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This article was downloaded by: [Moskow State Univ Bibliote] On: 18 February 2014, At: 21:00 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Communications in Soil Science and Plant Analysis Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/lcss20 Effect of Barium on Growth and Macronutrient Nutrition in Tanzania Guineagrass Grown in Nutrient Solution Francisco Antonio Monteiro a , Roberta Corrêa Nogueirol a , Leônidas Carrijo Azevedo Melo a , Adriana Guirado Artur a & Fabiana da Rocha b a Department of Soil Science , University of São Paulo/ESALQ , Piracicaba, Brazil b Department of Soil Science and Rural Engineering , Federal University of Mato Grosso , Cuiabá, Brazil Published online: 27 Jun 2011. To cite this article: Francisco Antonio Monteiro , Roberta Corrêa Nogueirol , Leônidas Carrijo Azevedo Melo , Adriana Guirado Artur & Fabiana da Rocha (2011) Effect of Barium on Growth and Macronutrient Nutrition in Tanzania Guineagrass Grown in Nutrient Solution, Communications in Soil Science and Plant Analysis, 42:13, 1510-1521, DOI: 10.1080/00103624.2011.581725 To link to this article: http://dx.doi.org/10.1080/00103624.2011.581725 PLEASE SCROLL DOWN FOR ARTICLE Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”) contained in the publications on our platform. However, Taylor & Francis, our agents, and our licensors make no representations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of the Content. Any opinions and views expressed in this publication are the opinions and views of the authors, and are not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be relied upon and should be independently verified with primary sources of information. Taylor and Francis shall not be liable for any losses, actions, claims, proceedings, demands, costs, expenses, damages, and other liabilities whatsoever or howsoever caused arising directly or indirectly in connection with, in relation to or arising out of the use of the Content. This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. Terms &
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Page 1: Effect of Barium on Growth and Macronutrient Nutrition in Tanzania Guineagrass Grown in Nutrient Solution

This article was downloaded by: [Moskow State Univ Bibliote]On: 18 February 2014, At: 21:00Publisher: Taylor & FrancisInforma Ltd Registered in England and Wales Registered Number: 1072954 Registeredoffice: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK

Communications in Soil Science andPlant AnalysisPublication details, including instructions for authors andsubscription information:http://www.tandfonline.com/loi/lcss20

Effect of Barium on Growth andMacronutrient Nutrition in TanzaniaGuineagrass Grown in Nutrient SolutionFrancisco Antonio Monteiro a , Roberta Corrêa Nogueirol a , LeônidasCarrijo Azevedo Melo a , Adriana Guirado Artur a & Fabiana da Rochab

a Department of Soil Science , University of São Paulo/ESALQ ,Piracicaba, Brazilb Department of Soil Science and Rural Engineering , FederalUniversity of Mato Grosso , Cuiabá, BrazilPublished online: 27 Jun 2011.

To cite this article: Francisco Antonio Monteiro , Roberta Corrêa Nogueirol , Leônidas CarrijoAzevedo Melo , Adriana Guirado Artur & Fabiana da Rocha (2011) Effect of Barium on Growth andMacronutrient Nutrition in Tanzania Guineagrass Grown in Nutrient Solution, Communications in SoilScience and Plant Analysis, 42:13, 1510-1521, DOI: 10.1080/00103624.2011.581725

To link to this article: http://dx.doi.org/10.1080/00103624.2011.581725

PLEASE SCROLL DOWN FOR ARTICLE

Taylor & Francis makes every effort to ensure the accuracy of all the information (the“Content”) contained in the publications on our platform. However, Taylor & Francis,our agents, and our licensors make no representations or warranties whatsoever as tothe accuracy, completeness, or suitability for any purpose of the Content. Any opinionsand views expressed in this publication are the opinions and views of the authors,and are not the views of or endorsed by Taylor & Francis. The accuracy of the Contentshould not be relied upon and should be independently verified with primary sourcesof information. Taylor and Francis shall not be liable for any losses, actions, claims,proceedings, demands, costs, expenses, damages, and other liabilities whatsoever orhowsoever caused arising directly or indirectly in connection with, in relation to or arisingout of the use of the Content.

