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VII. Other Interactions Involving Root Exudates

VII. Other Interactions Involving Root Exudates

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stantial part of the increase in phosphatase activity near plant roots (McLachlan,

1980). In addition, the excretion of phosphatase by plant roots has been shown to

be stimulated by P deficiency (McLachlan, 1980; Amann and Amberger, 1989;

Grimal et al., 1992), suggesting that the release of these enzymes should be considered as an adaptative response of the P-deficient plant. However, in addition to

their possible degradation by microbes, the competition of these enzymes with microbial phosphatases and their possible inactivation by adsorption onto soil reactive components such as clay minerals (Quiquampoix et al., 1995; Leprince and

Quiquampoix, 1996) bring into question their effective role in soil environments.

Further work is needed in this area, since in many soils organic Pcontributes a major proportion of soil P.

Major root exudates are the so-called mucilage-a gelatinous material made of

high-molecular-weight polysaccharides (Curl and Truelove, 1986). Polyuronic

acids that are well known for their important role in the cation exchange capacity

of the root cell walls account for a large proportion of this mucilagenous exudate.

The consequent exchange properties of mucilage explain their ability to bind

heavy metals such as Pb and Cd or micronutrients such as Cu and Zn (Morel et al.,

1986; Mench et al., 1987). In acid soils, Al can similarly be detoxified by a massive adsorption on mucilage (Horst et al., 1982). In addition to these binding properties of polyuronic sites in mucilage with respect to metal cations, polyuronate

ions may help desorb some anions, such as phosphate ions sorbed on soil minerals as shown for polygalacturonate by Nagarajah et al. (1970). Such a process

agrees with the findings of Grimal er al. (1995). They showed that mucilage excreted by axenic-grown plants was sorbed on goethite, whereas phosphate was

desorbed from goethite in the rhizosphere of maize. Many other benefits have been

attributed to mucilage, including their role in establishing a better contact between

the roots and the porous soil matrix (Uren and Reisenauer, 1988), thereby improving the transfer of water and mineral nutrients to the roots.


The rhizosphere, i.e., the volume of soil that is influenced by root activities, can

exhibit drastically different conditions compared with the bulk soil. Since the rhizosphere conditions are those that are encountered by plant roots, understanding

them is critical to improving our knowledge of root functioning and plant nutrition. The rhizosphere was once recognized only for its singular microbiology.

However, over the last two or three decades, evidence has accumulated that severe

changes in chemical conditions relative to the bulk soil are a major trait of the rhizosphere. This review has concentrated on those peculiar modifications of chemical conditions that are occurring in the rhizosphere as a direct consequence of the



activity of plant roots. Obviously, some of these, such as many changes in ionic

concentrations and pH, are due simply to the uptake activity of the root. Although

changes in pH have received considerable attention, modifications of ionic concentrations certainly have to be considered as equally important and universally

widespread features of rhizosphere chemistry. Indeed, ionic concentrations and pH

are critical parameters that control many chemical reactions occumng at the

root-soil interface. Root-induced changes of these conditions will therefore influence the dynamics of many nutrients in the rhizosphere and ultimately their acquisition by plant roots. The uptake of nutrients thus operates as a major driving

force in nutrient acquisition. It should, however, no longer be solely considered as

the ultimate mechanism involved in plant nutrition. One has to bear in mind that

nutrient uptake has a major effect on the chemical conditions occumng in the rhizosphere, which will reciprocally determine the uptake activity of the root.

As evidenced mostly over the last decade, in addition to these interactions, other chemical processes, such as redox reactions, complexation, or enzymatic catalyses, can take place in the rhizosphere as a direct consequence of the exudation of

more or less specifically oriented metabolites produced by plant roots. The exudation of organic acids and enzymes, for instance, can contribute a significant proportion of the supply of major nutrients such as P to plant roots. Similarly, the exudation of phytosiderophores by roots of grass species plays a major role in the

acquisition of poorly mobile micronutrients such as Fe and many other metals. A

better understanding of these peculiar chemical processes occurring at the

root-soil interface is thus a prerequisite for more accurately predicting the nutrition needs of plants and the risks of undesirable micropollutants such as heavy metals entering the food chain.

Many of the aforementioned processes seem to be induced or stimulated in response to nutrient deficiencies. This suggests that they may be regarded as strategies of plant nutrition that evolved among higher plants to overcome adverse soil

chemical conditions (Marschner, 1995). Whether these processes can be considered as such, it should be borne in mind that the acquisition of mineral nutrients

not only relies on these diverse chemical processes but is largely influenced by (1)

the colonization of the soil by the root system and (2) the physical properties of

the intimate contact between the roots and the solid, liquid, and gaseous phases of

the soil. Considerable progress has been made in improving our knowledge of root

growth and rooting patterns (architecture of the root systems). In comparison, only

a limited amount of scientific data is relevant to the physical dimension of root-soil

interactions occurring in the rhizosphere. Thus, further investigations are needed

in this area.

