Plant and Soil 54, 339-357 (1980)

Ms. 4115

pH OPTIMA FOR CROP GROWTH

RESULTS OF A FLOWING SOLUTION CULTURE EXPERIMENT W I T H SIX S P E C I E S by A. K. M. S. I S L A M , D. G. E D W A R D S a n d C. J. A S H E R

Department of Agriculture, University of Queensland, St. Lucia, Queensland, 4067, Australia

KEY WORDS Calcium Cassava Chemical composition Control of solutionpH Copper Flowing solution culture French bean Ginger Hydrogen ion injury Magnesium Maize Manganese Nitrogen Optimum pH range pH Plant growth Root weight ratio Tomato Wheat SUMMARY Ginger, cassava, maize, wheat, french bean and tomato were grown for periods up to six weeks in continuously flowing nutrient solutions at seven constant pH values ranging from 3.3 to 8.5. All species achievedmaximum or near-maximum growth in the pH range 5.5 to 6.5. However, there were substantial differencesin the ability of speciesto grow outside this range. Ginger and cassava were the most tolerant species to low solution pH, while ginger and tomato were the only species to show no yield depression at the highest solution pH. Roots of all species at pH 3.3 and some speciesat pH 4.0 exhibited symptoms of hydrogen ion injury. In addition, the concentrations of magnesiumin the tops of all six species,of nitrogen in the tops of tomato and cassava, and of manganese in the tops of maize at these pH values were inadequate for optimal growth. Growth depression at high solution pH was associated with iron deficiencyin maize and wheat and with nitrogen and/or copper deficiencyin cassava. The relevanceof the present results to crop growth under fieldconditions is discussed.The complex interplay of plant and soil characteristics militates against precisedefinition of an optimum pH range for the growth of a particular crop unless the soil is also specified. INTRODUCTION Soil p H is an i m p o r t a n t factor affecting the growth of crops a n d pastures, a n d the d i s t r i b u t i o n of native p l a n t species 15,21,34, 52, 54, 58, 64. Often the effects of p H are complex a n d it is difficult to separate direct effects of excess h y d r o g e n or hydroxyl ions from indirect effects associated with changes in the solubility of various biologically i m p o r t a n t m i n e r a l elements. S o l u t i o n culture experiments offer a m e a n s of studying direct effects of p H o n p l a n t growth in the absence of large changes in other root e n v i r o n m e n t parameters. However precise c o n t r o l of s o l u t i o n p H a n d n u t r i e n t ion c o n c e n t r a t i o n s is difficult to achieve using c o n v e n t i o n a l s o l u t i o n culture methods. The m a g n i 339

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A. K. M. S. ISLAM, D. G . E D W A R D S A N D C. J. A S H E R

t u d e o f this p r o b l e m is i l l u s t r a t e d by t h e w e l l - k n o w n s t u d y of A r n o n a n d J o h n s o n a w h o h a d t o e m p l o y l a r g e v o l u m e s of well stirred n u t r i e n t s o l u t i o n (10 to 201 p l a n t - 1), p l u s c o n t i n u o u s s l o w a d d i t i o n of a c i d o r alkali to e a c h c u l t u r e vessel a n d t w i c e d a i l y c h e c k s of s o l u t i o n p H to c o n t r o l p H to w i t h i n _ 0.2 units. M o d e r n f l o w i n g c u l t u r e t e c h n i q u e s 5, 7,13 are ideally s u i t e d to m e e t i n g the t w i n p r o b l e m s of p r e c i s e p H c o n t r o l a n d t h e m a i n t e n a n c e of a d e q u a t e n u t r i e n t c o n c e n t r a t i o n s at all p H levels, b u t d o n o t a p p e a r to h a v e b e e n u s e d for this p u r p o s e u p till n o w * . In the p r e s e n t study, g i n g e r (Zingiber officinale R o s c o e ) a n d five o t h e r species w e r e g r o w n in c o n t i n u o u s l y f l o w i n g n u t r i e n t s o l u t i o n s m a i n t a i n e d at c o n s t a n t p H v a l u e s r a n g i n g f r o m 3.3 to 8.5. T h e s t u d y was c o n c e r n e d p r i m a r i l y w i t h e s t a b l i s h i n g the o p t i m u m p H r a n g e for ginger, b u t o t h e r species w e r e i n c l u d e d to facilitate c o m p a r i s o n s w i t h e a r l i e r studies a n d to p r o v i d e , a m o r e a d e q u a t e basis for g e n e r a l i z a t i o n s c o n c e r n i n g effects of p H o n p l a n t g r o w t h .