This article may be used for research, teaching, and private study purposes. Anysubstantial or systematic reproduction, redistribution, reselling, loan, sub-licensing,systematic supply, or distribution in any form to anyone is expressly forbidden. Terms &

Page 2: Effect of Barium on Growth and Macronutrient Nutrition in Tanzania Guineagrass Grown in Nutrient Solution

Conditions of access and use can be found at http://www.tandfonline.com/page/terms-and-conditions

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Communications in Soil Science and Plant Analysis, 42:1510–1521, 2011Copyright © Taylor & Francis Group, LLCISSN: 0010-3624 print / 1532-2416 onlineDOI: 10.1080/00103624.2011.581725

Effect of Barium on Growth and MacronutrientNutrition in Tanzania Guineagrass Grown in

Nutrient Solution

FRANCISCO ANTONIO MONTEIRO,1 ROBERTA CORRÊANOGUEIROL,1 LEÔNIDAS CARRIJO AZEVEDO MELO,1

ADRIANA GUIRADO ARTUR,1 AND FABIANA DA ROCHA2

1Department of Soil Science, University of São Paulo/ESALQ, Piracicaba,Brazil2Department of Soil Science and Rural Engineering, Federal University of MatoGrosso, Cuiabá, Brazil

Barium has been identified as a toxic element to most plants, although for grasses thetoxicity has not been determined. A greenhouse experiment was performed to evaluatethe effect of barium on growth parameters, barium accumulation, and macronutrientconcentration in Tanzania guineagrass (Panicum maximum Jacq.), cultivated in nutri-ent solution. Five barium rates and a control were set in a complete randomized blockdesign, with four replications. Forage yield, leaf area, barium, and macronutrient con-centrations and accumulation were measured. Leaf area and yield sharply decreasedwith increase of barium concentration in the nutrient solution. The greatest bariumconcentration and accumulation were found in culms and sheaths. Toxic barium con-centrations were estimated to be 1.24 mmol L−1 (170 mg L−1) in nutrient solution and225 mg kg−1 in the diagnostic leaf, and the main symptoms of toxicity were interveinalchlorosis followed by necrotic spots in the leaf laminae of the grass.

Keywords Barium toxicity, nutritional disorders, Panicum maximum, toxicitysymptoms

Introduction

Barium (Ba) is contained in recycled wastes, and agricultural fields may accumulate it withtime and loading. The inappropriate deposition of wastes on pasture lands may increase theamount of heavy metals, including Ba, in the food chain, resulting in serious environmentaland health risks. Copper (Cu) accumulation in beef cattle was observed in the Galiciaregion (Spain) after the use of swine waste as pasture fertilizer (López Alonso et al. 2000).Barium concentration in tissues of wild asparagus (Asparagus officinalis) grown near wastedeposition sites reached a maximum of 22 mg kg−1 and no visual symptoms of toxicityappeared, as reported by Brandt and Rickard (1996).

Barium mean contents in most plants range from 2 to 13 mg kg−1, with an exceptionof blueberries, in which Ba levels of 160 mg kg−1 are reported. The greatest Ba contentswere reported for Brazil nuts from up to 3,000 to 4,000 mg kg−1 (Kabata-Pendias andMukherjee 2007). Barium can be found in several soil types, and all plants usually show

Received 18 November 2009; accepted 15 July 2000.Address correspondence to Francisco Antonio Monteiro, Department of Soil Science,

University of São Paulo/ESALQ, 9, 13418-900, Piracicaba, Brazil. E-mail: [email protected]

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Barium in a Forage Grass 1511

at least 4 to 50 mg kg−1 (Chaudhry, Wallace, and Mueller 1977). In soils under pasture,for example, Ba selenate is a common source of selenium applied to supplement cattlenutrition. Despite this, Whelan (1993) did not find an increase in Ba concentration in redclover (Trifolium subterraneum), even after three years of Ba selenate applications. Bariumuptake by plants and its transport from roots to shoots may increase exposure to Ba inhumans and animals through vegetable or forage consumption, respectively. Greater Bashoot than root concentrations have been found by Llugany, Poschenrieder, and Barceló(2000) in bean (Phaseolus vulgaris) plants.

