Pertanika 15(3),189,197 (1992)

Effect of Toxic Concentrations of H+ and AI onNonsymbiotic Growth of Groundnut in Solution Culture

ROSMIN KASRAN ", ZULKIFLI H. SHAMSUDDIN * and D.C. EDWARDS"* Depaft11lent of Soil Science, Universiti Pertanian Malaysia.

43400 UPM Serdang, Selangor, Malaysia.*.~, Depa1tment oj Ab'T'icultU1'e, The University of Queensland,

Queensland, Australia 4072.

Keywords: groundnut, H· toxicity, aluminium toxicity, solution culture

ABSTRAK

Dua percubaan kultura nutrien telah dijalankan untuk mengkaji kesan kepekatan toksik H' dan aluminiumterhadap pertumbuhan bukan-simbiotik kacang tallah (Arachis h)'pogaea L) cu. Matjam. Lima paras pH yangdigull,aka,n ialah 3.5, 4.0, 4.8, 5.5 dan 6.5 manakala jumlah ak/jvil; spesies Al monomerik(.~aA"",,) ialah 0, 1.5, 3.0, 15, 35 dan 80 I'-M pada paras pH 4.3. &pekarall iOIl H' pada pH 3,5 tidakmenjejaskan berat kering bahagian alas pokok [elapi menjejaskan pertumbuhan akar dan pemanjangannya.Peningkatan pH kepada 4.0 11lenambahkan lagi pertumbulwn akttT. Paras pH yang disyorkan untuk tumbesarankacang tanah iatah ~ 4.4. Kepekatan aluminium yang tidak toksik untuk peltumlmhan ballagian atas danpemanJangan akar ialah ~ 12.2 J..LM 4a.~,t1tOI'" ( ==: 17.11J.M Al-monomeric alau 20.3 J..LI\t( mon01llerik Alju71lIah).Peninglwlan kepekatan La

MmVMkepada 80 p,M merendahkan kepekatan magnesium pada daun lermuda

berkembang penuh (YEL) kepada paras tidak mencukupi, 0.18%. Kepelwran magnesium di dalam YEL lebihsensitiJ terhadap ketoksikan aluminium daripada kepekalan magnesium di da/am akar. Keputusan jugamenrlapati kacang tanak member; res/Jon berat kering a!wT dan pemanjangan akar yang positiI pada hepeka,tanaluminium yang rendah (1.5 IJ.M ~a.~f""'"); kesan ini lidok ditunjukkan pada beral kering bahagian alas IJOkol~.

ABSTRACT

Two solution culture experiments We1'e conducted to stt,d)' the effects of toxic concentrations ofH and aluminiumon. nOllsymbiotic growth ojgroundnut (Arachis hypogaea L.) cu. Maljam. TheJive pH lroels used were 3,5, 4,0,4.8.5.5 and 6.5, and the sum of activities o/monomeric Al species (LaM_"/I) were 0. 1.5. 3.0, 15,35 and 80I'-M at pH 4.3. The H' iOIl concentratian at pH 3.5 did not aiJed top dry weight but markedly decreased root d,)'weight and kngth. Raisillg the solution pH 10 4.0 illlp1VlJed the grawth oj roots. The pH level recom-mended Jargrowth of groundnut was ~ 4.4. The non-toxic concentration of aluminium for growth of groundnut and motelongation was ~ 12.2 p,M La,u_",,(==: 17.1 J..L1W monomeric-Al or 20.3 J-LM monomeric total-Ai). Increasing the};a-\/Jfl""/I to 80 J..LM decreased the magnesium concentration in the youngest expanded leaf (YeL) to 0.18%. a valueknown to be deficient for groundnut. J\1agnesium concentmtion in the YEL was more sensitive to aluminium. toxicitythan magnesium concentration in the roots. Tlu! results also showed that root dry weight and root length ·respondedpositively to low concentrations oj aluminium (1.5 1·L1W ~aAI,""'i"); this effect was not observed for dry' weight

oj tops.

INTRODUCTION

Many soils of the humid u'opics have pH valuesof less than 5.0 (1:5 soil: waler) and low levels ofexchangeable calcium (Pearson 1975), Low pHper se seems unlikely to inhibit growth of legumesseverely, even in acid-sensitive species, such asMedicago sativa and Phaseolus vulgaris. These plants

have been shown to tolerate acid conditions (pH4.5) in solution culture with a low concentrationof aluminium (~ 10 J.lM). provided there was anadequate supply of calcium and combined nitro­gen (Asher and Edwards 1978; Franco and Munns1982).