MATERIALS AND METHODS

Propagation, selection o f species

Ginger rhizome pieces weighing 10-15 g were germinated in peat moss following successive pretreatments with water at 51~ for 10 minutes, benlate, and 1000 ppm ethrel as described elsewhere 26. Other species included were cassava (Manihot esculenta Crantz, cv. Nina), tomato (Lycopersicon esculentum cv. Grosse Lisse), french bean (Phaseolus vulgaris cv. Redland Pioneer), wheat (Triticum aestivum cv. Gatcher), and maize (Zea mays cv. NK 195). With the exception of tomato, the mineral nutrition of all of these species had been the subject of other flowing culture studies within the Department of Agriculture at the University of Queensland. Tomato was included to allow a comparison of,the results with those obtained for that species by Arnon and Johnson 3. Cassava plants were raised from stem tip cuttings in a mist propagation chamber as described by Forno et al? ~ The remaining four species were raised from seed. In each case the seed was surface sterilized for 1 minute in 0.1% HgC1z solution, rinsed thoroughly in deionized water, then soaked in aerated deionized water for either two hours (tomato, french bean) or 12 hours (wheat, maize). The partially imbibed seeds were then germinated between moist paper towels in an incubator at 25~ The seeds were transferred to the nutrient solutions when the radicles were 5 to 10mm long. Propagation of individual species was commenced at differing times so that all species would be ready for transplanting at approximately the same time.

Flowing culture equipment

The flowing culture equipment used in the present study is of the continuously recirculating type similar in principle to that described by Asher et al.7. A brief description of the equipment was given by Edwards and Asher 13 and a more detailed description by Asher and Edwards s.

* N~teaddedinpr~j~.Sincethispaperwasaccepted~rpub~icati~nthef~wingreferencehasc~met~theauth~rs~
attention: Moritsugu, M. and Kawasaki, T. 1979 A new system of automatic pH regulation in solution culture. Bet. Ohara Inst. landw. Biol. 17, 171-178.

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Details of treatments, experimental design

The composition of the basal nutrient solution was as follows (aM): Ca 250, K 250, N (as nitrate ) 250, S 260-560, P 15, Mg 10, Na 40-63, Fe (as sequestrene 138 ~20, Si 10. CI 5, B 3, Zn 0.5, Mn 0.25, Cu 0.1, Co 0.04, and Mo 0.02. The temperature of the nutrient solutions was maintained at 25 _ _ +1 ~ For the first five days after transplanting the nutrient solution in all flowing culture units was maintained at pH 6.0 + 0.1 and the plants were lightly shaded to aid establishment. The pH was then adjusted with 0.25 M H2SO 4 or 0.5 M NaOH to values of 3.3, 4.0, 4.8, 5.5, 6.5, 7.5 or 8.5. Thereafter the pH was held approximately constant by the automatic pH control equipment incorporated in each flowing culture unit. A Rikadenki multi-channel chart recorder (Rikadenki Kogyo Co. Ltd. Model PBR-I 2) recorded the pH in each unit at one minute intervals as a check on the short term stability of pH control. Long term stability was assessed on the basis of twice daily pH meter readings over the six week experimental period. Within each of the seven flowing culture units there were 18 replicate cultures of ginger, and eight replicates of each of the other species. The possible role of iron deficiency as a factor limiting growth at high solution pH values was assessed by spraying half the replicates of all species at each pH with 0.5 per cent (w/v) ferrous sulphate solution three times per week. To ensure an adequate supply of all essential elements the nutrient solutions were completely renewed at weekly intervals. In addition, solution nitrogen and phosphorus concentrations were checked every 3 to 4 days and adjusted as necessary. Nitrate concentrations were measured with an Orion Model 404 Specific Ion Meter and solution phosphorus analyses were conducted using the method of Taras et al. 63.