In this chapter, chemical processes that occur in the rhizosphere as a direct consequence of root activity were addressed. However, other processes that significantly contribute to plant nutrition occur as a result of rhizosphere microflora. This

phenomenon can be regarded as an indirect effect of plant roots, since the activi-



ty of microorganisms in the rhizosphere is largely supported by root exudation of

C compounds. Although this “rhizosphere effect” has been studied over almost a

century, many questions remain, especially regarding its actual benefit for plant

nutrition and plant growth. For instance, rhizosphere microorganisms are likely to

rapidly degrade those exudates-such as organic anions, phytosiderophores, and

enzymes-that are supposed to assist the plant in acquiring some mineral nutrients. They also compete with plant roots for mineral nutrients. Rhizosphere microflora can thus have a detrimental effect on plant nutrition. The energetic cost of

rhizosphere microflora has been particularly addressed in the case of symbiotic

rhizosphere microorganisms such as N,-fixing bacteria and mycorrhizal fungi.

Nevertheless, mycorrhized plants often have a better P status than do nonmycorrhized plants, and over 95% of plant species are indeed mycorrhized. More interestingly, some species that are never mycorrhized, such as oilseed rape and white

lupin, among crops, and many members of the Proteaceae family, among wild

species (Harley and Harley, 1987; Brundrett and Abbott, 1991), have been reported as being some of the most efficient species for mobilizing soil P. This is attributed to their peculiar ability to excrete considerable amounts of protons and/or organic anions, such as citrate in particular. One may thus question whether these

root-induced chemical processes evolved in these species to compensate for the

lack of mycorrhizal support in P acquisition.Whatever the answer, the occurrence

of such plant species suggests that some rhizosphere characteristics may be worth

taking into account in plant breeding programs.

In today’s world, where conventional, intensive agricultural practices are being

challenged for both economic and environmental reasons, we should no longer

breed crops and pasture species that give a maximum yield under optimal growing conditions.From the plant-nutritionview point, this practice assumes that such

optimal conditions can be achieved with an adequate, and most often massive, use

of fertilizers. Sustainableagriculture, however, requires moderate consumption of

fertilizers. In this perspective, we should aim instead at selecting those species and

varieties that can most efficiently cope with a range of nonoptimal soil conditions.

A prerequisite to incorporating such considerations into our breeding programs is

a better understanding of the actual, combined effect on nutrient acquisition of the

various processes that occur in the rhizosphere. New experimental tools derived

from molecular biology, such as using mutants and genetic manipulations,will certainly help in ascertaining the relative contribution of the numerous mechanisms

that are involved. Moreover, using mathematical models of the combined phenomena involved in the process of mineral nutrient acquisition will also help us to

improve our understanding of plant nutrition. For this purpose, as pointed out by

Darrah (1993), a more integrative and quantitative approach of rhizosphere

processes is indeed required. This is a fundamentalprerequisite to managing plant

nutrition in agricultural, forested, and natural environments.




This chapter is dedicated to the memory of Professor Horst Marschner, who contributed much to our

understanding of the chemical processes involved in the rhizosphere. I also thank Professor R. J. Gilkes,

Dr. J. C. Arvieu, and Dr. B. Jaillard for their comments on an earlier version of this chapter.


Adams, M. A., and Pate, J. S. (1992). Availability of organic and inorganic forms of phosphorus to

lupins (Lupinus spp.). PIanr Soil 145, 107-1 13.

Ae, N., Arihara, J.. Okada, K..Yoshihara. T., and Johansen, C. (1990). Phosphorus uptake by pigeon

pea and its role in cropping systems of the Indian subcontinent. Science 248,477480.

Aguilar, S. A., and van Diest, A. (1981). Rock phosphate mobilization induced by the alkaline pattern

of legumes utilizing symbiotically fixed nitrogen. Pfant Soil 61,2742.

Ahmad, A. R., and Nye, P. H. (1990). Coupled diffusion and oxidation of ferrous iron in soils. 1. Kinetics of oxygenation of ferrous iron in soil suspension. J. Soil Sci. 41,395409.

Amann, C., and Amberger, A. (1989). Phosphorus efficiency of buckwheat (Fagopyrum esculentum).

2. Pjlanzenem. Bodenk. 152, 181-189.

Ando, T., Yoshida, S., and Nishiyama, I. (1983). Nature of oxidizing power of rice roots. Plant Soil 72,

57-7 1.

Armstrong. W. (1967). The use of polarography in the assay of oxygen diffusing from roots in anaerobic media. Physiol. Plant. 20,540-553.

Arvieu, J. C. (1980). RCactions des phosphates minkraux en milieu calcaire; constquence sur I’Ctat et

la solubilitk du phosphore. Sci. Sol 3, 179-190.

Asher, C. J., and Loneragan, J. F. (1967). Response of plants to phosphate concentration in solution

culture: I. Growth and phosphorus content. Soil Sci 103,225-233.

Bacha. R. E., and Hossner, L. R. (1977). Characteristics of coating formed on rice roots as affected by

iron and manganese additions. Soil Sci. Soc. Am. J. 41,931-935.

Baethgen, W. E.. and Alley, M. M. (1987). Nonexchangeable ammonium nitrogen contribution to plant

available nitrogen. Soil Sci. SOC.Am. J. 51, 110-1 15.