A study with Scotland soils by Mitchell (1957) reported average Ba concentra-tions of 42 mg kg−1 in shoots of Trifolium subterraneum and 18 mg kg−1 in Loliumperenne. In another study, no symptoms of Ba toxicity were observed in shoots ofsunflower (Helianthus annuus), mustard (Brassica juncea), and castor bean (Ricinus com-munis) grown under greenhouse conditions in a Rhodic Hapludox soil supplied with 150and 300 mg kg−1 of Ba as Ba sulfate (BaSO4), and the greatest Ba concentration wasobserved in sunflower (21 mg kg−1), followed by mustard (19 mg kg−1) and castor bean(11 mg kg−1) (Coscione and Berton 2009). Research on forage plants has shown thatBa concentration varies with the sampled plant part and also depends upon the soil typewhere they were grown (Chamberlain and Miller 1982). In nutrient solution, Debnath andMukherji (1982) found growth inhibition of Vigna radiate with a 0.1 mmol L−1 Ba chlo-ride (BaCl2). Suwa et al. (2008) emphasized that literature information on Ba toxicity toplants is scarce, but the available reported results confirm its potential phytotoxicity.

Barium toxicity can be reduced by calcium (Ca), magnesium (Mg), and sulfur (S)supplied through the growth medium, as a result of the antagonistic interaction betweenthese macronutrients and Ba, both in soil and inside the plant (Kabata-Pendias and Pendias1992). The antagonistic interaction between Ba and S was studied by Wang (1988) andbetween Ba and Ca by Wallace and Romney (1971). To avoid Ba precipitation with sulfate,one strategy adopted to investigate Ba toxicity in plants cultivated in nutrient solution isthe use of Ba chloride; however, this strategy may lead to S deficiency in plants.

Barium phytotoxicity seems to be rare, although few researchers have focused on crit-ical toxicity concentrations (Llugany, Poschenrieder, and Barceló 2000). Because there isa lack of information about toxic concentrations and toxicity symptoms of Ba on tropicalforage plants, this study used Tanzania guineagrass to investigate (a) how the dry-matteryield and leaf area behaved when plants were exposed to increasing concentrations of Bain the growth medium; (b) concentration and accumulation of Ba in plant parts and therelated toxicity symptoms; (c) how Ba rates influence concentration of macronutrients inplant tissues; and (d) the toxic concentration of Ba.

Material and Methods

The experiment was carried out in a greenhouse at Piracicaba, State of São Paulo, Brazil,with the tropical forage grass Panicum maximum Jacq. cv. Tanzânia. The seeds were ger-minated in plastic trays containing washed sand. Fourteen days after sowing, when theseedlings were about 4 cm high, batches of 15 homogeneous seedlings were transplantedto 3.6-L plastic pots filled with ground quartz. Just after transplanting, each pot received 1 Lof Hoagland and Arnon (1950) solution diluted to 20% ionic strength, during 7 days. Fromthe 8th day on, undiluted Hoagland and Arnon (1950) solution was used for another 7-dayperiod, and then plants were thinned to five plants per pot, taking care to select the mosthomogeneous to minimize the difference between the plant materials used. A randomizedcomplete block design with four replications was used.

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1512 F. A. Monteiro et al.

To avoid precipitation of Ba sulfate and at the same time supply S properly to theplants, two types of nutrient solutions were used: one complete and the other completewithout S and containing Ba (Llugany, Poschenrieder, and Barceló 2000). Barium con-centrations were 0.5, 2.5, 5.0, 10.0, and 20.0 mmol L−1, and a control (no-Ba) solutionwas used. The supply of these nutrient solutions to the plants were alternated each 48 h,during 22 days (12 days in Ba solution). To oxygenate the roots and homogenize solu-tions, pot content was swirled three to four times per day, and solutions were completelydrained at the end of each day. Both nutrient solutions were replaced after 2 weeks. Asthe medium of growth was a nutrient solution, the MINTEQ speciation program was usedto estimate free Ba (Ba2+) concentration in each treatment. Thirty-six days after trans-planting, plants were harvested by cutting shoots at the substrate level and taking apart thetwo recently expanded leaf laminae (diagnostic leaves, RL), the other leaf laminae (OL),and culms + sheaths (CS). Leaf area was measured in a leaf area meter (LI-3000, Licor,Lincoln, Neb.) just after plant harvesting. The aboveground biomass (RL, OL, and CS) wasdried at 70 ◦C until constant mass was reached, and then the dry mass was recorded.