Harvestin9, chemical analysis

Immediately prior to imposing the pH treatments, a thinning harvest was conducted leaving one plant per pot of all species except wheat which was thinned to four plants per pot. Initial weights of roots and tops of each species at the time of imposing the pH treatments were obtained from this thinning harvest. The differing absolute growth rates among the species and the need to avoid shading by the faster growing species resulted in the harvest of plants after treatment periods ranging from two weeks (tomato, french bean) to six weeks (ginger). The remaining three species were harvested after four weeks. At each harvest, the roots were separated from the tops, rinsed free of nutrient solution and blotted dry. Fresh and dry weights of tops and roots were then obtained, a 70~ forced draft oven being used to dry all plant samples. Selected samples of plant tops which had not been sprayed with ferrous sulphate were ground in a cast iron laboratory hammer mill and subjected to multi-element analysis by direct reading emission spectroscopy using a briquetting technique as described by Johnson and Simons 3~.

RESULTS

(a) Control of solution pH

B o t h s h o r t t e r m a n d l o n g t e r m s t a b i l i t y o f p H in t h e s e v e n t r e a t m e n t s w e r e f o u n d t o b e h i g h l y s a t i s f a c t o r y . A s t u d y o f c h a r t r e c o r d i n g s s h o w s t h a t v e r y few o f t h e m o r e t h a n 1400 o b s e r v a t i o n s p e r u n i t d u r i n g a 24 h o u r p e r i o d lay m o r e t h a n 0.1

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A. K . M. S. ISLAM, D. G . E D W A R D S A N D C. J. A S H E R

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p H units above or below the mean (Fig. 1). The mean pH values and their 95 per cent confidence limits calculated from the twice daily pH measurements over the six week experimental period were as follows: 3.3 + 0.01, 4.0 + 0.01, 4.8 + 0.01, 5.5 + 0.01, 6.5 _ 0.02, 7.5 + 0.03, and 8.5 +__0.02.

(b) Effect of solution pH on plant growth

Solution pH had large effects on growth of all six species (Fig. 2). Ginger was the most tolerant species to extreme solution acidity achieving a mean relative yield of 44.5 per cent at pH 3.3. Cassava was the next most tolerant species at this pH, followed by tomato, french bean, wheat and maize. The latter two species achieved only 12 and 8 per cent of m a x i m u m yield respectively at this pH. At pH 3.3 and also 4.0 tomato showed symptoms similar to mild nitrogen deficiency 61, while maize and wheat showed symptoms suggestive of magnesium deficiency 61. Most species reached or closely approached maximum yield at pH 5.5 and no significant positive growth responses to increasing pH were observed above this p H value. Increasing the p H from 5.5 to 8.5 had no significant effect on the growth of ginger or tomato but depressed the growth of all other species. In maize and wheat the growth reduction in the unsprayed series was accompanied by the development of iron deficiency chlorosis 61. Spraying with ferrous sulphate significantly reduced the growth depression in maize and reduced the intensity of iron chlorosis in both species. In wheat, however, the iron spraying also caused some leaf injury. The growth of wheat was significantly reduced by the spray

P H OPTIMA FOR GROWTH OF SIX CROP SPECIES

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A. K. M. S. ISLAM, D. G. EDWARDS AND C. J. ASHER

treatment at pH 4.8 and 5.5 and the effectiveness of spraying in reducing growth depression at higher pH values was substantially less than in maize (Fig. 2). Spraying with ferrous sulphate also caused some leaf injury and growth depression in french bean (Fig. 2). In cassava, a substantial growth depression occurred at pH 8.5 in both sprayed and unsprayed plants but no symptoms of iron deficiency or of any other micronutrient deficiency were observed. Spraying with ferrous sulphate did not improve growth of cassava in the higher pH treatments.

(c) Root injury at low pH

Damaging effects of low solution pH on root growth were observed inall species at pH 3.3, and in some species at pH 4.0. At pH 3.3, roots were short, thickened, comparatively few in number, and discoloured brown or dull grey. Lateral root growth was severely inhibited in all species. At pH 4.0, root growth appeared healthy in ginger, cassava and maize. By contrast, at pH 4.0, roots of french bean and wheat exhibited necrotic root tips and short laterals. The roots of french bean were discoloured grey while those of wheat and tomato were brown. In tomato, at this pH, healthy root growth occurred only in the moist air gap above the nutrient solution. The relatively coarse, discoloured roots present in all species at low pH contrasted markedly with the fine, light coloured, profusely branched and symptom-free roots present at pH 4.8 and above.