+ Present address:Cocoa and CoconUl Research Division, MARDI Hilir Perak, P.O. Box 25, 36307, Sungai Sumun, Perak. D.R., Malaysia

ROSMIN KASR~N, ZULKIFLI H. SHAMSUDDIN AND D.G. EDWARDS

Aluminium LOxicity is a major problem forthe growth of many plant species in acid soils ofpH (1:5 soil: water) below 5.0, or even below pH5.5 in kaolinitic soils (Foy 1984; Kamprath andFoy 1985). Excess available aluminium in subsoilscan reduce plant access to subsoil water andnutrients due to limited foot proliferation inlOsubsoil horizons (Foy 1988). The visual symp­toms of aluminium toxicity in legumes includemarked stunting and thickening of roots (Munns1965 a,b,c; B1amey et al. 1983; Haug 1983).There is often a close relationship between alu·minium status in soils and soil pH; increases inexchangeable aluminium are often associated withan increase in soil acidity (Blarney et al. 1987).Repons on the effects of H' and aluminium onnon-symbiotic growth of groundnut are limited.In relation to this, two solution culture experi­ments were undertaken to determine the toxicconcentrations of H' and aluminium for non­symbiotic growth of groundnut. Solution cultureexperiments have been used to overcome thedifficulty in studying the effects of soil pH per seon plalll growth.

MATERIALS AND METHODS

Two solution culture experiments were conductedat Universiti Pertanian Malaysia, Serdang, Malaysia\\'here daily temperatures of the nutrient solutionranged from 280 to 32°C. The minimum andmaximum air temperatures during the experi­mental period were 21.5° and 33.8°C, respec­tively. Solutions in the 51 pots were aerated bycontinuous bubbling using an aquarium pumpwith four outlets, one to each pot. Groundnut cv.Ma~am seeds were surface-sterilised with 95%alcohol for one minute, followed by 0.1 % HgCI,for twO minutes and five rinses with steriledeionised "later for one minute. Four-day-oldseedlings, germinated on sterilised sand, weretransferred by wrapping a sponge around eachseed and suspending the roots in the 51 nutdentsolution, three plants per pot.

pH Expe>imenlThe experiment was conducted using the deplet­ing nutrient method (Hewitt ] 966; Asher andEdwards 1983). Solution pH was measured dailyand adjusted with 0.05 M HCl and 0.05 M KOHto the desired pH, if necessary. The five pHvalues used were 3.5, 4.0, 4.8, 5.5 and 6.5. Theu·eatmenLS were arranged in a randomized com­plete block design with four replicates and a totalof 20 pots.

The initial nuu-ieot solution contained (JlM):N 1000, P 100, K500, Ca 400, Mg 100, S 677, Na114, CI 100, Fe 14, Zn 0.38, Mo 0.05, Mn 2.2, B23, Co 0.02, and eu 0.16. These concentrationswere achieved by calculating the total amount ofthe respective element required in 51 volumes ofsolu lion. No additional nutrient was given duringthe experiments. Plants were grown for 21 daysin the absence of aluminium.

Aluminium Expe1iment

The experiment was conducted using the Pro~

grammed Nutrient Addition Technique (Asherand Blarney 1987), The treatments were arrangedin a randomized complete block design with fourreplicates and a total of 24 pots. Six concenU'a~

tions of aluminium, as Al2(S04)3. 16H20, were

applied, viz. 0, 4, 8, 25, 50, and 110 JlM total-AI,equivalent to 0, 2, 4, 21, 47 and 108 JlMmonomeric-AI or 0, 1.5, 3, 15, 35 and 80 JlM(aAhnono (sum of activities of monomeric-All.