Roots were recovered from the substrate by washing through a nest of sieves, 1.00and 0.25 mm, and rinsing with deionized water. Twenty percent of the fresh root masswas transferred to a beaker containing a solution of 50 g L−1 gentian violet. The coloredroots were scanned, and their images were analyzed by the Integrated System for Root andSoil Coverage Analysis (SIARCS) to measure the length and the surface of the roots foreach pot (Crestana et al. 1994). Total dry mass of roots was recorded after drying at 70 ◦Cuntil constant mass. Specific length (root length / root dry mass) and specific surface (rootsurface / root dry mass) of the roots were calculated.

All plant material (RL, OL, CS, and roots) was ground in a Wiley mill with a 0.6-mmscreen and macronutrients and Ba concentrations were measured after nitric–perchloricdigestion. Calcium (Ca) and magnesium (Mg) concentrations were determined by atomicabsorption spectroscopy (AAS), phosphorus (P) was found by the colorimetric vanadate–molybdate method, potassium (K) was measured with flame photometry, and S wasdetermined by the turbidimetric method. Barium concentration was determined by induc-tively coupled plasma with optical emission spectrometry (ICP-OES). Total accumulationof these elements (uptake) was calculated by using the plant tissue element concentrationsand the plant dry mass. A 10% decrease in the maximum plant yield was used to estimatethe toxic concentration of Ba for this specie (Lima et al. 2007).

All data were subjected to analysis of variance (ANOVA), and if significant (P < 0.05),regression analysis (GLM) was done using the Statistical Analysis System software version6.08 (SAS 1996).

Results and Discussion

Symptoms of Nutritional Disorders

Plants grown in the 0.5 mmol L−1 Ba solution did not show any visible symptoms ofnutritional disorder and had growth similar to those in the control (no-Ba) treatment. Evenplants in the 2.5 mmol L−1 Ba rate had apparent normal development, except that the youngleaf laminae presented interveinal chlorotic strips. In the 5.0 mmol L−1 Ba rate, plantsshowed decreases in growth and interveinal chlorosis in all leaf laminae. With the supplyof 10.0 mmol L−1 of Ba in the solution, plants had retarded growth, stimulated senes-cence, interveinal chlorosis, and marginal necrotic spots in leaf laminae. Visible symptomsbecame more acute in plants grown in the 20 mmol L−1 Ba solution, which induced

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Barium in a Forage Grass 1513

chlorosis and necrosis in all leaf laminae, and anticipated the occurrence of necrosis inall mature laminae, as compared to the other Ba rates.

Assessment of Barium Toxicity

Since the solutions containing Ba and sulfate were supplied separately, it was possibleto estimate the Ba activity in the nutrient solution. Approximately 93% of the total Bawas present as free Ba (Ba2+) in the solution when 0.5 mmol L−1 was supplied andslightly reduced to about 88% when the Ba concentration was increased to 20.0 mmol L−1

(Table 1). Such a reduction was mainly due to precipitation with chloride (Cl−). Therefore,most Ba was available for plant uptake.

Dry-mass yield of shoots and roots decreased with increasing Ba concentration inthe nutrient solutions (Figures 1 and 2). These effects were particularly highlighted insolutions with Ba concentration up to 5.0 mmol L−1, and at the greatest Ba concentration(20 mmol L−1) all shoot parts yielded about 23% of that found in plants of the controltreatment. By supplying 0.1 and 1.0 mmol L−1 of Ba in the nutrient solution during 14 daysfor soybean (Glycine max), Suwa et al. (2008) found 15% and 40% decreases in dry-mass yield, respectively, as compared to the control plants. It has been shown that legumespecies are less tolerant to metal toxicity as compared to cereals and grasses (Obata andUmebayashi 1997).

To establish the critical level of boron toxicity for maize (Zea mays), Lima et al.(2007) adopted a 10% decrease in dry-mass yield. Considering that there is no reportedtoxic Ba rate for forage grass grown in nutrient solutions, a 10% decrease in shootyield was also adopted to estimate Ba toxic levels for Tanzania guineagrass. The exter-nal toxic critical level of total Ba in the nutrient solution was 1.24 mmol L−1. Planttissue critical concentrations for Ba toxicity were 225, 383, 562, and 156 mg kg−1 inthe two recently expanded leaf laminae (RL), other leaf laminae (OL), culms + sheaths(CS), and roots, respectively. These observed Ba concentrations could be used as atemporary reference to assess the risk of this metal contamination in Tanzania guinea-grass. However, further studies should be developed in pasture soils naturally rich inBa, or even in areas to which Ba-containing residues were added, to establish val-ues in a field situation. Also, it should be pointed out that these concentrations cannotnecessarily act as indicators of toxic levels for animals because plants usually tolerategreater levels of toxic metals than animals (Mengel and Kirkby 2001). For example, there