Barber, S. A. (1995). “Soil Nutrient Bioavailability: A Mechanistic Approach,” 2nd ed. John Wiley,

New York.

Barrow, N. J. (1983). On the reversibility of phosphate sorption by soils. J. Soil Sci. 34,751-758.

Barrow, N. J. (1984). Modelling the effect of pH on phosphate sorption by soils. J. Soil Sci. 35,


Begg, C. B. M., Kirk, G. 1. D., MacKenzie, A. E, and Neue, H.-U. (1994). Root-induced iron oxidation and pH changes in the lowland rice rhizosphere. New Phyfol. 128,469477.

Bekele, T., Cino, B. J., Ehlert, P. A. I., van der Maas, A. A,, and van Diest, A. (1983). An evaluation of

plant-borne factors promoting the solubilization of alkaline rock phosphates. Planr Soil 75,


Bertsch, P. M., and Thomas, G. W. (1985). Potassium status of temperate region soils. In “Potassium

in Agriculture” (R. D. Munson, ed.), pp. 131-162. American Soc. of Agron., Crop Sci. Soc.Am.

and Soil Sci. Soc.Am., Madison, W.

Bhat, K. K. S., and Nye. P. H. (1973). Diffusion of phosphate to plant roots in soil. I. Quantitative autoradiography of the depletion zone. Plunr Soil 38, 161-175.

Bhat, K. K. S., Nye, P. H., and Baldwin, J. P. (1976). Diffusion of phosphate to plant roots in soil. IV.



The concentration distance profile in the rhizosphere of roots with root hairs in a low-P soil. Planr

Soil 44,63-72.

Bienfait, H. F., Bino, R. J., van der Bliek, A. M., Duivenvoorden, J. F., and Fontdine, J. M. (1983).

Characterization of ferric reducing activity in roots of Fe-deficient Phaseolus vulgaris. Physiol.

Plant. 59, 196202.

Bolan, N. S. (1991). A critical review on the role of mycorrhizal fungi in the uptake of phosphorus by

plants. Plant Soil 134, 189-207.

Bolan, N. S., Naidu, R., Mahimairaja, S., and Baskaran, S. (1994). Influence of low-molecular-weight

organic acids on the solubilization of phosphates. Biol. Fer/. Soils 18,311-319.

Bosc, M. (1988). Enseignements fournis par des essais de longue durke sur la fumure phosphatke et

potassique. 3. Essai sur la fumure potassique. In “Phosphore et potassium dans les relations solplante: conskquences sur la fertilisation” (L. Gachon, ed.), pp. 403-466. Institut National de la

Recherche Agronomique, Paris.

Bowen, G. D. (1980). Coping with low nutrients. In “The Biology of Australian Plants” (J. S. Pate and

A. J. McComb, eds.), pp. 33-64. Univ. of Western Australia Press, Nedlands.

Breeze, V. G., Wild, A,. Hopper, M. J., and Jones, L. H. P. (1984). The uptake of phosphate by plants

from flowing nutrient solution. 11. Growth of Lolium perenne L. at constant phosphate concentrations. J. Exp. Bor. 35, 1210-1221.

Brewster, J. L., Bhat, K. K. S., and Nye, P. H. (1976). The possibility of predicting solute uptake and

plant growth response from independently measured soil and plant characteristics. V. The growth

and phosphorus uptake of rape in soil at a range of phosphorus concentrations and a comparison

of results with the predictions of a simulation model. Plant Soil 44,295-328.

Brown, J. C., and Ambler, J. E. (1973).“Reductants” released by roots of Fe-deficient soybeans. Agran.

J. 65,311-314.

Brundrett, M. C., and Abbott, L. K. (1991). Roots of jarrah forest plants. I. Mycorrhizal associations

of shrubs and herbaceous plants. Ausr. J. Bot. 39,445-457.

Bruand, A,, Cousin, I.. Nicoullaud, B., Duval, O., and BCgon, J. C. (1996). Backscattered electron scanning images of soil porosity for analysing soil compaction around roots. Soil Sci. SOC.Am. J . 60,


Chaney, R. L., Brown, J. C., andTiffin, L. 0. (1972). Obligatory reduction of ferric chelates in iron uptake by soybeans. Plant Physiol. 50,208-213.

Chen, C. C., Dixon, J. B., and Turner, F. T.(1980). Iron coatings on rice roots: Mineralogy and quantity influencing factors. Soil Sci. Soc. Am. J. 44,635-639.

Claassen, N., and Jungk, A. (1982). Kaliumdynamik im wurzelnahen Boden in Beziehung zur Kaliumaufnahme von Maispflanzen. 2. Pfanzenern. Bodenk. 145,s 13-525.

Clarkson, D. T. (1985). Factors affecting mineral nutrient acquisition by plants. Ann. Rev. Planr Physiol. 36,7771 15.

Curl, E. A., and Truelove, B. (1 986). “The Rhizosphere.” Springer-Verlag. Berlin-Heidelberg.

Darrah, P. R. (1991). Models of the rhizosphere. 1. Microbial population dynamics around a root releasing soluble and insoluble carbon. Planr Soil 133, 187-199.

