nutrient supply by mass flow and diffusion to maize plants ... · ermelinda maria mota oliveira...

NUTRIENT SUPPLY BY MASS FLOW AND DIFFUSION TO MAIZE PLANTS IN RESPONSE... 317

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NUTRIENT SUPPLY BY MASS FLOW AND DIFFUSION TO

MAIZE PLANTS IN RESPONSE TO SOIL AGGREGATE SIZE

AND WATER POTENTIAL(1)

Ermelinda Maria Mota Oliveira(2), Hugo Alberto Ruiz(3),Víctor Hugo

Alvarez V.(3), Paulo Afonso Ferreira(4), Fernanda Oliveira Costa(5) &

Ivan Carlos Carreiro Almeida(5)

Nutrients are basically transported to the roots by mass flow and diffusion.The aim of this study was to quantify the contribution of these two mechanisms tothe acquisition of macronutrients (N, P, K, Ca, Mg, and S) and cationicmicronutrients (Fe, Mn, Zn, and Cu) by maize plants as well as xylem exudatevolume and composition in response to soil aggregate size and water availability.The experiment was conducted in a greenhouse with samples of an Oxisol, fromunder two management systems: a region of natural savanna-like vegetation(Cerradão, CER) and continuous maize under conventional management for over30 years (CCM). The treatments were arranged in a factorial [2 x (1 + 2) x 2] design,with two management systems (CER and CCM), (1 + 2) soil sifted through a 4 mmsieve and two aggregate classes (< 0.5 mm and 0.5 - 4.0 mm) and two soil matricpotentials (-40 and -10 kPa). These were evaluated in a randomized block designwith four replications. The experiment was conducted for 70 days after sowing.The influence of soil aggregate size and water potential on the nutrient transportmechanisms was highest in soil samples with higher nutrient concentrations insolution, in the CER system; diffusion became more relevant when water availabilitywas higher and in aggregates < 0.5 mm. The volume of xylem exudate collectedfrom maize plants increased with the decrease in aggregate size and the increasedavailability of soil water in the CER system. The highest Ca and Mg concentrationsin the xylem exudate of plants grown on samples from the CER system were relatedto the high concentrations of these nutrients in the soil solution of this managementsystem.

Index terms: Cerradão, continuous maize , soil water potential.

(1) Parte da Tese de Doutorado do Primeiro autor. Recebido para publicação em março de 2008 e aprovado em janeiro de 2010.(2) Doutora em Solos e Nutrição de Plantas, Universidade Federal de Viçosa – UFV. Av. Peter Henry Rolfs s/n, CEP 36570-000

Viçosa (MG). E-mail: [email protected].(3) Professor do Departamento de Solos, UFV. Bolsista CNPq. E-mails: [email protected]; [email protected](4) Professor do Departamento de Engenharia Agrícola, UFV. Bolsista CNPq. E-mail: [email protected](5) Mestrando do Programa de Pós-graduação em Solos e Nutrição de Plantas, UFV. E-mails: [email protected];

[email protected]

318 Ermelinda Maria Mota Oliveira et al.

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RESUMO: IMPORTÂNCIA DO FLUXO DE MASSA E DA DIFUSÃO NOSUPRIMENTO DE NUTRIENTES AO MILHO E DA COMPOSIÇÃODO EXSUDATO XILÉMATICO EM RESPOSTA AO TAMANHODOS AGREGADOS E AO POTENCIAL DA ÁGUA DO SOLO

O transporte de nutrientes até as raízes é essencialmente efetuado por fluxo de massa edifusão. O objetivo deste trabalho foi quantificar a contribuição desses dois mecanismos naaquisição de macronutrientes (N, P, K, Ca, Mg e S) e de micronutrientes catiônicos (Fe, Mn, Zne Cu) por plantas de milho, e o volume e a composição do exsudato xilemático em resposta aotamanho dos agregados e à disponibilidade de água. O experimento foi conduzido em casa devegetação com amostras de um Latossolo Vermelho, retiradas sob dois sistemas de manejo:cerradão sem manejo (CER) e cultivo contínuo com milho em manejo convencional por maisde 30 anos (CCM). Os tratamentos corresponderam ao arranjo fatorial [2 x (1 + 2) x 2], sendodois sistemas de manejo (CER e CCM), (1 + 2) solo passado em peneira de 4 mm e duas classesde agregados (< 0,5 mm e entre 0,5 e 4,0 mm), e dois potenciais de água do solo (-40 e -10 kPa).Eles foram dispostos em delineamento em blocos casualizados, com quatro repetições. Aos 70dias após a semeadura, finalizou-se o ensaio. A influência do tamanho do agregado e dopotencial de água do solo nos mecanismos de transporte de nutrientes é maior nas amostras desolo com maiores concentrações de nutrientes em solução, sistema CER, tendo a difusão crescidoem importância na condição de maior disponibilidade de água e nos agregados < 0,5 mm. Ovolume de exsudato xilemático colhido de plantas de milho aumenta com a diminuição dotamanho do agregado e com o aumento da disponibilidade de água do solo, no sistema CER.As maiores concentrações de Ca e Mg no exsudato xilemático das plantas cultivadas em amostrasretiradas do sistema CER, dependeram das altas concentrações desses nutrientes na soluçãodo solo neste sistema de manejo.