(d) Effects of solution pH on root weight ratio

Effects of the iron spraying treatment on root weight ratio were not significant at P = 0.05 in 39 of the 42 species pH treatment combinations studied. Hence, it was considered legitimate to pool that data for sprayed and unsprayed plants. There were large effects of solution pH on the root weight ratio of all species. In each case, root and top gro~vth was poor at pH 3.3. However, as the solution pH was raised from 3.3 to 4.0, the root weight ratio increased sharply in cassava, tomato and ginger as a result of the greater response in root growth than shoot growth to the increase in pH (Fig. 3). By contrast, the root weight ratio remained unchanged in french bean and fell rapidly in wheat and maize as the solution pH was increased from 3.3 to 4.0. This reduction in root weight ratio was entirely due to increased top growth at pH 4.0. Raising the pH from 4.0 to 4.8 increased the root weight ratio of all species except ginger, again indicating a greater response in root growth than top growth to increase in solution pH. The root weight ratios of cassava, tomato and ginger decreased while those of maize and wheat in-

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Fig. 3. Effect of solution pH on root weight ratio of six crop species grown in flowing culture. (Means of 18 replications for ginger and eight replications for other species; means followed by the same letter are not statistically different at P = 0.05). The root weight ratios of plants prior to imposition of treatments were: ginger not determined, cassava 0.20, tomato 0.14, french bean 0.11, wheat 0.37, maize 0.50.

creased significantly as the solution p H was raised from 4.8 to 5.5. T h e r o o t weight r a t i o of maize c o n t i n u e d to increase, b u t that of the o t h e r species showed little c h a n g e over the p H r a n g e 5.5 to 7.5. The r o o t weight r a t i o of all species increased significantly with further increase in p H to 8.5. A l t h o u g h effects of p H on r o o t weight r a t i o differed s u b s t a n t i a l l y a m o n g species, n o clear difference was evident between m o n o c o t y l e d o n s as a g r o u p a n d d i c o t y l e d o n s (Fig. 3). In n u t r i t i o n a l studies, lower r o o t weight ratios generally tend to occur in those t r e a t m e n t s m o s t f a v o u r a b l e to p l a n t g r o w t h 1'3'59. H o w e v e r , in the present e x p e r i m e n t no simple r e l a t i o n s h i p between yield level a n d r o o t weight r a t i o

346

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P H OPTIMA FOR GROWTH OF SIX CROP SPECIES

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emerged. Further research is needed to explain these responses. Differences in root weight ratio prior to imposition of the p H treatments (Fig. 3) and subsequent effects of these treatments on both root and top growth may be involved.

(e) Chemical composition of plant tops

Analysis of whole tops of unsprayed plants from selected treatments showed that pH had very large effects on chemical composition (Table 1). Increasing solution p H from 3.3 to 5.5 strongly increased the concentrations of potassium, calcium and magnesium in the tops of all species. A further increase in solution p H to 8.5 resulted in increased magnesium concentrations in all species, increased calcium concentrations in french bean and maize, and an increase in potassium concentration in tomato. Nitrogen concentrations in the tops of all species except ginger increased as the solution p H was raised from 3.3 to 5.5. The nitrogen concentration in cassava and wheat was strongly decreased and that in tomato increased as the solution pH was further increased to 8.5. Phosphorus concentrations generally increased with increase in solution p H from 3.3 to 5.5, then declined as the solution p H was increased to 8.5. Sulphur concentrations were less in the tops of all species except cassava at pH 4.0 than at pH 3.3, and either increased or showed little change as the pH was raised to 5.5 and 8.5. Manganese and zinc concentrations in the tops of all species increased with increasing solution p H from 3.3 to 8.5. Copper concentrations decreased in ginger and maize and increased in wheat as the solution pH was increased. Iron concentrations were maximal in wheat at pH 3.3 and in all other species at p H 4.0 and were strongly depressed by further increase in solution pH in all species except tomato. Boron concentrations were little affected by solution p H from 3.3 to 8.5, while molybdenum concentrations increased as the solution p H was raised from 3.3 to 5.5, and then declined in most species as the pH was further raised to 8.5.
DISCUSSION

(a) Apparent sensitivity of plants to variation in pH

On the basis of carefully conducted solution culture experiments with tomato, lettuce and Bermuda grass, Arnon and Johnson 3 concluded: 'Within a relatively wide range of pH between 4 and 8, fluctuations in the hydrogen ion concentration are tolerated by plants provided an adequate supply of all nutrient elements is maintained.'