Table 1Percentage of free Ba2+ and other precipitated forms as related to total barium

concentrations supplied in the nutrient solution

Ba treatments (mmol L−1) Ba2+ BaNO3+ BaHPO4 (aq) BaCl+

0.0 (control: Ba free) — — — —0.5 92.9 5.2 1.6 0.32.5 92.7 5.3 1.4 0.65.0 92.2 5.5 1.4 1.010.0 91.1 5.8 1.3 1.820.0 88.2 6.7 1.3 3.9

Note. The speciation was performed using MINTEQ speciation program with the pHfixed at 7.0.

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Page 7: Effect of Barium on Growth and Macronutrient Nutrition in Tanzania Guineagrass Grown in Nutrient Solution

1514 F. A. Monteiro et al.

20.010.05.02.50.5

10

20

30

40

Barium rates (mmol L–1)

Shoot dry

mass (

g p

er

pot)

;24.00

R2 = 0.86**Ba

y = 10.78 +

Figure 1. Total dry-mass yield of Tanzania guineagrass shoots as related to barium rates in nutrientsolution.

Dry

mass (

g p

er

pot)

Barium rates (mmol L–1)

20.010.05.02.50.5

2

4

6

8

10 CS

; R2 = 0.79∗∗7.08

Bay = 3.12 +

; R2 = 0.77∗∗68.7

Bay = 3.56 +

; R2 = 0.82∗∗9.25

Bay = 4.10 +

; R2 = 0.83∗∗31.0 Bay = 8.05 −

OL

RL

ROOT

Figure 2. Dry-mass yield in shoots and roots components of Tanzania guineagrass as related tobarium rates in nutrient solution.

is no recommended minimum concentration of Ba in plants or animal diets in Australia(Whelan 1993).

Tanzania guineagrass leaf area showed similar responses to Ba rates as those observedfor shoot yield (Figure 3). Even at the lowest Ba rates, the grass leaf area was depressed.Because the absorption of light energy in greater plants involves chlorophylls andcarotenoids, located mainly in leaves, the depression in plant growth was a consequence ofthe low energy acquisition by plants with small leaf area.

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Page 8: Effect of Barium on Growth and Macronutrient Nutrition in Tanzania Guineagrass Grown in Nutrient Solution

Barium in a Forage Grass 1515

20.010.05.02.50.50

1000

2000

3000

4000

Leaf A

rea (

cm

2 p

er

pot)

Barium rates (mmol L–1)

;2410.61 R2 = 0.88∗∗Ba

y = 977.85 +

Figure 3. Total leaf area of Tanzania guineagrass as related to barium rates in nutrient solution.

The total root length and root surface were not significantly changed by Ba rates.However the specific root length and root surface linearly increased as the Ba rates wereincreased (Figure 4). Fitter (1996) outlined that greater specific root length indicates rel-ative greater proportion of thin roots, which grow as a strategy to search for nutrients inmedia with low nutrient availability. In this study, this root behavior may suggest an attemptto grow away from toxic Ba concentrations, which resulted in a stress to the roots.

Barium and Macronutrient Concentrations in Plant Tissues

Barium concentrations in RL and CS linearly increased and varied according to a second-degree model in the OL and roots as Ba rates were increased in the nutrient solution(Figure 5). Roots had the lowest and CS the greatest Ba concentrations in plant tissues.Chaudhry, Wallace, and Mueller (1977) found greater Ba concentrations in the leaves(ca. 22,000 mg kg−1) than in the stems (ca. 11,000 mg kg−1) when Ba rates were suppliedto common beans (Phaseolus vulgaris).

Llugany, Poschenrieder, and Barceló (2000) reported greater Ba concentrations in thetrifoliolate and primary leaves (ca. 9,000 mg kg−1) than in the stems (ca. 4,000 mg kg−1)or roots (ca. 6,000 mg kg−1) of beans, when Ba was applied at 5 mmol L−1 in the nutrientsolution. These results reinforce the statement that plant Ba concentrations greatly varyamong plant species (Nielsen 1986).