Termos de indexação: cerradão, cultivo contínuo com milho, conteúdo de água do solo.

INTRODUCTION

To be taken up by plants, the nutrients in the soilsolution must be transported to the root surface. Themechanisms of solute transport to the roots are,essentially, mass flow and diffusion.

The mass flow is associated with the total potentialgradient regulating the water movement in the soil-plant-atmosphere. Thus, the soil solution concentrationand plant transpiration rate determine the quantityof ions transported through this mechanism (Barber,1974).

The relative importance of mass flow for nutrientsupply depends on the supply capacity of the soil,besides the plant demand, characterized by differencesin transpiration rate and nutrient uptake, whichvaries according to the plant species, root activity,soil water content and nutrient (Marschner, 1995).Mathematically, the quantity transported by massflow (FM, kg m-2 s-1) is given by:

FM = q C (1)

where q is the water flow (m3 m-2 s-1) and C is theaverage solute concentration (kg m-3).

Nutrient diffusion, in turn, occurs through randomthermal motion of ions towards the root, due to theconcentration gradient near the root surface by the

absorption process (Barber, 1984). The soil watercontent, the interaction of nutrient with soil colloidsand the distance the nutrients must overcome to reachthe root surface are the main factors governing thistransport mechanism (Wild, 1981). The diffusioncoefficient (DS, m2 s-1) is calculated by the equation:

DS = D1 θ f dCl/dCS (2)

where Dl is the diffusion coefficient in a pure solution(m2 s-1), θ is the soil water content, based on volume(m3 m-3); f is the impedance factor (m m-1), whichincludes, among other variables, the tortuosity ofporous media, and dCl/dCS is the inverse value of thebuffering capacity, where Cl is the nutrientconcentration in the soil solution, the intensity factor(kg L-1), and Cs the adsorbed nutrient, the quantityfactor (kg m-3) (Novais & Smyth, 1999).

Some authors also cite root interception as one ofthe mechanisms responsible for plant nutrient supply.This mechanism occurs when nutrients areintercepted by the roots during the growth process.Marschner (1995) reported that a small percentage ofthe total nutrient requirement is met by this process.Other authors believe that root interception shouldbe disregarded because there is no possibility of a directexchange between soil particles and plant rootswithout a liquid environment (Ruiz et al., 1999).


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After root uptake, the nutrients are translocatedwithin the plant, reaching stems and leaves(Marschner, 1995). This transport occurs throughthe xylem and subsequently, nutrients areredistributed in the phloem. Nutrients are quicklydistributed through the corrente transpiracional/transpiration current (Larcher, 2000). It is thereforeimportant to know the composition of the xylemexudate to evaluate nutrient absorption andtranslocation from roots to the aerial part. The studyof the xylem sap is, in essence, one of the mostrepresentative of the actual conditions of plantnutrition (Vale et al., 1984).

The predominant mechanism for the transport ofa nutrient is determined by its concentration in thesolution moving toward the roots in response to thepotential difference. When the nutrient amount thatreaches the roots is equal to or exceeds the absorbedquantity, mass flow is the only relevant mechanism.In this case, the excess nutrients would accumulatein the rhizosphere (Barber, 1974). When thenutritional requirement of the plant exceeds theamount carried by mass flow, diffusion becomes thecomplementary mechanism, which may even exceedin importance the mass flow (Barber, 1962; Araújo etal., 2003).

Decreased water availability in the soil reducesthe nutrient movement by mass flow and diffusion.These conditions lead to stomatal closure andconsequent reduction of both flow transpiration as wellas mass flow. In addition, it also decreases theamount of CO2 entering the stomata, and the rootwater and nutrient uptake. Under these conditions,plant growth is reduced and the amount of plant-available water becomes a limiting factor for growth(Costa et al., 1997; Barros & Comerford, 2002).

The contribution of mass flow and diffusion totransport nutrients to the roots can be calculatedroughly by measuring the individual contributionsand comparing them with the total plant nutrientuptake. In practice, the total nutrient accumulationand the contribution of mass flow are generallydetermined; the difference between the total amountabsorbed and the quantity transported by mass flowwas attributed to diffusion (Barber, 1984). Thistechnique allows estimating the average contributionsduring the test period.

In soils, the nutrient movement to the rootsincludes two steps. At first, the transport from theaggregate interior to the surface occurs by diffusionin motionless solution (intra-aggregate porosity). Inthe second, from the aggregate surface to the roots,transport in the mobile solution is necessarily carriedout by mass flow. However, in weathered soils suchas Oxisols with a low nutrient concentration in thesoil solution, the mass flow is mostly insufficient tomeet the plant demand for some nutrients. Thus,diffusion becomes a complementary mechanism, oreven the predominant mechanism in the nutrient

supply to the plant roots, as observed by Ruiz et al.(1999) and Araújo et al. (2003).