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A. K . M. S. ISLAM, D. G . E D W A R D S A N D C. J. A S H E R

Their paper, which has been widely quoted in the literature, often has been interpreted to mean that any major effects of pH on plant growth between pH 4 and 8 are likely to be indirect effects associated with pH dependent changes in the solubility of micronutrients or other biologically important elements in the soil. Some support for the conclusions of Arnon and Johnson 3 comes from studies on the nutrition of non-nodulated legumes in which little or no growth depression occurred in solution cultures or sand cultures at pH values as low as 4.5 (and sometimes 4.0) provided the plants were adequately supplied with calcium, mineral nitrogen and other essential elements ~'39"47. However, a careful examination of the data of Arnon and Johnson (Ref. 3, Table 2) shows that at pH 4 only Bermuda grass grew well; the yields of tomato and lettuce were reduced to less than 35 per cent of maximum. Again at pH 8, the growth of the three species studied was reduced to between 58 per cent (tomato) and 69 per cent (Bermuda grass) of maximum. In the present study also, yields of all six species were markedly reduced at pH 4, ranging from 14 per cent of maximum for maize to 62 per cent for cassava (mean of sprayed and unsprayed treatments). Similarly, substantial yield reductions occurred in the region o f p H 8 in cassava, wheat and maize (Fig. 2). Since in both studies a conscious effort was made to maintain 'an adequate supply of all nutrient elements' we must conclude that either plants are rather more sensitive to extremes o f p H than has been previously supposed, or that the problems of maintaining an adequate nutrient supply at low or high pH values have not been fully overcome.
(b) Injury by H + or O H - ions

Short term studies have shown that at low pH, ion transport may be impaired, especially at low calcium concentrations 2~, and that sufficient membrane damage may occur to allow the loss of previously absorbed solutes 29"45. Longer term studies on several plant species 3,2,~.57.62 have shown that prolonged exposure of roots to low pH leads to suppression of lateral root development and, in extreme cases, to death of the root tips. In the present study, all species showed these symptoms at pH 3.3, and french bean, wheat and tomato showed them at pH 4.0. Less is known about the effects of O H - ions than of H + ions on plant growth, but directly injurious effects within the range of pH values normally encountered in soils seem unlikely. Thus even at pH 8.5, the highest pH studied in the present experiment, the O H - ion concentration would have been only 3.2/aM. This value may be compared with H + ion concentrations of 100/aM (pH 4) to 500/aM (pH 3.3) associated with visible root injury in the present study.

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(c) Ef[ects of pH on ion uptake

Several short term studies have shown large effects o f p H on the rate of uptake of various cations. Thus, Rashid et al. 51 showed that reducing the pH of the test solution from 5.5 to 4.5 decreased zinc absorption rates by factors of 1000 and 10000 in the rice cultivars IR-6 and Basmati-370 respectively. In another study, decreasing the solution pH from 5.0 to 3.0 reduced zinc absorption by a factor of about 100 in wheat seedlings 1~ Similarly, large decreases in the rate of absorption of manganese 9'43' 53, calcium42, magnesium 4, 37,4.4,46, potassium28 and copper 9 with decreasing pH have been reported. In the present study, tissue concentrations of all essential elements for which we analysed were adequate for healthy growth at pH 5.5 (Table 1) 6 ' 1 5 ' 3 2 ' 4 0 However, the magnesium concentrations in the tops of all six species at pH 3.3 and 4.0 (0.03-0.16~) were sufficiently low to be either deficient or marginally limiting for plant growth 6,19, 2o, 30, 33, 60, 65, 66. Manganese concentrations in the tops of maize at pH 3.3 and 4.0 (12 and 14 pg.g- ~ respectively) were also in the range considered to be inadequate for healthy growth 32, 36. In all species except ginger, nitrogen concentrations in the tops decreased with decreasing pH over the range 5.5 to 3.3 (Table 1) and in tomato the concentrations at pH 3.3 and 4.0 (1.2 and 1.3~ respectively) were clearly in the deficient range. Nitrogen concentrations in cassava tops at pH 3.3 and 4.0 (2.3 and 2.6~ respectively) were also well below the critical nitrogen concentration of 5.1 ~o in the fourth and fifth fully expanded leaves of cv. Llanera 18 and the normal nitrogen concentrations ranging from 4.5 to 6.5 ~o in young fully-opened leaves ~1. No satisfactory explanation can be given for the decline in plant nitrogen concentrations at low pH. Bassioni 8 reported that nitrate uptake by excised barley roots was less at pH 4 than at pH 6. However, this result is somewhat suspect as the test solutions apparently did not contain calcium. In more recent work, Rao and Rains 50 reported higher rates of nitrate absorption in short-term uptake experiments with barley roots at pH 4.0 than at pH 5.7 or 8.5. Similarly, in a flowing solution culture experiment, Forno 16 found that mean rates of nitrogen uptake (as nitrate) per unit root weight were either higher at pH 4.4 than at pH 6.8 (cassava cv. M Aus 3, cotton) or approximately the same (cassava cvs Nina, Ceiba). Calcium concentrations in maize at pH 3.3 and 4.0 (0.39 and 0.37~ respectively) were below the concentration normally considered adequate for healthy growth (0.5~o)15; however, Loneragan and Snowball 4~ obtained maximum yield of young maize plants in flowing solution culture when the calcium concentration in the tops was only 0.12~. In the same experiment, toma~to and the wheat cultivars Gabo and Wongoondy achieved maximum yield with cal-