Barium stayed in the nutrient solution mainly (88.2% to 92.9%) as free Ba (Table 1).The concentration of this element was sharply increased in plant shoot tissues in agree-ment with the increased free Ba activity in the nutrient solution (Figure 5 and Table 1).On the other hand, by increasing Ba concentration there was a clear decrease in Ca andMg concentrations in the mature leaves (Table 2), which reflects an antagonistic relation-ship between those divalent cations, as observed by Lombnaes and Singh (2003), withmanganese and zinc.

Barium concentration in the roots was much lower than in shoot tissues (Figure 5). Asdescribed by Mengel and Kirkby (2001) for cadmium in plants, Ba was probably readilytransported from the nutrient solution through the grass roots to the upper plant parts. It can

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Page 9: Effect of Barium on Growth and Macronutrient Nutrition in Tanzania Guineagrass Grown in Nutrient Solution

1516 F. A. Monteiro et al.

a

Barium rates (mmol L–1)

20.010.05.02.50.5

Specific

root surf

ace (

cm

2 p

er

g)

400

600

800

1000

1200

y = 522.70 + 21.73 Ba; R2 = 0.94∗∗

y = 74.99 + 2.89 Ba; R2 = 0.94∗∗

Barium rates (mmol L–1)

20.010.05.02.50.5

b

Specific

root le

nght (c

m p

er

g)

60

90

120

150

Figure 4. Specific root surface (a) and specific root length (b) of Tanzania guineagrass as related tobarium rates in nutrient solution.

be hypothesized that free Ba was absorbed and readily transported in the upward movementof water in the xylem, in a way similar to that reported by Lombnaes and Singh (2003) forfree manganese.

By supplying Ba at low rates (≤0.5 mmol L−1), the accumulation of this ele-ment was similar in RL, OL, CS, and roots. Barium increases in the solution sharplyincreased Ba accumulation in CS, which was almost twice the amount found in RL orOL (Figure 6).

Phosphorus and K concentrations in RL and OL linearly increased as the Ba rateswere increased (Table 2). Potassium concentration in CS and roots increased with Ba ratesup to the 5.0 mmol L−1 treatment concentration, where the greatest K concentration inthese plant tissues was found. Potassium accumulation in CS of Panicum maximum (42–55% of the total amount of K in the shoots in the first harvest and 51–62% for the second

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Page 10: Effect of Barium on Growth and Macronutrient Nutrition in Tanzania Guineagrass Grown in Nutrient Solution

Barium in a Forage Grass 1517

Tis

sue B

a c

oncentr

ation (

mg p

er

kg)

20.010.05.02.50.50

3200

6400

9600

12800

y = –205.17 + 618.41Ba; R2 = 0.99**

y = –354.93 + 467.93Ba; R2 = 0.92**

y = 62.22 + 245.08Ba + 11.2 Ba2 ; R2 = 0.99**

y = 178.10–27.45Ba + 7.68 Ba2; R2 = 0.96**

CS

OL

ROOT

RL

Figure 5. Barium concentration in parts of Tanzania guineagrass as related to barium rates innutrient solution.

harvest of forage) was previously reported by Mattos et al. (2002). Increasing Ba supplyabove 5.0 mmol L−1 resulted in decreases in K concentration in CS and roots. These resultsagree with those reported by Suwa et al. (2008), who suggested that Ba would inhibit theopening of K channels in the membrane, resulting in reduced K absorption. A generaldecrease in both Ca and Mg concentrations was found in OL and roots as Ba rates wereincreased.

Sulfur was the macronutrient whose concentration linearly decreased in all plant parts,as Ba rate increased (Table 2). Since Ba and sulfate were supplied through alternatingsolutions, this effect may be explained by the occurrence of precipitation of this nutrient asBa sulfate within the plant. Barium sulfate has a very low solubility (Kps = 1.1 × 10−10)even in acidic media (Tubino and Simoni 2007). As demonstrated by Menzie et al. (2008),the solubility of most Ba compounds is not strongly pH dependent, and from pH 2.0 to 7.0the solubility of Ba sulfate remains much below that of the other Ba compounds, such aschloride, acetate, or nitrate.