Darrah,P. R. (1993). The rhizosphere and plant nutrition: Aquantitative approach. Plant Soil 159156,


Dexter, A. R. (1987). Compression of soil around roots. Plant Soil 97,401406.

Dinkelaker, B., and Marschner. H. (1992). In vivo demonstration of acid phosphatase activity in the

rhizosphere of soil-grown plants. Plunt Soil 144, 199-205.

Dinkelaker, B., Romheld, V., and Marschner, H. (1989). Citric acid excretion and precipitation of calcium citrate in the rhizosphere of white lupin (Lupinus albus L.). PIanr Cell Envi,: 12,285-292.

Dinkelaker, B., Hahn. G., and Marschner, H. (1993).Non-destructive methods for demonstrating chemical changes in the rhizosphere. LI. Application of methods. Plant Soil 155/156,73-76.

Dinkelaker, B., Hengeler, C., and Marschner, H. (1995). Distribution and function of proteoid roots

and other root clusters. Bor. Acra 108, 183-200.



Drew. M. C. (1988). Effect of flooding and oxygen deficiency on plant mineral nutrition. Adv. Plant

Nutr: 3, 115-159.

Drew, M. C., and Nye, P. H. (1969). The supply of nutrient ions by diffusion to plant roots in soil. 11.

The effect of root hairs on the uptake of potassium by roots of ryegrass (Lolium multiflorum).

Plant Soil 31,407424.

Epstein, E., and Hagen, C. E. (1952).A kinetic study of the absorption of alkali cations by barley roots.

Plant Physiol. 27,457474.

Fanning. D. S.. Keramidas. V. 2..and El-Desoky, M. A. (1989). Micas. In ”Minerals in Soil Environment,” 2nd ed. (J. B. Dixon and S. B. Weed, eds.), pp. 55 1-634. Soil Sci. Soc.Am., Madison, WI.

Fan; E., Vaidyanathan, L. V., and Nye, P. H. (1969). Measurement of ionic concentration gradients in

soil near roots. Soil Sci. 107, 385-9 1.

Fernandes Barros, 0. N., and Hinsinger, P. (1994). Basalt weathering as induced by the roots of higher plants. In “Proc. 15th ISSS Cong.” Acapulco.

Findenegg, G. R., and Nelemans, J. A. (1993). The effect of phytase on the availability of P from myoinositol hexaphosphate (phytate) for maize roots. Plant Soil 154, 189-196.

Fisher, H. M., and Stone, E. L. (1991). Iron oxidation at the surfaces of slash pine roots from saturated soil. Soil Sci. Soc. Am. J. 55, I 123- I 129.

Flessa, H., and Fischer, W. R. (1992). Plant-induced changes in the redox potentials of rice rhizospheres. Plant Soil 143,55-60.

Fohse, D., Claassen. N.. and Jungk, A. (1988). Phosphorus efficiency of plants. I. External and internal requirement and P uptake efficiency of different plant species. Plant Soil 110, 101-109.

Fox, R. L. ( I 98 I). External phosphorus requirements ofcrops. In “Chemistry in the Soil Environment.”

pp. 223-239. Am. SOC.Agron., Soil Sci. Soc.Am., Madison, WI.

Freeman. J. S.. and Rowell, D. L. (1981). The adsorption and precipitation of phosphate onto calcite.

J. Soil Sci. 32,75584.

Fried, M., and Shapiro, R.E. (1961). Soil plant relationships in ion uptake. Ann. Rev. Plant Physiol.


Gahoonia, T. S. (1993). Influence of root-induced pH on the solubility of soil aluminium in the rhizosphere. Plant Soil 149,289-291.

Gahoonia, T. S., and Nielsen, N. E. (1992). Control of pH at the soil-root interface. Planf Soil 140,


Gahoonia, T. S., Claassen, N., and Jungk, A. (1992a). Mobilization of phosphate in different soils by

ryegrass supplied with ammonium or nitrate. Plant Soil 140,241-248.

Gahoonia, T. S., Claassen, N., and Jungk, A. (1992b). Mobilization of residual phosphate of different

phosphate fertilizers in relation to pH in the rhizosphere of ryegrass. Ferr. Res. 33,229-237.

Gardner, W. K., Parberry, D. G., and Barber, D. A. ( I 982). The acquisition of phosphorus by Lupinus

albus L. I. Some characteristics of the soil/root interface. Plant Soil 68, 19-32.

Gardner, W. K., Barber, D. A,, and Parbeny, D. G. (1983). The acquisition of phosphorus by Lupinus

albus L. 111. The probable mechanism by which phosphorus movement in the soil/root interface

is enhanced. Plant Soil 70,107-124.

Gerke, J. (1994). Kinetics of soils phosphate desorption

ffected by citric acid. Z. Pjanzenern. Bodenk. 157, 17-22.

Gerke, J., Romer. W., and Jungk, A. (1994). The excretion of citric and malic acid by proteoid roots of

Lupinus albus L.: Effects on solubility of phosphate, iron, and aluminum in the proteoid rhizosphere in samples of an oxisol and a luvisol. 2. Pjanzenern. Bodenk. 157,289-294.