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A. K. M. S. ISLAM, D. G . E D W A R D S A N D C. J. A S H E R

cium concentrations in the plant tops of 1.29 ~o, 0.15 ~o and 0.32 ~o respectively 40. These calcium concentrations are well below those obtained in the tops of tomato and wheat cv. Gatcher in the present experiment at pH 3.3 and 4.0. It would thus appear that calcium was not growth-limiting for maize, tomato, wheat or probably for any other species at any solution pH in the present experiment. Increasing the solution pH from 5.5 to 8.5 caused substantial reductions in nitrogen concentrations in shoots of cassava and wheat, and smaller reductions in maize, french bean and ginger (Table 1). Decreases in rate of phosphate absorption with increasing pH are well documented 22'23 and Arnon and Johnson 3 considered that phosphorus deficiency contributed to the poor growth of their plants at the higher pH values. However, in the present experiment, tissue phosphorus concentrations were in all cases adequate at pH 8.5. Poor iron nutrition depressed the growth of maize and wheat at pH 7.5 and 8.5, despite the use of F e E D D H A (sequestrene t38) as an iron source. This compound is reported to be stable over the pH range 4 to 9 48 but was not entirely satisfactory under the conditions of our experiment, particularly at the higher pH values. In this connection, Lindsay and Halvorson 38 have reported that in nutrient solutions absorption of iron by the plant roots leaves free chelating agent in solution which then competes with the plant roots for the ferric ions remaining in solution. Iron concentrations in the tops of tomato, ginger and french bean which showed little yield depression at pH 8.5 were higher than those in cassava, maize and wheat in which yields were severely depressed (Fig. 2, Table 1). In fact, the iron concentration in the tops of maize grown at pH 8.5 (85/tg.g- 1) is in the range that has been considered deficient for this species 3s, an observation confirmed by the development of severe symptoms of iron chlorosis (cf. Sprague61). It would appear that tomato, ginger and french bean were more efficient than the other three species in obtaining their iron requirements from the nutrient solution at pH 8.5. Tomato was common to both the present study and that of Arnon and Johnson 3 and hence provides the most satisfactory basis for comparison of the results obtained in the two studies. Above pH 7 growth in the present study was superior to that in Arnon and Johnson's study, presumably because of the better control of phosphorus nutrition achieved under flowing culture conditions. However, over the pH range 4 to 6.5 better growth was obtained in Arnon and Johnson's study (Fig. 4), partly because of inadequate nitrogen and magnesium uptake at lower pH values in the present study. Arnon and Johnson attributed much of their growth reduction below pH 5 to inadequate calcium absorption. In a subsidiary experiment with lettuce and tomato they showed that raising the initial calcium concentration in the nutrient solution from 2000 to 7000/zM