Conclusions

Increase in Ba supply through nutrient solutions sharply reduced the leaf area and dry-massyield of Tanzania guineagrass. Culms and sheaths contained the greatest Ba concentrationsand accumulation. Besides the retarded growth, visible symptoms of Ba toxicity wereinterveinal chlorosis and marginal necrotic spots in the leaf laminae. As Ba supply wasincreased, S concentration decreased in all plant parts, whereas K, P, and Mg increased inthe two recently expanded leaf laminae (diagnostic leaves) of the grass. The toxic level offree Ba (Ba2+) in the nutrient solution was 1.24 mmol L−1, and the toxic Ba concentrationin the two expanded leaf laminae of the Tanzania guineagrass was 225 mg kg−1. Thesetoxic Ba concentrations and leaf symptoms in this grass can be used as a first field guide toidentify Ba toxicity in natural or contained tropical grasslands.

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Page 11: Effect of Barium on Growth and Macronutrient Nutrition in Tanzania Guineagrass Grown in Nutrient Solution

Tabl

e2

Mac

ronu

trie

ntco

ncen

trat

ions

inpa

rts

ofTa

nzan

iagu

inea

gras

sas

rela

ted

toba

rium

rate

sin

nutr

ient

solu

tion

Bar

ium

rate

sP

KC

aM

gS

Plan

tpar

t(m

mol

L−1

)(g

kg−1

)(g

kg−1

)(g

kg−1

)(g

kg−1

)(g

kg−1

)

Oth

erle

afla

min

a(O

L)

0.0

2.2

20.6

10.3

4.5

1.18

0.5

2.6

23.5

11.1

4.3

1.25

2.5

2.7

28.8

9.5

4.1

1.13

5.0

2.7

32.4

9.0

3.7

0.90

10.0

3.4

34.1

9.2

3.9

0.12

20.0

3.5

35.4

7.9

3.9

0.06

Eff

ect

L∗∗

L∗∗

L∗∗

L∗

L∗∗

CV

(%)

16.2

15.4

14.6

10.2

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Two

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ntly

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nded

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inae

(RL

)0.

02.

215

.46.

63.

11.

26

0.5

2.6

18.3

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1.23

2.5

3.2

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7.9

3.7

1.36

5.0

3.2

25.4

6.7

3.0

0.63

10.0

3.6

30.2

7.2

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20.0

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33.0

8.1

4.1

0.08

Eff

ect

L∗∗

L∗∗

nsL

∗L

∗∗C

V(%

)17

.615

.615

.316

.541

.0C

ulm

s+

shea

ths

(CS)

0.0

2.5

34.7

5.5

5.2

0.93

0.5

3.0

38.4

6.3

5.7

1.15

2.5

2.8

46.4

6.0

7.0

0.87

5.0

2.6

46.7

5.5

6.0

0.56

10.0

3.2

41.2

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7.0

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20.0

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31.0

4.7

5.4

0.07

Eff

ect

L∗∗

nsL

∗L

∗L

∗∗

1518

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Page 12: Effect of Barium on Growth and Macronutrient Nutrition in Tanzania Guineagrass Grown in Nutrient Solution

CV

(%)

13.8

13.2

14.0

12.1

49.8

Roo

ts0.

01.

812

.56.

52.

02.

510.

52.

014

.66.

12.

02.

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51.

916

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82.

10.

875.

02.

317

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41.

40.

7010

.02.

415

.43.

11.

20.

3920

.02.

19.

12.

40.

80.

10E

ffec

tL

∗∗ns

L∗∗

L∗∗

L∗∗

CV

(%)

11.7

18.3

27.0

18.5

56.2

Not

es.L

,lin

ear;

ns,n

otsi

gnifi

cant

;CV

,coe

ffici

ento

fva

riat

ion.

∗∗Si

gnifi

cant

atP

<0.

01.

∗ Sig

nific

anta

tP<

0.05

.

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Page 13: Effect of Barium on Growth and Macronutrient Nutrition in Tanzania Guineagrass Grown in Nutrient Solution

1520 F. A. Monteiro et al.

Barium rates (mmol L–1)

20.010.05.02.50.5

Tis

sue B

a a

ccu

mula

tion (

mg p

er

pot)

0

8500

17000

25500

34000

CSOL

ROOT

RL

y = 511.51 + 3858.40 Ba–112.89 Ba2 ; R2 = 0.98**

y = 2320.79 + 929.02 Ba; R2 = 0.97**

y = 1289.49–97.06 Ba + 24.22 Ba2; R2 = 0.86**

y = –682.15 + 2266.48 Ba–60.89 Ba2; R2 = 0.91**

Figure 6. Barium accumulation in parts of Tanzania guineagrass as related to barium rates innutrient solution.

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