Godo, G. H.. and Reisenauer. H. M. (1980). Plant effects on soil manganese availability. Soil Sci. Soc.

Am. J. 44,993-995.

Green, M. S., and Etherington, J. R. (1977). Oxidation of ferrous iron by rice (Uryza sativa L.) roots:

A mechanism for waterlogging tolerance? J. Exp. Bot. 28,678-690.

Grierson, P. E (1992). Organic acids in the rhizosphere of Banksia integrifolia L. f. Plant Soil 144,




Grimal, J. Y., Delauney, A,, Frossard, E., and Morel, J. L. (1992). Utilisation de phosphore organique

par un mays cultivt en conditions stbriles. In “Proc. 4th Intl. IMPHOS Conf., pp. 629-63 I . Ghent,


Grimal, J. Y., Frossard, E., and Morel, J. L. (1995). Acquisition of phosphorus sorbed on goethite by

Zea mays. In “Soil Management in Sustainable Agriculture” (H. Cook and H. Lee, eds.). Wye College Press, London.

Grinsted, M. J., Hedley, M. J., White, R. E., and Nye, P. H. (1982). Plant-induced changes in the rhizosphere of rape (Brassica napus var. Emerald). 1. pH changes and the increase in P concentration in the soil solution. New Phyfol. 91, 19-29.

Hamza, M. A., and Aylmore, L. A. G. (1992). Soil solute concentration and water uptake by single lupin

and radish plant roots. I. Water extraction and solute accumulation. Plant Soil 145, 187-196.

Harley, J. L., and Harley, E. L. (1987).A check-list of mycorrhiza in the British flora. New Phyrol. 105,


Haussling, M.. and Marschner, H. (1989). Organic and inorganic soil phosphates and acid phosphatase

activity in the rhizosphere of 80-year-old Norway spruce (Picea abies (L.) Karst) trees. Biol. Fen.

Soils 8, 128-133.

Haussling, M., Leisen, E., Marschner, H., and Romheld, V. (1985). An improved method for non-destructive measurements of the pH at the root-soil interface (rhizosphere). J. Plant Physiol. 117,


Haynes. R. J.. (1990).Active ion uptake and maintenance of cation-anion balance: Acritical examination of their role in regulating rhizosphere pH. Plant Soil 126,247-264.

Hedley, M. J., Nye, P. H.,and White, R. E. (1982a). Plant-induced changes in the rhizosphere of rape

(Brassica napus var. Emerald) seedlings. II. Origin of the pH change. New Phyrol. 91,3144.

Hedley, M. J., White, R. E., and Nye, P. H. (1982b). Plant-induced changes in the rhizosphere of rape

(Brassica napus var. Emerald) seedlings. IU.Changes in L value, soil phosphate fractions, and

phosphatase activity. New Phyrol. 91,45-56.

Helal, H. M., and Sauerbeck, D. (1989). Carbon turnover in the rhizosphere. Z. Pjanzenem. Bodenk.


Hendriks, L., Claassen, N., and Jungk, A. (1981). Phosphatverarmung des wurzelnahen Bodens und

Phosphataufnahme von Mais und Raps. Z. Pflanzenern. Bodenk. 144,486-499.

Hiltner, L. (1904). Uber neuere Ehrfahrungen und Problem auf dem Gebiet der Bodenbakteriologie

unter besonderer Beriicksichtigung der Grundiingung und Brache. Arb. Drsch. Landwirr. Ges. 98,


Hinsinger, P. (1994). The acquisition of mineral nutrients by roots: Rhizosphere processes. In “Proc.

3rd Cong. European Soc. for Agron.” (M. Bonn and M. Sattin, eds.), pp. 428437. European SOC.

for Agron., Colmar, France.

Hinsinger, P., and Gilkes, R. J. (1995). Root-induced dissolution of phosphate rock in the rhizosphere

of lupins grown in alkaline soil. Ausr. J. Soil Res. 33,477489.

Hinsinger, P., and Gilkes, R. J. (1996). Mobilization of phosphate from phosphate rock and aluminasorbed phosphate by the roots of ryegrass and clover as related to rhizosphere pH. Eur: J. Soil Sci.


Hinsinger, P., and Gilkes, R. J. (1997). Dissolution of phosphate rock in the rhizosphere of five plant

species grown in an acid, P-fixing mineral substrate. Geoderma 75,231-249.

Hinsinger, P., and Jaillard, B. (1993). Root-induced release of interlayer potassium and vermiculitization of phlogopite as related to potassium depletion in the rhizosphere of ryegrass. J. Soil Sci. 44,


Hinsinger, P., Jaillard, B., and Dufey, J. E. (1992). Rapid weathering of a trioctahedral mica by the roots

of ryegrass. Soil Sci. SOC.Am. J. 56,977-982.

Hinsinger, P.,Elsass. F.,Jaillard, B., and Robert, M. (1993). Root-induced irreversible transformation

of a trioctahedral mica in the rhizosphere of rape. J. Soil Sci. 44,535-545.