Al monomeric = AI'- + AI(OH)'- + AI(OH)o' +AI (OH)O, +AI(SO.,)- -

(Equation I)

The culture solution \Vas continuously aeratedfor seven days LO allow for equilibration of nutri­ents and treatments imposed (equilibration pe­riod). On the seventh day, the solution was agi­tated and allowed 10 stand for Ih before solutionsalnples were taken at a depth of 4cm from l..he17 em-high pots. Solution pH and electricalconductivity (mS em"l) were measured. Molarionic strength was estimated as 0.013 3 electricalconductivity (Griffin and ]urinak 1973; Gillmanand Bell 1978). Phosphate (5 JlM) and pH (4.3)levels were kept low to maintain the levels of AI­treaOllents as described by Kim (1985), to avoidformation of soluble, non-phytotoxic aluminium­polymers and to prevent precipitation of amor­phous aluminium phosphate. Solution pH wasmeasured daily and adjusted to pH 4.3 with 0.05M HCl or 0.05 M NaOH, if necessary. Phospho­rus concentrations in the solution were measureddaily by the molybdenum-blue method (Jackson1957) and maintained at 5 j,tM by addition ofNaH,lO~.2H20. All other nutrients were sup­plied daily for 28 days using the ProgrammedNutrient Addition Technique (Asher and Blarney1987) to give total nuuient solution concentra­tions as follows (JlM): 582, K522, Ca 400, Mg154, S 96, Fe 4.5, B 4.5, Mn 2.2, CI 3.05, Cu 0.5,Zn 0.2 and Mo 0.05.

190 PERTANIKA VOL. 15 NO.3, 1992

TOXIC CONCENTRATIONS OF H' AJ."IJD AL ON NONSYMBIOTIC GROWTH OF GROUNDNUT

- log f = (Az'V~) / (I + Bav'~) (Equation 2)

Al (OH); • = = =: » AI" + 20H- pK 19.3

AI (OH),O « = = = » AI" + 30H- pK 26.8

AI (SO)' « == = » Al:» + 5°-12- pK 3.2

where f = activity coefficient of the ion; ~ = ionicstrength of the solution; z = valence of the ion;a = ionic diameter; A = 0.5115; and B = 0.3291(Kielland 1937), and the following equations(Marion et at. 1976):

RESULTS AND DISCUSSION

1. Effects of pH per se on Growth of Groundnuta. Dry Matter Yield of Tops and Roots, and

Root LengthThe H+ ion concentration at pH 3.5 did notaffect the top dry weight of groundnut and theplants produced 90% of maximum total dry matteryeild (Table 2). There was no significant differ­ence between pH values from 3.5 to 6.5 on drymatter yield of tops. In contrast, Blarney et at.(1982) found that a solution pH of 3.5 was lethalfor plant top growth of sunflower (Helianthusannuus). In that study, the critical solution pHcorresponding to 90% of maximum total drymatter yield for four sunflower cultivars rangedfrom 3.9 to 4.5. In the present experiment, theroot dry weight and root length were reduced atthe lower pH (3.5) compared to the higher pHtreatments (4.0-6.5) (Table 2). At pH 3.5, the

The ionic diameters used were as follows:AI" &0.9 nm; AI (OH)" 0.7 nm; AI (OH)', &0.6nm; AI(Sa,)' 0.43 nm; and 50/- 0.4 nm (Kielland1937; Nair and Prenze! 1978). The solution com­positions are shmvn in Table 1.

Leaf tissues and roots required for chemicalanalyses were dried in an oven at 65·C for 4 daysand later ground using a 'Wiley mill ,\~th stainlesssteel blades and lining. After dry ashing (Anon1980). phosphorus and potassium concentrationswere determined using an autoanalyzer, and cal­cium and magnesium were determined usingatomic absorption spectrophotometer. After wetashing (Anon 1980), nitrogen was determinedwith an autoanalyzer.

The following data. were recorded: top androot dry weight. root length, and nitrogen, phos­phorus, potassium, calcium and magnesium con­centrations in the YEL and rooLS.

pK 8.99AI>- + OH-AI (OH)'-

Total-AI in solution was determinedcolorimeu-ically, using the aluminon method ofHsu (1963). The concenn-atlon of monomeric-A]in solution was determined by the modifiedaluminon method (Blarney et at. 1983). Thismethod employed the same procedure as thatdescribed by Hsu (1963) for total-AI, but withoutthe acid digestion (addition of HCI and heatingfor 30 min in a waterbath at 80°C). Colourintensity was read 30 min after the addition ofaluminon butler solution. The LaAllnOnO was calcu­lated from solution parameters as described byBlarney et at. (1983), which assumed the presenceof sulphate in solution, and pH range to bebetween 4.0 and 6.0.