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increased yields at both pH 4 and 5, whereas lowering the initial calcium concentrations to 500~M lowered plant yields. Further evidence of the interaction between effects of low pH and calcium concentration comes from studies in legume nutrition. Thus, Munns 47 showed that lucerne plants supplied with combined nitrogen grew equally well at pH 4 and 5 with a calcium concentration of 5000~M, but that growth was poorer at pH 4 when lower calcium concentrations were used. Calcium deficiency symptoms were not observed in the lucerne tops at pH 4 when plants were grown at the lowest solution calcium concentration used (1000/tM). Andrew i showed that the growth of six temperate pasture legumes plus the tropical legumes Glycine wightii and Desmodium uncinatum supplied with inorganic nitrogen was more severely restricted at pH 4 when calcium was supplied at 125pM than at 2000pM. At this pH, D. uncinatum, Macroptiliurn lathyroides, lucerne and G. wightii displayed calcium deficiency symptoms with calcium concentrations in the plant tops of 0.13, 0.15, 0.16, and 0.21 ~o respectively 2. The other species which showed yield reduction s at low calcium ( 125 pM) at pH 4 had somewhat higher concentrations in the plant tops and did not exhibit any calcium deficiency symptoms. Loneragan et al. 41 determined the solution calcium concentrations necessary for maximal growth of 30 grasses, cereals, herbs and legumes in flowing culture solutions at pH 5.7. Wheat cv. Gabo and maize achieved maximal growth at a solution calcium concentration of lOpM, while wheat cv. Wongoondy and tomato achieved maximal growth at 100pM calcium. In the present study, a constant calcium concentration of 250/~M was maintained throughout the

loo

2c

pH OF NU"/RIENT SOLUTION

Fig. 4. Effects of solution pH on growth of tomato: O-----O Data from Arnon and Johnson 3. O - - - - O Data from current experiment.

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A. K . M. S. ISLAM, D. G . E D W A R D S A N D C. J. A S H E R

experiment at all solution p H values. The failure to observe any calcium deficiency in the plant tops and the adequate calcium concentration in the plant tops, even at p H 3.3 and 4.0, indicate there was no direct calcium limitation on plant growth. However, the possibility exists that better growth of at least some species may have occurred at low p H in the present experiment if a higher solution calcium concentration had been selected. Protection of plant tissues against hydrogen ion injury by an elevated calcium concentration 28,45 provides the most likely explanation for such an effect. The above observations are consistent with the view that if sufficient attention is given to the formulation of nutrient solutions at low and high pH values, vigorous plant growth may be obtained over a rather broad range in pH. However, as far as the authors are aware, no one has yet succeeded in demonstrating an optimum range as wide as the 4 pH units proposed by Arnon and Johnson 3.
(d) Relevance to plant growth in the field

Large differences were observed in the ability of the six crop species used in the present study to grow at low pH values. Thus at pH 3.3 and 4.0 species ranked in the following order of relative yield: ginger > cassava > tomato > french bean > wheat > maize (Fig. 2). These results are in good agreement with those obtained in a recent field study in which the ability of species to grow in a highly acid oxisol with 0 or 0.5 tonnes lime per hectare ranked in the order cassava > beans > maize (CIAT) 12. The behaviour of ginger also accords well with field experience. In recent field studies, Islam et al. 27 obtained rhizome yields /> 9 0 ~ of maximum at comparatively low pH values on a majority of soils. Thus, of eight soils examined, optimum yields were obtained on four at soil pH values (measured in 1 : 2 soil/0.01 M CaCla) ranging from 4.3 to 4.6, and in a further three at pH values ranging from 5.0 to 5.2. The p H required for 90~o of m a x i m u m yield in the remaining soil was 6.4. The wide range of optimum soil pH values observed by Islam et al. 27 for the same crop on different soils (pH 4.3 to 6.4) calls into question the concept of an optimum pH for the growth of a particular crop. In the field, response to varying pH will be influenced not only by susceptibility of the crop to hydrogen ion injury at low p H and to various deficiencies at low and high p H discussed in (c) above, but also by large pH dependent changes in the solubility of aluminium, phosphorus, manganese, iron, zinc, copper and molybdenum. Susceptibility to adverse effects of high concentrations of soluble aluminium or manganese present in some soils at low p H is also likely to be a major factor determining the optimum