Hoffland, E. (1992). Quantitative evaluation of the role of organic acid exudation in the mobilization

of rock phosphate by rape. Planr Soil 140,279-289.

Hoffland, E., Findenegg, G. R., and Nelemans, J. A. (1989). Solubilization of rock phosphate by rape.

11. Local root exudation of organic acids as a response to P-starvation. Plant Soil 113, 161-165.

Hoffland, E., Van den Boogaard, R.. Nelemans, J. A., and Findenegg, C. R. (1992). Biosynthesis and

root exudation of citric and malic acids in phosphate-starved rape plants. New Phyrol. 122,


Horst, W. J., Wagner, A., and Marschner, H. (1982). Mucilage protects root meristems from aluminium injury. 2.Pfanzenern. Bodenk. 105,435-444.

Hiibel, F., and Beck, E. (1993). In-situ determination of the P-relations around the primary root of maize

with respect to inorganic and phytate-P. Plant Soil 157, 1-9.

Hue, N. V., Craddock. G. R.. and Adams, F. (1986). Effect of organic acids on aluminum toxicity in

subsoils. Soil Sci. SOC.Am. J. 50,28-34.

Jaillard, B. (1984). Mise en evidence de la neogentse de sables calcaires sous I’influence des racines:

Incidence sur la granulometrie du sol. Agronomie 4,91-100.

Jaillard, B. (1985). Activitt! racinaire et rhizostructures en milieu carbonate. Pidologie 35, 297-3 13.

Jaillard, B. (1987). Les structures rhizomorphes calcaires: modtle de reorganisation des minkraux du

sol par les racines. Ph.D. thesis, Universitt! des Sciences et Techniques du Languedoc, Montpellier, France.

Jaillard, B., Guyon, A., and Maurin. A. F. (1991).Structure and composition of calcified roots, and their

identification in calcareous soils. Geodema 50, 197-210.

Jaillard. B., Ruiz, L., and Arvieu, J. C. (1996). pH mapping in transparent gel using color indicator

videodensitometry. Plant Soil 183,85-95.

Jakobsen, I., Abbott, L. K., and Robson, A. D. (1992). External hyphae of vesicular-arbuscular mycorrhizal fungi associated with Trifolium subrerruneum L. I . Spread of hyphae and phosphorus

inflow into roots. New Phytol. 120,371-380.

Jarvis, S . C., and Robson, A. D. (1983).The effects of nitrogen nutrition of plants on the development

of acidity in Western Australian soils. 1. Effects with subterranean clover grown under leaching

conditions. Aust. J. Agric. Res. 34,341-353.

Jones, D. L., and Darrah, P.R. (1994). Role of root derived organic acids in the mobilization of nutrients from the rhizosphere. Planr Soil 166,247-257.

Jones, D. L.. and Ddrrah, P.R. (1995). Influx and efflux of organic acids across the soil-root interface

of Zea mays L. and its implications in rhizosphere C flow. Plant Soil 173, 103-109.

Jones, D. L., Darrah, P. R.,and Kochian, L. V. (1996a). Critical evaluation of organic acid mediated

dissolution in the rhizosphere and its potential role in root iron uptake. Plunt Soil 180,5746.

Jones, D. L., Prabowo, A. M., and Kochian, L. V. (1996b).Kinetics of malate transport and decomposition in acid soils and isolated bacterial populations: The effect of microorganisms on root exudation of malate under Al stress. Plant Soil 182,239-247.

Jungk, A. (1996). Dynamics of nutrient movement at the soil-root interface. In “Plant Roots. The Hidden Half” 2nd ed. (Y.Waisel, A. Eshel, and U. Kafkafi, eds.), pp. 529-556. Marcel Dekker, New


Jungk, A,. and Claassen, N. (1986). Availability of phosphate and potassium as the result of interactions between root and soil in the rhizosphere. Z. Pfunzenern. Bodenk. 149,411-427.

Keerthisinghe, G., De Datta, S. K., and Mengel, K. (1985). Importance of exchangeable and nonexchangeable NH; in nitrogen nutrition of lowland rice. Soil Sci. 140, 194-201.

Khasawneh, F. E., and Doll, E. C. (1978).The use of phosphate rock for direct application to soils. Adv.

Agron. 30, 159-206.

Kinraide, T. B. (1991). Identity of the rhizotoxic aluminium species. Plant Soil 134, 167-178.

Kirk, G. J. D., and Nye, P. H. (1986). A simple model for predicting the rates of dissolution of sparingly soluble calcium phosphates in soil. I. The basic model. J. Soil Sci. 37,529-540.



Kirk, G. J. D., and Saleque, M. A. (1995). Solubilization of phosphate by rice plants growing in reduced soil: Prediction of the amount solubilized and the resultant increase in uptake. European J.

Soil Sci. 46,247-255.

Kodama, H., Nelson, S., Yang. F., and Kohyama, N. (1994).Mineralogy of rhizospheric and non-rhizospheric soils in corn fields. Clays Clay Min. 42,755-763.

Kraffczyk, I., Trolldenier, G., and Beringer, H. (1984). Soluble root exudates of maize: Influence of

potassium supply and rhizosphere microorganisms. Soil Biol. Biochem. 16,3 15-322.

