved similarly to copper in its influence on Zn*-uptake and translocation.It is concluded that the general effect of added cations on Zn*-uptake and translocation in intact bean plants is predominantly to inhibit uptake, not internal distribution.Supported in part by a grant from the United States Atomic Energy Commission.     

Potassium and phosphate uptake of cucumber plants at different ammonia supply

Summary Experiments on cucumber plants grown in nutrient solution were conducted in order to study long and short time effects of ammonia on growth, nutrient element uptake and respiration of roots.Shoot yield and potassium concentration in tissue of plants treated 18 days with varied ammonia concentration were decreased. However, it was not assumed that K deficiency caused the yield reduction. The ammonia effect on K content was more pronounced in roots than in shoots.The decreased K concentration of plant tissue was linked to a diminished ability of plant roots to absorb potassium. The maximum rate of potassium uptake was lowered by ammonia during both, long- and short-time treatment. The results indicated that the NH₃ influence on potassium uptake was due to effects on metabolism and permeability of roots because changes of K uptake rate occurred immediately after starting the NH₃ treatment. Furthermore, it is shown that ammonia inhibited respiration of roots.During the short-time treatment net potassium efflux of roots was observed at higher NH₃ concentrations. The extent of K efflux depended on K concentration of both, root tissue and nutrient solution.Pretreating the plants for 12 hours with ammonia also resulted a decline in K uptake rate. However, plant roots subjected to ammonia concentrations up to 0.09 mM completely recovered during 24 hours after removing the NH₃ treatment whereas at higher NH₃ concentrations only a partial recovery occurred.Furthermore, it was shown that ammonia also influenced P uptake by plant roots.     

Effects of Pyrenochaeta lycopersici infection on nutrient uptake by tomato plants

The growth of young tomato plants in nutrient solution or in soil and infected with Pyrenochaeta lycopersici Schneider & Gerlach, the cause of tomato brown root rot, was decreased relative to that of uninfected plants. The roots of plants grown in nutrient solution and infected with a mycelial mat of the pathogen contained lower concentrations of potassium and higher concentrations of calcium than roots of uninfected plants. These changes occurred largely in the visibly affected tissue, as opposed to the root system as a whole. The concentrations of magnesium, total nitrogen and phosphorus in the roots of infected plants were not significantly different from those of control plants. Magnesium, nitrogen and phosphorus concentrations in the tops of infected plants were also not significantly different from those of healthy plants, but no consistent changes were found in the concentrations of calcium and potassium. Young tomato plants grown in soil infested with P. lycopersici contained lower concentrations of phosphorus and potassium in the tops than plants grown in sterilized soil. It was not possible to separate intact damaged root systems of infected plants from soil. The changes in composition found in infected plants are discussed in relation to possible methods of manipulating the nutrition of the plant to offset the effects of the disease on crop yield.     

Impact of auxin-compounds produced by the antagonistic fungus Pythium oligandrum or the minor pathogen Pythium group F on plant growth

Plant growth promotion induced by the antagonistic fungus, Pythium oligandrum, is the result of a complex interaction which includes an indirect effect through control of pathogens in the rhizosphere and/or a direct one mediated by plant-induced resistance. The present study shows an increased plant growth associated with direct interaction between P. oligandrum and roots, which is mediated by a fungus-produced auxin compound, tryptamine (TNH₂). In vitro experiments provided evidence that P. oligandrum metabolised specifically indole derivatives, such as tryptophan and indole-3-acetaldehyde, to produce THN₂ through the tryptamine pathway. When P. oligandrum grew in sterile root exudates, it also produced an auxin-like compound. Additional experiments on P. oligandrum–root interaction showed that, in amended nutrient solution of plants, the antagonist metabolised Trp into TNH₂ and that root absorption of this newly formed auxin-compound in appropriate concentrations was associated with enhancement of plant growth. This phenomenon was observed only when nutrient solution was amended with low tryptophan (Trp) concentrations, i.e. 0.05 and 0.1 mM; higher concentration (0.5 and 1 mM Trp) induced abnormal root development. Similar experiments were performed with Pythium group F, a minor pathogen known for its ability to produce auxin-compounds through the tryptamine pathway. In this case, irregular root development was always noticed with all Trp concentrations added to the nutrient solution of plants. Moreover, Pythium group F colonization of roots was associated with leakage of auxin-compounds in the nutrient solution. Our results, therefore, highlight that the production of similar auxin-compounds by two Pythium species has contrary effects on plant development.     