The sum of activities of the five monomeric­Al species (~a..mO") (Equation 1) was calculatedusing solution pH, electrical conductivity, totalsulphate and the measured concentrations oftotal and monomeric-AI in solution by the AlmanaComputer Programme, the Debye-Hucke1 equa­tion

TABLE 1Calculated activities of aluminium monomers in nutrient solution, total·Al. monomeric-AI and electrical

conductivity at pH 4.3 prior to planting

Total-AI Mono-AI EC ~aAlmonoAI:1• AI(OH)'· AI(OH); AI (OH)," AI(SO,)·

(~M) (~M) (mS/cm) (~M) (~M) (~M) (~M) (~M) (~M)

0 0 0.29 0 0 0 0 0 04 2 0.30 1.5 0.59 0.09 0.29 0.002 0.528 4 0.30 3.0 1.17 0.19 0.59 0.003 1.05

25 21 0.31 15.0 5.94 0.84 2.93 0.02 5.4150 47 0.33 35.0 13.42 2.05 6.72 0.03 12.83

110 108 0.35 80.0 29.64 4.65 14.81 0.07 30.82

PERTANIK.A. VOL. 151 0.3,1992 191

ROSyllN KASRAN, ZULKIFU H, SHAMSUDDIN Al'lD D,G, EDWARDS

TABLE 2Relative yield of tops, top and root dry weights

and root length of groundnut grown at different solution pH

Solution ReI. Yield Top dry Root dry Root

pH of tops weight weight length(%) (g plant") (g pLant ") (em)

3,5 90' 3.45~ 0,59' 30,3'4,0 94' 3.58a 0,79' 39,0'4,8 100' 3,83' 0,77' 45,2'5.5 95" 3,63' 0,75' 45,8'6.5 94' 3,59' 0,72' 45,2'

Values followed by different lelters ,...i.I..hin a column arc significantly different at 5% level by Duncan's Multi­ple Range Test

'"

tion were not apparent. Soybean tap fOOts grownin the nutrient solution portion of a split me­dium also showed a lower calcium concentrationat pH 4,5 than at 5,6 (Lund 1970), Several nutri­ent solution studies have shown that a high H+ion concentration decreases the uptake of cal­cium, magnesium and potassium (Islam et al.1980), manganese (Robson and Loneragan 1970),

roots were shon, thickened, and brown. Islam etal. (1980) showed that lateral root developmentof J\!lanihot esculenta (cassava), Phaseolus vulgaris(French bean), lingiber officinaie (ginger), leamays (maize), L),copersicon esculentum (tomato) andTriticum aestivum (wheat) was suppressed and insome cases the rout tips were killed at pH 3.3. Asolution pH < 4.2 has been shown lO greatlyreduce the growth rate of the primary roots ofcotton (Gossypiwn spp.) (Howard and Adams1965), Suthpradit (19SS) found that groundnutcv. Red Spanish grew best in nowing culture withadequate inorganic nitrogen at pH 3.5 com­pared to other higher pH treatments (maximumpH 5.5). Groundnut was the most tolerant croplegume SUldied and reached maximal growth ofshoots and roOlS after 25 days at pH 3.5.

In contrast to roots of plants grown at lowpH, roots of plants grown at pH ~ 4.8 were light­coloured and finely branched. Relative root lengthincreased with increase in solution pH from 3.5to 4.8 (Fig. 1); interpolation indicates that 90% ofmaximum relative root length was achieved atpH 4.4. Thus, raising the solution pH was morebeneficial to growth of roots than tops.

u,,,

'"

'",< ~••~ 40

~"

1 • 125.'1, 60·02.639.'

, • OM!

b. Nutrient Concentrations in Youngest Expanded

Leaves

Cation uptake is suppressed in the presence ofhigh H' concentrations (Islam et al. 1980), Thiswas evident in the present experiment where pH3.5 markedly reduced the calcium concentrationin the VEL compared to pH .. 4,0 (Table 3), Incontrast to the stfong effects of H' ions in inhib­iting calcium absorption at pH 3.5, effects of H'ions at pH 3.5 in inhibiting magnesium absorp-

'"

'"

Fig. 1: Relative Toot length ofgroundnut at differentsolution pH

192 PERTANlKA VOL 15 NO, 3, 1992

TOXIC CONCENTRATIONS OF W ,'u'lD AL ON NONSYMBIOTIC GROWTH OF GROUNDNUT

70

90

".

Y. 101· 1.1SX .. 0.00'72X'.

1_ 0.989

20

50

.."