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353

pH. In some soils, effects of pH on the ratio of a m m o n i u m to nitrate-nitrogen may be important also. Hence a combination of soil and crop characteristics will determine the optimum p H at any particular site. A further difficulty in characterizing the optimum p H for growth of a particular crop concerns the measurement of soil pH. Different methods of measurement often yield markedly different pH values for the same soil. However, a consensus appears to be emerging that the measurement of soil p H in a dilute electrolyte such as 0.01 M CaC12 solution provides the best index of the concentration of hydrogen ions present in the soil49' 55, 56, 67. Soil pH values measured in 0.01 M CaCI 2 are often 0.5 to 1.0 p H units lower than values measured at the same soil/solution ratio in water 67. Unfortunately, a wide variety of p H measurements are currently in use and it often happens that the method of measurement is not specified precisely when crop response data are reported. Recent research on the liming of cassava soils may serve to illustrate the problems in arriving at an optimum soil pH for the growth of a particular crop. Spain et al.58 found that most of the 138 cassava cultivars grown on a highly acid oxisol in Colombia reached m a x i m u m yields at a low rate of liming (0.5 t.ha-1) corresponding with a p H (1 : 1 soil/water suspension) of 4.4. Raising the pH further to 4.7 or 5.3 caused moderate to severe yield depression through the induction of zinc deficiency. Edwards and Kang14 obtained somewhat similar results on the lime requirements of two cassava cultivars on a highly acid ultisol in Nigeria. In this case, the optimum pH (1 : 1 soil/water suspension) was in the range 4.6 to 5.1. Higher liming rates reduced yields drastically due to zinc deficiency, with tuber yields reduced to zero at an application rate (5 t.ha-1) corresponding with a soil pH of approximately 7.0. The range of p H values corresponding with m a x i m u m yield in these two studies (4.4 to 5.1) falls outside the optimum range found for cassava of 5.5 to 6.5 in the present study (Fig. 2). Since the measured p H values of the Colombian oxisol and the Nigerian ultisol were both above the zero point of charge, the discrepancy between the optimum pH ranges would have been even wider had the soil pH's been measured in a dilute electrolyte instead of in water. More recent work on the Colombian oxisol has shown that when adequate amounts of zinc and other micronutrients were included in the fertilizer mixture, cassava showed a marked positive growth response from 0.5 to 2 tonnes lime per hectare and a smaller positive growth response up to 6 tonnes lime per hectare at which rate the soil p H had increased to 5.325. No information is available on the effect of supplying zinc on the p H optimum in the Nigerian soil studied by Edwards and Kang 14. The foregoing considerations lead us to the conclusion that while it appears possible to rank crops in terms of their sensitivity to extremes of soil pH, it is

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A. K. M. S. ISLAM, D. G. EDWARDS AND C. J. ASHER

e x t r e m e l y difficult t o d e f i n e p r e c i s e l y t h e o p t i m u m p H r a n g e f o r t h e g r o w t h of a p a r t i c u l a r c r o p . E v e n in c a r e f u l l y c o n d u c t e d s o l u t i o n c u l t u r e e x p e r i m e n t s it is difficult t o e n s u r e o p t i m u m levels o f s u p p l y o f all e s s e n t i a l e l e m e n t s a t all p H on nutrient solubility further complicate the v a l u e s . I n soils, effects o f p H

s i t u a t i o n . H e n c e p u b l i s h e d v a l u e s for t h e o p t i m u m p H o f a c r o p m a y h a v e r a t h e r l i m i t e d a p p l i c a b i l i t y o u t s i d e t h e p a r t i c u l a r set o f c o n d i t i o n s u n d e r w h i c h t h e y were determined.

ACKNOWLEDGEMENTS Grateful acknowledgement is made to the Reserve Bank of Australia for support of A.K.M.S. Islam in the form of a Rural Credits Development Fund research grant. Thanks are also due to Buderim Ginger Growers' Cooperative, Buderim, Queensland for the supply of planting material, to Mr. G. Kerven, Mr. H. Mighell and Mrs. J. Mercer for technical assistance, and to Mr. B. Kenyon for assistance with the preparation of the manuscript. Special thanks are due to Mr. A. D. Johnson, CSIRO Cunningham Laboratory, St. Lucia, Queensland for running multi-element analyses of plant samples from selected pH treatments. Received 12 June 1979. Revised August 1979

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