Effects of aluminium on growth and root reactions of phosphorus stressed Betula pendula seedlings

The impact of two constant non-toxic levels of Al stress (0.2 and 0.4 mM) on growth and ³²P uptake capacity on sub-optimal (P-limited) Betula pendula seedlings grown in sand culture was examined.Seedling growth was under optimum controlled conditions in a growth chamber where nutrient additions were made at a predetermined relative addition rate (R_A) of 10% day^-1. Three treatment groups of seedlings 0, 0.2 and 0.4 mM Al were harvested at 15, 29 and 42 days. The excised roots were exposed to a ³²P-labelled solution for 15 minutes to measure their capacity for P uptake. Growth was determined by weighing the roots, stems and leaves of the seedlings.Growth data showed that relative growth rate (R_G) should equal the R_A of P the most limiting nutrient, which was supplied at P/N 3% instead of an optimal 15%. Therefore, Ingestad's theory can also be used succesfully in sand culture and this may be particularly important for future studies of root and rhizosphere exudates. Low levels of Al (< 0.2 mM) in combination with low P supply significantly lowered the R_G of the birch seedlings by further reducing P supply. However, previous studies of birch seedling growth and nutrient uptake using Ingestad's solution culture technique with optimumal P supply did not show any effect of Al on growth untill the Al was in excess of 3 mM. Aluminium was not directly toxic to the plants and therefore roots could respond to the ³²P bioassay.     

Regulation of hormonal responses of sweet pepper as affected by salinity and elevated CO2 concentration

This study examines the extent to which the predicted CO₂‐protective effects on the inhibition of growth, impairment of photosynthesis and nutrient imbalance caused by saline stress are mediated by an effective adaptation of the endogenous plant hormonal balance. Therefore, sweet pepper plants (Capsicum annuum, cv. Ciclón) were grown at ambient or elevated [CO₂] (400 or 800 µmol mol^–1) with a nutrient solution containing 0 or 80 mM NaCl. The results show that, under saline conditions, elevated [CO₂] increased plant dry weight, leaf area, leaf relative water content and net photosynthesis compared with ambient [CO₂], whilst the maximum potential quantum efficiency of photosystem II was not modified. In salt‐stressed plants, elevated [CO₂] increased leaf NO₃^– concentration and reduced Cl^– concentration. Salinity stress induced ABA accumulation in the leaves but it was reduced in the roots at high [CO₂], being correlated with the stomatal response. Under non‐stressed conditions, IAA was dramatically reduced in the roots when high [CO₂] was applied, which resulted in greater root DW and root respiration. Additionally, the observed high CK concentration in the roots (especially tZR) could prevent downregulation of photosynthesis at high [CO₂], as the N level in the leaves was increased compared with the ambient [CO₂], under salt‐stress conditions. These results demonstrate that the hormonal balance was altered by the [CO₂], which resulted in significant changes at the growth, gas exchange and nutritional levels.     

The role of sucrose and nitrogen in adventitious root formation on cultured rose shoots

Cultured shoots ofRosa ‘Improved Blaze’ were used to determine the effects of sucrose and inorganic nitrogen on adventitious root formation. Shoots grown in media containing high sucrose concentrations (146.07–262.93 mM) produced more and longer roots than those grown in media containing 0–87.64 mM sucrose. This response to sucrose was related to the metabolism of sucrose rather than its osmotic properties since the use of mannitol and 3-O-methyl-α-D-glucopyranoside as osmotic substitutes did not reproduce the effect on rooting. The number and length of roots increased when the shoots were grown in media with the nitrogen concentration of the Murashige-Skoog (MS) salt formulation reduced from 60 to 7.5 mM. Neither nitrate (NO ₃ ⁻ ) nor ammonium (NH ₄ ⁺ ) alone at any of the concentrations tested had the effect on rooting that both had together in the ratio of the MS salt formulation. When the sucrose and nitrogen concentrations were both varied, the greatest rate of root initiation occurred on shoots grown in media with a high sucrose to nitrogen concentration ratio.     

Influence of potassium and manganese on growth and uptake of magnesium by soybeans (Glycine max (L.) Merr. cv. Bragg)

Summary Soybean (Glycine max (L) Merr. cv. Bragg) seedlings were grown in nutrient solutions to evaluate the response to manganese nutrition as affected by potassium supply. In solutions containing 275 M manganese, increasing the solution concentration of potassium from 1 mM to 10 mM alleviated symptoms of manganese toxicity, decreased manganese concentrations in the leaves and increased dry matter yields of the plants. The reduction in manganese toxicity was brought about by a reduced rate of root absorption of manganese at high potassium supply levels.Increasing the supply of either potassium or manganese decreased the leaf concentration of magnesium although there were no apparent symptoms of magnesium deficiency in any treatment. The reduced concentration of magnesium in the leaves was due to effects of potassium and manganese on the rate of root absorption of magnesium.Under manganese deficiency conditions, growth was reduced and manganese concentrations in plant parts were very low; there was no effect of potassium supply when manganese was absent from the nutrient solution.