This study aimed to quantify the contribution ofmass flow and diffusion to the acquisition ofmacronutrients (N, P, K, Ca, Mg and S) and cationicmicronutrients (Fe, Mn, Zn, and Cu) by maize, andthe volume and composition of xylem exudate inresponse to soil aggregate size and water potential insamples of an Oxisol, sampled in two managementsystems: savanna-like with no management (CER)and continuous maize under conventionalmanagement for more than 30 years (CCM).

MATERIAL AND METHODS

The experiment was conducted in a greenhouseusing samples of an Oxisol collected at Embrapa Maizeand Sorghum in Sete Lagoas, Minas Gerais, Brazil.The samples were collected in the surface layer (2–15cm) in two adjacent areas under differentmanagements, one with natural Cerradão vegetation(CER), and the other with continuous maize underconventional system for more than 30 years (CCM).

These samples were air-dried and sieved througha 4 mm sieve. A part was used to separate dryaggregates into two diameter classes: < 0.5 mm and0.5 - 4.0 mm, in a sieve shaker (Produtest) with 50 x 50cm sieves. No fertilizer was added to the soil samples.Both soil and aggregates were physically andchemically characterized (Table 1).

The test was conducted in a factorial [2 x(1 + 2) x 2] design with two management systems:savanna-like with no management (CER) andcontinuous maize under conventional management(CCM); (1 + 2) soil sieved through 4 mm mesh andtwo aggregate classes (< 0.5 and 0.5–4.0 mm); andtwo soil water potentials (-40 and -10 kPa), arrangedin randomized blocks with four replications. Theexperimental units consisted of plastic pots with 5dm3 of soil or aggregates, with a previously adjustedmoisture content of -10 kPa.

After the period required for uniform moisture,maize was sown (variety BRS-3060) with six seedsper pot and thinned to three plants per pot seven daysafter germination. From sowing to thinning, the soilwater potential in pots was adjusted to -10 kPa. Inthe treatments of -40 kPa no water was added untilthe desired potential was reached. The soil waterpotential was controlled daily during the maizecultivation period, using one tensiometer per pot.Based on the data of the water amount applied,evapotranspiration was calculated. In addition to thepotted plants two pots with no plants for eachtreatment were prepared, to monitor water loss byevaporation. Plant transpiration in each experimentalunit was calculated as the difference between thewater volume of evapotranspiration and of evaporation.


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Plants were harvested 70 days after emerging anddivided in aerial parts and roots. After cutting theaerial part the xylem exudate was collected by directcontact with capillary tubs in the incision region for2 h. The exudate was always collected in the earlymorning, sampling two blocks per day.

The xylem exudate was placed in glass vialspreviously weighed with an accuracy of ± 0.0001 g,determining the whole mass of the set. The collectedvolume was estimated by weight difference, assuminga value of 10,000 kg L-1 for solution density. In thismaterial, the concentration of P, K, Ca, Mg, S, Fe,Mn, Zn, and Cu was determined by direct reading inan Inductively Coupled Plasma Optical EmissionSpectrometer (ICP-OES). Using the concentrationsof these nutrients and the amount of exudate collectedin each experimental unit, the content of P, K, Ca,Mg, S, Fe, Mn, Zn, and Cu in the xylem exudate wascalculated.

Soil and aggregate samples of all treatments weredried and sieved to 2 mm. Then, three sub-sampleswere taken for soil solution extraction. To each 1 dm3

sub-sample of either soil or aggregate the water volumerequired to achieve a moisture content of -5 kPa wasadded. After a period of 8 h of equilibrium, the soilsolution of each of these sub-samples was extractedwith pressure membrane extractor at 1000 kPa for12 h. The concentration of K, Ca, Mg, S, Fe, Mn, Zn,and Cu was determined, by direct reading in ICP-OES in each extracted solution. P, N-NO3

- and N-NH4

+ were determined by colorimetry (Braga &Defelipo, 1974; Kempers & Zweers, 1986; Yang et al.,1998).

The amounts of N, P, K, Ca, Mg, S, Fe, Mn, Zn,and Cu transported by mass flow were determined bymultiplying the concentration of each nutrient in thesoil solution by the water volume transpired in thetest period. The diffusion was calculated bysubtracting the amount transported by mass flowfrom the total nutrient uptake (aerial part + roots)(Ruiz et al., 1999; Rosolem et al., 2003; Oliveira etal., 2004). With the values of mass flow and diffusion,the relative contribution of each mechanism to nutrienttransport was calculated.

Table 1. Physical and chemical characterization of soil and aggregates (< 0.5 and 0.5–4.0 mm) in samples ofan Red Oxisol under two management systems(1)

(1) Natural Cerradão vegetation without management (CER) and continuous maize under conventional management (CCM).(2) Ruiz (2005). (3) Dry sieving (Embrapa, 1997). (4) Yang et al. (1998). (5) Kempers & Zweers (1986). (6) Mehlich-1 extraction solution(Defelipo & Ribeiro, 1981). (7)Extraction solution KCl 1 mol L-1 (Embrapa, 1997). (8) Extraction solution calcium acetate 0.5 mol L-1

pH 7.0 (Embrapa, 1997). (9) Yeomans & Bremner (1988).