..

zinc (Rashid et al. 1976) and copper (Bowen1969). FawZj' et al. (1954) showed thaIprelTeaunent of barley roots for 3 hours at pH 3in the presence of Api and Ca2

+ considerablyreduced the loss of endogenous potassium. Thepresence of these ions in the preu'eaonent phasehad a protective effect on the subsequent abilityof barley fOOts to absorb potassium at pH 5.0.However, in the present solution cullUre study,nitrogen, phosphorus, potassium and magnesiumconcentration in the YEL were not limited byexcess W even at a pH as low as 3.5 (Table 3).This ability of groundnut to better maintain ad­equate concentrations of macronutrients in itstissues than other plant species when grown atlow pH makes it one of the morc acid-tolerantlegume species. in keeping with the findings ofAdams and Peal'son (1970).

2. Effects oj Aluminium on Groundnut

a. Dry Matte,. Yield oj Tops and Roots, and RootLength

The relative yield of groundnut LOps was inde­pendent of ~aAlmono from 0 to 15 )lM, but de­clined strongly v"ith further increase in La"'l",,,no to35 ~M and 80 ~M (Table 4). The critical LaAJrnOO"

corresponding to 90% relative yield of tops was12.2~M (fig. 2). Root dry weight was lower in thezero aluminium control treatment and in thehighest aluminium treatment (LaAllll01l0 of 80 J.lM)than in all other treatments (Table 4).

Symptoms of aluminium toxici[}' were notapparent on the leaves but a thickening of theroOlS and inhibition in root growth were obviouswithin 3 days of applying the higher concentra­tions of aluminium. The root systems in the

"

10 20 30 .40 50 60 70 III

··A1I'1\01'1DI.,oMI

Hg. 2: Relative yield ojgroll1ldnut tops at different values

of2:a'\I..",,~

La,\lmOnO of 35 and 80 )lM treatmenL'i were mark­edly inhited in length, coralloid shaped, andbrown, and the lateral roots were stubby with nofine branching, symptoms characteristic of alu~

minium toxicity (Foy 1984). Bouma et al. (1981)found that in AI-injured plants, aluminium typi­cally accumulates in or on roots but not in tops.

A stimulation in root length was observed atthe lower (LaA1mo,OO of 1.5 JlM) treatment while atthe higher (La,\Imono ~ 5 JlM) treatments a marked

TABLE 3Nitrogen, phosphorus, potassium, calcium and magnesium concentrations in

VEL of groundnut grown at different solution pH

Solution Nutrient concentrations in VEL (%)pH

Nitrogen Phosphorus Potassium Calcium Magnesium

3.5 2.33' 0.15' 2.34' 0.2!}' 0.46"4.0 2.29' 0.15' 2.51' 0.76' 0.41'''4.8 2.60' 0.15' 2.64' 0.76" 0.38'5.5 2.65' 0.14' 2.87' 0.87"< 0.36'6.5 2.77' 0.15' 3.35b 0.9[' 0.40'

Values followed by different letters wi.thin a column are significantly different at 5% levelby Duncan's Multiple Range Test

PERTANlKA VOL. 15 NO.3, 1992 193

ROSMIN KASRAN, ZULKIFLI H, SHAMSUDDIN AND D.G. EDWARDS

TABLE 4Relative yield of tops, top and rOOl dry weights, and

root length of groundnut grown at different values of (aMmon"

Total-AI Mono-AI (aAlmono ReI. yield Top dry Root dry Rootin soln. of tops weight weight length

(11M) (11M) (11M) (%) (g planr') (g plant') (em)

0 0 0 98< 2.79< 0.33' 38.9<4 2 1.5 100" 2.85c 0.40b 43.1'8 4 3.0 92(; 2.61" 0.40b 41.6'"

25 21 15.0 91" 2.59c O.44b 42.4'50 47 35.0 61b 1.75b 0.40b 22.Gb

110 108 80.0 37& 1.05" 0.33' 14.P

Values followed by different letters within a column are significantly different at 5% level by Duncan's Multi­ple Range Test

reduction in root length (Table 4) was observed.Other sLUdies have also shown a positive growthresponse of u'opical legumes at low concentra­tions of aluminium, although aluminium is notregarded as an essential plant nutrient (Andrewet al. 1973; Foy et al. 1978; Foy and Fleming1978).

b. Nutrient Concentrations in YEL

In general, an increase in (aAlmono resulted in anincrease in the nitrogen, phosphorus, potassiumand calcium concentrations in the YEL; a reversetrend was observed for magnesium (Table 5).Truman et al. (1986) found that aluminium wasantagonistic to the uptake of calcium and magne­sium but increased the uptake of potassium by