Kraus, M., Fusseder. A., and Beck, E. (1987). In siru determination of the phosphate-gradient around

a root by radioautography of frozen soil sections. Plant Soil 97,407418,

Kuchenbuch, R., and Jungk. A. (1982).A method for determining concentration profiles at the soil-root

interface by thin slicing rhizospheric soil. Plant Soil 68,3914.

Kuchenbuch, R., and Jungk, A. (1984).Wirkung der Kaliumdiingung auf die Kaliumverfiigbarkeit in

der Rhizosph~evon Raps. Z. Pjanzenern. Bodenk. 147,435-448.

Lamont, B. (1982). Mechanisms for enhancing nutrient uptake in plants with particular reference to

mediterranean South Africa and Western Australia. Bot. Rev. 48,597-689.

Leprince, F., and Quiquampoix, H. (1996). Extracellularenzyme activity in soil: Effect of pH and ionic strength on the interaction with montmorillonite of two acid phosphatases secreted by the ectomycorrhizal fungus Heheloma cylindrosporum. Eur: J. Soil Sci. 47,5 1 1-522.

Lewis, D. G., and Quirk, J. P. (1967).Phosphate diffusion in soil and uptake by plants. 111. P”’-movement and uptake by plants as indicated by P’2-autoradiogrdphy. Plant Soil 27,445453,

Li, X.-L., George, E., and Marschner, H. (1991). Extension of the phosphorus depletion zone in VAmycorrhizal white clover in a calcareous soil. Plant Soil 136,4148.

Lindsay, W. L. (1954). Role of chelation in micronutrient availability. In “The Plant Root and Its Environment” (E. W. Carson, ed.), pp. 507-524. University Press of Virginia.

Lindsay, W. L. (1979). “Chemical Equilibria in Soils.” John Wiley and Sons, New York.

Lindsay, W. L., Vlek, P. L. G., and Chien, S. H. (1989). Phosphate minerals. In “Minerals in Soil Environment,” 2nd ed. (J. B. Dixon and S. B. Weed, eds.), pp. 1089-1130. Soil Sci. Soc. of Am.,

Madison, WI.

Lipton, D. S., Blanchar, R. W., and Blevins, D. G. (1987). Citrate, malate, and succinate concentration

in exudates from P-sufficient and P-stressed Medicago sariva L. seedlings. Plant Physiol. 85,

3 15-3 1 7.

Lorenz, S. E., Hamon, R. E.. and McGrath, S. P. (1994). Differences between soil solutions obtained

from rhizosphere and non-rhizosphere soils by water displacement and soil centrifugation. Eua J.

Soil Sci. 4 5 4 3 1-438.

Loss, S. P., Robson, A. D.. and Ritchie, G. S. P. (1993). H+/OH- excretion and nutrient uptake in upper and lower parts of lupin (Lupinus angustijolius L.) root systems. Annals Bor. 72,3 15-20.

Malzer, G. L., and Barber, S. A. (1975). Precipitation of calcium and strontium sulfates around plant

roots and its evaluation. Soil Sci. SOC.Amer: Pmc. 39,492495.

Mdrschner, H. (1990). Mobilization of mineral nutrients in the rhizosphere In “Proc. 1st Cong. European Soc. Agron.” (A. Scaife, ed.), Session 3, 0 00, pp. 1-11. European Soc. Agron., Colmar,


Marschner, H. (1995). “Mineral Nutrition of Higher Plants, 2nd ed. Academic Press. London.

Mdrschner, H., and Dell, B. (1994). Nutrient uptake in rnycorrhizal symbiosis. Plant Soil 159,89-102.

Marschner, H., and Romheld, V. (1983). In vivo measurement of root-induced pH changes at the

soil-root interface: Effect of plant species and nitrogen source. Z. Ppbnzenphysiol. 111,241-251.

Marschner, H.. Romheld, V., and Ossenberg-Neuhaus, H. (1982). Rapid method for measuring changes

in pH and reducing processes along roots of intact plants. 2.Pflanzenphysiol. 105,407416.

Marschner, H., Romheld, V., Horst, W. J., and Martin, P. (1986). Root-induced changes in the rhizosphere: Importance for the mineral nutrition of plants. Z. Pfanzenern. Bodenk. 149,441456.

Marschner, H., Romheld, V., and Kissel, M. (1987). Localization of phytosiderophore release and iron

uptake along intact barley roots. Physiol. Plant. 71, 157-162.



Marschner, H.,Treeby, M., and Romheld, V. (1989). Role of root-induced changes in the rhizosphere

for iron acquisition in higher plants. Z. P’anzenern. Bodenk. 152, 197-204.

Marschner, H., Hiussling, M.. and George, E. (1991). Ammonium and nitrate uptake rates and rhizosphere-pH in non-mycorrhizal roots of Norway spruce (Picea abies (L). Karst). Trees Strucr.

Funct. 5, 14-2 I .

McLachlan, K. D. (1980).Acid phosphatase activity of intact roots and phosphorus nutrition in plants.