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The results were submitted to analysis of variance.The effect of management system was compared andfive orthogonal contrasts were used in each soilmanagement system, CER and CCM (Table 2). Two

additional contrasts (CA1 and CA2) were included todecompose the effect of the potential within eachaggregate class. Mean contrasts were used, dividingthe result of each contrast by ½ Ó |ci|, where |ci|denotes the absolute value of its coefficient (AlvarezV. & Alvarez, 2006). The significance of contrastswas tested statistically by the F-test at 5 and 1 %.

RESULTS AND DISCUSSION

When comparing the management systemsnatural Cerradão vegetation (CER) versus continuousmaize under conventional management for over30 years (CCM), it appears that the nutrient uptakein plants grown in soil or aggregate samples from theCER system was greater, except in the case of K(Figures 1 and 2). This response may be due to a highernutrient concentration in the soil solution (Table 3)and also to the greater nutrient availability, withexception of P, K and S in the CER system (Table 1).

Table 2. Contrasts (C) used in the comparison of soilor aggregates (AG), and soil water potential (ΨΨΨΨΨ),in each management system(1)

(1) C1: soil vs. AG. C2: AG < 0.5 mm vs. AG 0.5–4.0 mm. C3:Ψ -40 kPa vs. Ψ -10 kPa for the soil. C4: Ψ -40 kPa vs. Ψ -10 kPafor AG < 0.5 mm. C5: Ψ -40 kPa vs. Ψ -10 kPa for AG 0.5–4.0 mm.

Figure 1. Total nitrogen, phosphorus and potassium accumulation in maize plants and quantities suppliedby mass flow (MF) and diffusion (D), considering the material used for testing, soil or aggregates (AG),the water potential (ΨΨΨΨΨ) and the management system. Natural vegetation or continuous maize cropping.C1: soil vs. AG. C2: AG < 0.5 mm vs. AG 0.5–4.0 mm. C3: ΨΨΨΨΨ - 40 kPa vs. ΨΨΨΨΨ -10 kPa for soil. C4: ΨΨΨΨΨ - 40 kPa vs. ΨΨΨΨΨ-10 kPa for AG < 0.5 mm. C5: ΨΨΨΨΨ - 40 kPa vs. ΨΨΨΨΨ -10 kPa for AG 0.5–4.0 mm. * and ** significant at 5 and 1%,respectively, by the F test.


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Chemical analysis (Table 1) showed that N is thedecisive nutrient for the cited differences, whichresulted in significantly higher values of nutrientuptake, especially of N, Ca and Mg under CER,compared to CCM (Figures 1 and 2). It is noteworthythat dry matter production under CER was also higherin the aerial part (14.47 mg/pot) and roots (5.43 mg/pot) than in CCM (8.91 mg/pot and 3.47 mg/pot). Thisexplains why the low N availability in the soil samplesfrom the area under CCM limited the dry matterproduction of plants grown in this managementsystem

In the case of the longstanding use of theconventional system with periodic fertilization over

30 years (CCM), the results of soil chemicalcharacterization showed that this period was sufficientto reduce the soil nutrient levels, except for P, K andS (Table 1). The first two nutrients are generallyincorporated in higher quantities, mainly in the caseof P, and S can be incorporated when ordinarysuperphosphate is used. This finding justifies thelower nutrient concentrations in the soil solution ofplants grown on samples from CCM (Table 3).

The low nutrient concentration in the soil solutionwas due to nutrient depletion in the soil solution inthe 70 days of the test (Table 3). It is noteworthythat these concentrations were used to estimatenutrient transport by mass flow. Assuming that the

Figure 2. Total calcium, magnesium and sulphur accumulation in maize plants and quantities supplied bymass flow (MF) and diffusion (D), considering the material used for testing, soil or aggregates (AG), thewater potential (ΨΨΨΨΨ) and the management system. Natural vegetation or continuous maize cropping. C1:soil vs. AG. C2: AG < 0.5 mm vs. AG 0.5–4.0 mm. C3: ΨΨΨΨΨ - 40 kPa vs. ΨΨΨΨΨ -10 kPa for soil. C4: ΨΨΨΨΨ - 40 kPa vs. ΨΨΨΨΨ -10 kPa for AG < 0.5 mm. C5: ΨΨΨΨΨ - 40 kPa vs. ΨΨΨΨΨ -10 kPa for AG 0.5–4.0 mm. * and ** significant at 5 and 1%,respectively, by the F test.


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nutrient concentration in the soil solution wasconstant from the beginning of the experiment mayhave been an underestimation of the contribution ofmass flow and overestimation of diffusion, since thelatter was determined by the difference between thetotal plant uptake and the amount supplied by massflow. The contribution of root interception wasdisregarded since the percentage of contribution of thismechanism to the process of nutrient transport toplant roots is negligible.