Pinus mdiata. Similar observations on alu·minium increasing the uptake of phosphorushave been reported (Andrew and VandenBerg 1973; Andrew et al. 1973; Murphy et ai,1984). Murphy et al. (1984) showed that thephosphorus concentration in the tops of fourtropical pasture legumes increased with in­crease in solution aluminium concentrationto 25 or 50 11M, and then decreased withfurther increase in solution aluminium con­centration to 100 or 125 J.lM. Thus, the toxiceffect of aluminium on the uptake of severalnutrient elements varies with plant speciesand genotype.

An increase in the activity of monomericaluminium species to 80 ~M depressed the

TABLE 5Nitrogen, phosphorus, potassium, calcium and magnesium concentrations in YEL and magnesium

concentrations in roots of groundnut grown at different values of {a"lmono

Nutrient concentrations (%) in:Total-AI Mono-AI (aAlmono VEL Rootin soIn.

(11M) (11 M) (11M) p K Ca Mg Mg

0 0 0 3.05' 0.28' l.i7~ 0.06' 0.29' 0.05'4 2 1.5 3.37:1b 0.33b 1.89' 0.19b 0.29' 0.07"8 4 3.0 3.70b 0.34b 1.93' 0.20b 0.30' 0.08'

25 21 15.0 3.69b 0.36b 1.93' O.25b 0.28' 0.08'50 47 35.0 5.49' O.52d 2.50b 0.36' O.24b 0.09'

110 108 80.0 5.80' 0.44' 3.08' 0.42' 0.\8' 0.07b

Values followed by different letters \vithin a column are significantly different at 5% level by Duncan'sMultiple Range Test

194 PERTANIKA VOL. 15 NO.3, 1992

TOXIC CONCENTRATIONS OF H' AND AL ON NONSYMBJOT1C GROWTH OF GROUNDNUT

magnesium concentration in the YEL to the de­ficient value of 0.18% (Reuter and Robinson1986). In this study, magnesium concentration inthe YEL and plant growth were significantly re­duced at LaAimono~ 35 J.1M; however, magnesiumdeficiency may have attenuated the growth re­sponse only at LaAllIIuno of 80 J.1M. The visualmagnesium deficiency symptoms appeared asgreyish-green areas on the leaf, first at the tip andthen along the leaf margin. Gradually this discol­oration changed to yellow and eventually brownwith the chlorosis extending interveina11y. Thedecrease in magnesium concentration in the YELat La

Aimonoof 35 IlM was not accompanied by a

corresponding decrease in magnesium concen­tration in the root (Table 5). This indicated thatthe translocation of magnesium to the plant tops,and not the uptake of magnesium by roots, wasreduced at high (~ 35 11M LaAlmono) aluminiumconcentrations. Thus, in agreement with Fay(1988), we concluded that for plants grown un­der aluminium stress, the accumulation of nutri­ents in plant tops probably reflects prior rootdamage resulting in differential transport of ele~

ments.The experiment showed that aluminium was

not toxic up to ~aA1l1lQ"O of 12.2 ~M, whichcorresponds to 90% of maximum growth ofgroundnut. The results also indicated that mag~nesium concentration in the YEL was more sen­sitive to aluminium toxicity than magnesium con~

centration in the roots.

CONCLUSION

The pH level recommended for growth of ground­nut was ~4.4. The H+ ion concentration at pH3.5 did not affect top dry weight of groundnutbut markedly decreased root dry weight andlength and reduced the uptake of calcium. Rais­ing the solution pH lO 4.0 was more beneficial togrowth of roots than tops.

The non-toxic level of aluminium for growthof groundnut tops and root elongation was LaAlmono";12.2 11M, which corresponds to a 10% depres­sion of growth of tops from the maximumachieved in the absence of aluminium while alow concentration of 4 11M total-AJ (= 1.5 11MLa

Almon) produced a positive growth response.

Magnesium concentration in the YEL was moresensitive to aluminium toxicity than magnesiumconcentration in the roots.

ACKNOWLEDGEMENTS

The authors are indebted to Universiti PertanianMalaysia and to the Australian Centre for Inter­national Agricultural Research (ACIAR). Thisresearch formed part of ACIAR Project 8375,"The Management of Soil Acidity for SustainedCrop Production".

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(Received 15 January 1992)

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