Of the nutrient supply mechanisms to plants roots,mass flow was the main mechanism for Ca, Mg andN (Figures 1 and 2), in agreement with resultsobtained for maize by Vargas et al. (1983) and for riceby Ruiz et al. (1999). In all treatments, the Ca amounttransported by mass flow exceeded the amountaccumulated in the plants, aside from those grownon smaller aggregates (< 0.5 mm), with reduced wateravailability (-40 kPa) in soil samples from the CCMsystem (Figure 2). This result confirms the existenceof mechanisms responsible for a decreased uptake ofthis nutrient when the plant demand is met, e.g., Caaccumulation on the root surface.

There is a pronounced difference in N, Ca, and Mguptake between samples from the CER and from theCCM system (Figures 1 and 2). This result can beexplained by the higher concentration of thesenutrients in soil solution samples from the CERsystem (Table 3), resulting in higher transport bymass flow (Figures 1 and 2).

The transport of sulfur to maize roots was not onlyrelated to mass flow, but also to diffusion, dependingon the management system, CER or CCM (Figure 2).Thus, mass flow contributed with 40 % of S supply tothe roots in the CER and 51 % in the CCM system.Obviously, the contribution of diffusion had an inverseeffect. This difference is mainly due to higherconcentrations of this nutrient in CCM soil solution

samples, favoring the mass-flow transport (Table 3).At adequate concentrations in the soil, the supply ofother nutrients that may be carried solely by massflow may be complemented by the diffusion process,when the concentrations of these nutrients are low insoil solution. This was observed for nitrogen (Okajima& Taniyama, 1980; Strebel & Duynisveld, 1989) andmagnesium (Al-Abbas & Barber, 1964; Vargas et al.,1983).

For K, the predominant mechanism was diffusion,contributing with nearly 85 and 95 % of K absorbedin the CER and CCM systems, respectively (Figure 1),which is consistent with results obtained by Barber(1974). This author cites diffusion as the maintransport mechanism of potassium in the soil solutionto plant roots, accounting for 86 % of its supply, whilemass flux and root interception represented 11 and3 %, respectively. In Brazil, working with differentsoils of Rio Grande do Sul, Vargas (1983) found thatthe diffusion mechanism contributed with 72 - 95 %of the potassium supply for maize. Ruiz (1999),Rosolem et al. (2003) and Fernandes (2006) also foundthat diffusion was the main supply mechanism of Kto plant roots.

It was observed that diffusion was also the maintransport mechanism for P, Fe, Mn, Zn, and Cu,mainly for aggregates < 0.5 mm and highest waterpotential (-10 kPa), regardless of the managementsystem CER or CCM (Figure 1 and Table 4). Thisresult is justified by the low concentration of thesenutrients in the soil solution and the greater wateravailability in smaller aggregates (Tables 1 and 3).These results corroborate those of other authors(Vargas et al., 1983; Nunes et al., 2004).

Cu transported by mass flow and by diffusion inthe CER and CCM samples was not included in Table 4since the element was not detected by the analyticalmethod used in the analyses of soil solution and of the

Table 3. Nutrient concentration in the soil solution, considering the material used for testing, soil oraggregates (AG), and the management system (MS)(1)

(1) Natural Cerradão vegetation (CER) and continuous maize under conventional management (CCM). (2) Different letters incolumn indicate differences at 1% by the F test for management system. (3) Concentration below the detection limit of ICP-OES.


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aggregates separated from the CCM sample (Table 3).The low Cu concentration in the solution extractedfrom CER samples indicates that more than 99 % ofthe transport of this nutrient must be attributed todiffusion. The lower Cu concentration in the CCMsamples suggests that under these conditions, thecontribution of diffusion would be proportionatelygreater.

The percentage contribution of transport by massflow for both management systems was: Ca > Mg > N> S > K > P ≈ Mn ≈ Zn ≈ Fe ≈ Cu. The mean for thefirst five is 100, 70, 60, 40 and 15 %, respectively (CER)and 98, 57, 52, 51 and 5 % (CCM). Diffusion is themain transport mechanism of K, P, Fe, Mn, Zn, andCu to the surface of maize roots in both managementsystems, accounting for values above 99 % except forK, where the value is 85 % for the CER system and95 % for the CCM system.

Comparing the aggregate size in both systems, CERand CCM (C2: aggregate size, AG <AG 0.5 mm vs.0.5 to 4.0 mm), it appears that the contribution of massflow to nutrient supply increases in larger aggregates(Figures 1 and 2), especially in the CER system: Ca(C2: 187.5**mg/pot), Mg (C2: 53.23** mg/pot), S (C2:6.35** mg/pot) and K (C2: 21.13** mg/pot), and underCCM: N (C2: 29.69** mg/pot), S (C2: 11.78** mg/pot)K (C2: 7.21** mg/pot). In the case of diffusion, thebehavior is obviously inversed. In this more sandymaterial (0.5–4.0 mm aggregates), the lowerinteraction of nutrients with the components of thesoil solid phase favored higher concentrations in soil

solution (Table 3). It is likely that this result is alsodue to the tortuosity of the porous environment. Inlarger, more sandy aggregates, tortuosity is higherunder unsaturated conditions than in smalleraggregates. So when there is a reduction in theimpedance factor of the material formed by largeraggregates, the nutrient supply by diffusion to theroot surface is reduced. This explanation is valid,because under non-saturated conditions the hydraulicconductivity decreases more sharply in sandy thanin clay soils (Hillel, 1971).

On the other hand, contribution of the transportmechanism by diffusion in aggregates < 0.5 mm wasgreater in both management systems, but moremarkedly in CER (Figures 1 and 2). The differencesbetween the aggregate sizes may be related toincreased water retention with decreasing aggregatesize (Table 1), due to increased clay and organicmatter content in those aggregates. The higher watercontent in smaller aggregates favors diffusion by itsdirect effect on the diffusion coefficient and by reducingthe tortuosity of the diffusive path, thus increasingthe impedance factor and, consequently, the coefficientof diffusion (Barber, 1984).

Another fact to consider is the high concentrationof fine roots in the aggregates < 0.5 mm, in the CERsystem. The higher values of root dry weight in theaggregates < 0.5 mm (12.44 g/pot) compared toaggregates 0.5–4.0 mm (8.48 g/pot) allow theconclusion that in aggregates < 0.5 mm the depletionzone around the roots that favored diffusion was larger

Table 4. Total Fe, Mn and Zn accumulation in maize plants and quantities supplied by mass flow (MF) anddiffusion (D), considering the material used for testing, soil or aggregates (AG), the water potential (ø)and the management system (MS)(1)

(1)Natural Cerradão vegetation (CER) and continuous maize under conventional management (CCM). (2)Different letters incolumn indicate differences at 1% by the F test for management system.


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(Figures 1 and 2). This also demonstrates thateventual problems of aeration or compaction due tothe predominance of small and more clayey aggregateswere not sufficient to reduce the transport by diffusionin these materials.

When comparing the soil water potential, - 40 vs.-10 kPa (C3, C4 and C5), the increase in wateravailability led to a greater nutrient uptake, favoringmass flow and diffusion in the CER (Figure 1 and 2).In CCM samples, the effect of water potential on thenutrient transport was almost negligible, due to thelow N availability in the system.

A positive and significant effect was observed inthe contrasts comparing the soil water potentials-40 with. -10 kPa in CER (C3, C4 and C5) (Figures 1and 2) and that the amount of nutrient transportedby diffusion increased under greater water availability,especially P (C3: 8.04**; C4: 11.39**; C5: 8.62**), S(C3:3.75**; C4: 3.22**; C5: 3.04**), Mg (C3: 8.18**; C4:25.42**) and K (C5: 42.81**) (Figures 1 and 2). Thisresult was also observed for Fe, Mn, Zn, and Cu(Table 3, contrasts not shown). The higher watercontent in soil encourages diffusion by its direct effecton the diffusion coefficient and by reducing thetortuosity of diffusion pathway, increasing thethickness of the water film within the pores. Themoisture can also influence the ion distributionbetween the soil solid and liquid phases. Nye & Tinker(1977) related the impedance factor (f) to soil moistureand found that in very dry soil, f showed very lowvalues (2 x 10-4 at a matric potential of -10 MPa and

10-2 at -1.5 MPa). When the potentials were between-0.1 and -1.0 MPa, the impedance factor increasedlinearly similar to the water content. According tothe authors, the impedance factor approaches zero inthe driest soils reaching values between 0.4 and 0.7in saturated soil.

The management systems CER and CCMinfluenced the exudate volume collected from maizeplants (Table 5). It was observed that the volumecollected within 2 h from plants grown on CERsamples was approximately eight times higher thanthat collected from plants grown on soil undercontinuous cropping with conventional tillage maize(CCM). This result is consistent with the values ofdry matter, which were higher in CER (19.9 g/pot)than in CCM (12.7 g/pot). This confirms thatconditions for transport by mass flow and diffusionwere more favorable in CER soil samples, whichresulted in higher dry matter production.

Nutrient concentrations in the maize exudates wereinfluenced by the management systems; significantdifferences were observed for all nutrients, except S(Table 5). It was not possible to determine N, since Ncannot be quantified by ICP-EOS and the exudatevolume was insufficient for the specific analyticaldetermination. Plants grown in CER soil sampleshad higher concentrations of Ca, Mg and Mn and lowerlevels of P, K, Fe, Zn, and Cu (Table 5). Consideringthe five macronutrients determined in the exudate,the concentration ratio of the soil solutions whencomparing the two management systems, CER and

Table 5. Volume (V) and mineral composition of the xylem exudate, considering the material used for testing,soil or aggregates (AG), the water potential (ΨΨΨΨΨ) and the management system (MS)(1)

(1) Natural Cerradão vegetation (CER) and continuous maize under conventional management (CCM). (2)Different letters incolumn indicate differences at 1 % by the F test for management system.


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MCC, were 2.71, 1.65, 7.04, 6.18 and 0.64, for P, K,Ca, Mg and S, respectively (Table 3). The appreciabledifference in the concentrations in soil solution Caand Mg allowed higher concentration of these nutrientsin the exudate removed from plants grown in CERsoil samples. In the case of P and K, similarconcentrations and significantly lower exudatevolumes in plants grown on samples of the CCMincreased the concentration of these elements. Inrelation to S, the lower concentration in the systemCER was compensated by a higher exudate volume,without statistical differences between the twomanagement systems. For the micronutrients, thelow concentrations in soil solution indicate that theabsorption of these elements responded more tometabolic than osmotic effects, leading to differentvalues (Table 5).

Based on the mean nutrient concentration in thexylem sap (Table 5) and the concentration in soilsolution (Table 3), the relationship between these twovariables was calculated. The values were 487.9 (P),30.1 (K), 5.7 (S), 4.5 (Mg) and 0.7 (Ca) in samples ofthe CER system. In addition, diffusion was responsiblefor the transport of 99.5, 84.8, 60.3, 29.7 and 0 % ofthese nutrients, respectively. Considering that rootpressure is a consequence of factors involved inmetabolic (active absorption) and osmotic processes(concentration gradient) (Zholkevich, 1991), theconclusion could be drawn that the higher thecontribution to transport by diffusion, the greater theparticipation of metabolic processes in the plantnutrient uptake.

Similar calculations were performed for samplesof the CCM system, where the values for therelationship between concentrations were: 6,072 (P),54.7 (K), 10.3 (Mg), 3.0 (S) and 2.2 (Ca). Since diffusionwas responsible for transporting 99.8 (P), 94.8 (K),49.5 (S), 43.3 (Mg), and 2.4 % (Ca), there is an invertedsequence in the comparison of Mg and S.

Analyzing the effect of aggregate size (C2: -1.00**),it appears that the largest exudate volume wascollected from plants grown in aggregates < 0.5 mmin the CER system only. This result was due to thehigher water availability in these than in largeraggregates (Table1). The exudate volume collectedfrom maize plants was influenced by water availability(C3: 1.47**, C4: 1.21** and C5: 0.93**) in CER soil only.This result demonstrates the strong dependence of theexudate volume on the soil water potential (Table 5).

CONCLUSIONS

1. A greater nutrient availability resulted inincreased mass flow and diffusion transport of N, Ca,Mg, S, P, Fe, Mn, Zn, and Cu to the roots in a comparisonof natural Cerradão vegetation (CER) with continuousmaize in a conventional system for more than 30 years(CCM).

2. The proportion of mass flow contribution tonutrient transport was the following: Ca > Mg > N >S > K > P ≈ Mn ≈ Zn ≈ Cu ≈ Fe. The mean values forthe first five were 100, 63, 56, 45 and 10 % respectively.

3. The diffusion was the main mechanism of K, P,Fe, Mn, Zn, and Cu transport, accounting for valuesabove 99 % except for K, where the value was 90 %.

4. The influence of aggregate size and soil waterpotential is higher in soil samples with higher soilsolution nutrient concentrations.

5. The xylem exudate volume increases with areduced aggregate size and increased soil wateravailability. The nutrient concentration in the xylemexudate is higher than in soil solution, with exceptionof Ca.

LITERATURE CITED

AL-ABBAS, H. & BARBER, S.A. Effects of root growth andmass-flow on the availability of soil calcium andmagnesium to soybeans in a greenhouse experiment.Soil Sci., 97:103-107,1964.

ALVAREZ V., V.H. & ALVAREZ, G.A.M. Comparação demédias ou teste de hipóteses? Contrastes! B. Inf. SBCS,31:24-34, 2006.

ARAÚJO, C.A.S.; RUIZ, H.A.; SILVA, D.J.; FERREIRA, P.A.;ALVAREZ V., V.H. & BAHIA FILHO, A.F.C. Eluição demagnésio, cálcio e potássio de acordo com o tempo dedifusão em colunas com agregados de um LatossoloVermelho distrófico típico. R. Bras. Ci. Solo, 27:231-238,2003.

BARBER, S.A. A diffusion and mass-flow concept of soilnutrient availability. Soil Sci., 93:39-49, 1962.

BARBER, S.A. Influence of the plant root on ion movement insoil. In: CARSON, E.W., ed. The plant root and itsenvironment. Charlottesville, University Press ofVirginia, 1974. p.525-564.

BARBER, S.A. Soil nutrient bioavailability: A mechanisticapproach. New York, John Wiley, 1984. 398p.

BARROS, N.F. & COMERFORD, N.B. Sustentabilidade daprodução de florestas plantadas na região tropical. In:ALVAREZ V., V.H.; SCHAEFER, C.E.G.R.; BARROS,N.F.; MELLO, J.W.V. & COSTA, L.M., eds. Tópicos emciência do solo. Viçosa, MG, Sociedade Brasileira emCiência do Solo, 2002. v.2. p.487-592.

BRAGA, J.M. & DEFELIPO, B.V. Determinação espectrofo-tométrica de fósforo em extratos de solos e plantas. R.Ceres, 21:73-85, 1974.

COSTA, L.C.; MORISON, J. & DENNETT, M. Effects of waterstress on photosynthesis, respiration and growth of Fababean (Vicia faba L.) growing under field conditions. R.Bras. Agron., 5:9-16, 1997.


R. Bras. Ci. Solo, 34:317-327, 2010

DEFELIPO, B.V. & RIBEIRO, A.C. Análise química de solo.Viçosa, MG, Universidade Federal de Viçosa, 1981. 17p.(Boletim de Extensão)

EMPRESA BRASILEIRA DE PESQUISA AGROPECUÁRIA -EMBRAPA. Centro Nacional de Pesquisa de Solos.Manual de métodos de análise de solo. 2.ed. Rio de Janeiro,Centro Nacional de Pesquisa de Solos, 1997. 212p.

FERNANDES, M.S. Nutrição mineral de plantas. Viçosa, MG,Sociedade Brasileira de Ciência do Solo, 2006. 432p.

HILLEL, D. Soil and water. Physical principles and processes.New York, Academic Press, 1971. 288p.

KEMPERS, A.J. & ZWEERS, A. Ammonium determinationin soil extracts by the salicylate method. Comm. Soil Sci.Plant. Anal., 17:715-723, 1986.

LARCHER, W. Ecofisiologia vegetal. São Carlos, RiMa, 2000.531p.

MARSCHNER, H. Mineral nutrition of higher plants. 2.ed.London, Academic Press, 1995. 889p.

NOVAIS, R.F. & SMYTH, T.J. Fósforo em solo e planta emcondições tropicais. Viçosa, MG, Universidade Federal deViçosa, 1999. 399p.

NUMES, F.N.; NOVAIS, R.F.; SILVA, I.R.; GEBRIM, F.O. &SÃO JOSÉ, J.F.B. Fluxo difusivo de ferro em solos sobinfluência de doses de fósforo e de níveis de acidez eumidade. R. Bras. Ci. Solo, 28: 423-430, 2004.

NYE, P.H. & TINKER, P.B. Solute movement in the soil-rootsystem. Berkeley, University of California Press, 1977.342p.

OLIVEIRA, R.H.; ROSOLEM, C.A. & TRIGUEIRO, R.M.Importância do fluxo de massa e da difusão no suprimentode potássio ao algodoeiro como variável de água e potássiono solo. R. Bras. Ci. Solo, 28:439-445, 2004.

OKAJIMA, H. & TANIYAMA, I. Significance of mass flow innitrate-nitrogen supply to plant roots. Soil Sci. Plant Nutr.,26:363-374, 1980.

ROSOLEM, C.A.; MATEUS, G.P.; GODOY, L.J.G.; FELTRAN,J.C. & BRANCALIÃO, S.R. Morfologia radicular esuprimento de potássio ás raízes de milheto de acordocom a disponibilidade de água e potássio. R. Bras. Ci.Solo, 27:875-884, 2003.

RUIZ, H.A. Incremento da exatidão da análise granulométricado solo por meio da coleta da suspensão (silte + argila). R.Bras. Ci. Solo, 29:297-300, 2005.

RUIZ, H.A.; MIRANDA, J. & CONCEIÇÃO, J.C.S.Contribuição dos mecanismos de fluxo de massa e dedifusão para o suprimento de K, Ca e Mg a plantas dearroz. R. Bras. Ci. Solo, 23:1015-1018, 1999.

STREBEL, O. & DUYNISVELD, W.H.M. Nitrogen supply tocereals and sugar beet by mass flow and diffusion on siltyloam soil. Z. Pflanzenernährung. Bodenk., 152:135-141,1989.

VALE, F.R.; NOVAIS, R.F.; SANT’ANNA, R. & BARROS, N.F.Absorção e translocação de fosfato em milho suprido comnitrato ou amônio e pré-tratado com alumínio. R. Bras.Ci. Solo, 8:219-222, 1984.

VARGAS, R.M.B.; MEURER, E.J. & ANGHINONI, I.Mecanismos de suprimento de fósforo, potássio, cálcio emagnésio às raízes de milho em solos do Rio Grande doSul. R. Bras. Ci. Solo, 7:143-148, 1983.

WILD, A. Mass flow and diffusion. In: GREENLAND, D.J. &HAYES, M.H.B., eds. The chemistry of soil processes.Chichester, John Wiley, 1981. p.37-80.

YANG, J.E.; SLOGLEY, E.O.; SCHAFF, B.E. & KIM, J.J.A.Simple spectrophotometric determination of nitrate inwater, resin and soil extracts. Soil Soc. Am. J., 62:1108-1115, 1998.

YEOMANS, J.C. & BREMNER, J.M. A rapid and precisemethod for routine determination of carbon in soil. Comm.Soil Sci. Plant Anal., 19:1467-1476, 1988.

ZHOLKEVICH, V.N. Root pressure. In: WAISEL, Y.; ESHEL,A. & KAFKAFI, U., eds. Plant roots: The hidden half.New York, Marcel Dekker, 1991. p.589-603.


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