I. Assay conditions and phosphatase activity. Ausr. J. Agric. Res. 3 1 , 4 2 9 4 0 .

Mench. M., and Fargues, S. (1995). Metal uptake by iron-efficient and inefficient oats. In “Iron Nutrition in Soils and Plants,” (J. Abadia, ed.), pp. 217-223. Kluwer Academic Publishers, Dordrecht,

the Netherlands.

Mench, M., and Martin, E. (1991). Mobilization of cadmium and other metals from two soils by root

exudates ofZeu m a w L., Nicotiana tahacum L., and Nicotiana rustica L. Plant Soil 132,187-196.

Mench, M., Morel, J. L.. and Guckert, A. (1987). Metal binding properties of high molecular weight

soluble exudates from maize (Zea mays L.) roots. Biol. Fen. Soils 3, 165-169.

Mengel. K., and Kirkby, E. A. (1987). “Principles of Plant Nutrition,” 4th ed. Intl. Potash Institute,

Bern, Switzerland.

Mengel, K., and Scherer, H.W. (198 I). Release of nonexchangeable (tixed) soil ammonium under field

conditions during the growing season. Soil Sci. 131.226232.

Mengel. K., Horn, D., and Tributh, H. (1990). Availability of interlayer ammonium as related to root

vicinity and mineral type. SoilSci. 149, 131-137.

Merckx, R., van Ginkel, J. H., Sinnaeve, J., and Cremers, A. (1986a). Plant-induced changes in the rhizosphere of maize and wheat. I. Production and turnover of root-derived material in the rhizosphere of maize and wheat. Plant Soil 96,85-93.

Merckx, R., van Ginkel, J. H.,Sinnaeve. J.. and Cremers, A. (1986b). Plant-induced changes in the rhizosphere of maize and wheat. 11. Complexation of cobalt, zinc, and manganese in the rhizosphere

of maize and wheal. Plant Soil 96,95-107.

Moorby, H., White, R. E., and Nye, P. H. (1988). The influence of phosphate nutrition on H ion efflux

from the roots of young rape plants. Plant Soil 105,247-56.

Morel, J. L., Mench, M., and Guckert,A. (1986).Measurement of PbZ+,Cu2+,and Cd2+ binding with

mucilage exudates from maize (&a mays L.) roots. B i d . Fert. Soils 2,29-34.

Murrrnann. R. P., and Peech. M. (1969). Effect of pH on labile and soluble phosphate in soils. Soil Sci.

Soc. Am. Proc. 33, 205-2 10.

Nagarajah, S., Posner. A. M., and Quirk, J. P. (1970). Competitive adsorptions of phosphate with polygalacturonate and other organic anions on kaolinite and oxide surfaces. Nature 228, 83-84.

Niebes, J.-F., Dufey, J. E., Jaillard, B., and Hinsinger, P. ( I 993). Release of nonexchangeable potassium from different size fractions of two highly K-fertilized soils in the rhizosphere of rape (Brassicu napus cv. Drakkar). Plant Soil 155/156,403406.

Nye, P. H.(1981). Changes of pH across the rhizosphere induced by roots. Plant Soil 61,7-26.

Nye, P. H.(1983). The diffusion of two interacting solutes in soil. J. Soil Sci. 34,677-691.

Nye, P. H.(1986). Acid-base changes in the rhizosphere. Adv. Plant NutK 2, 129-153.

Oertli, J. J., and Opoku, A. A. (1974). Interaction of potassium in the availability and uptake of iron

from ferric hydroxide. Soil Sci. Soc. Am. Proc. 38,451454.

Owusu-Bennoah, E., and Wild, A. (1979). Autoradiography of the depletion zone of phosphate around

onion roots in the presence of vesicular-arbuscular mycorrhiza. New Phytol. 82, 133-140.

Paauw, E van der. ( I 97 I ). An effective water extraction method for the determination of plant available soil phosphorus. Plant Soil 3 4 , 4 6 7 4 8 I .

Parfitt, R. L. (1978). Anion adsorption by soils and soil materials. Adv. Agron. 30, 1-50.

Parlitt, R. L. ( 1979).The availability of Pfrom phosphate-goethite bridging complexes. Desorption and

uptake by ryegrass. Plant Soil 5 3 , 5 5 4 5 .

Pellet, D. M., Grunes, D. L., and Kochian, L. V. (1995). Organic acid exudation as an aluminum-tolerance mechanism in maize (Zeu movs L.). Plunta 196,788-795.



Petersen, W.. and Bottger, M. (1991). Contribution of organic acids to the acidification of the rhizosphere of maize seedlings. Plant Soil 132, 159-163.

Qutmener. J. (1986). Important factors in potassium balance sheets. In “Nutrient Balances and the

Need for Potassium. Proc. 13th IPI Cong.,” pp. 41-72. Intl. Potash Institute, Bern, Switzerland.

Quiquampoix, H.. Abadie, J., Baron, M. H.,Leprince, F.. Matumoto-Pintro, P. T., Ratcliffe, R. G., and

Staunton, S. (1995). Mechanisms and consequences of protein adsorption on soil mineral surfaces.

In “Proteins at Interfaces 11. ACS Symposium Series 602” (T. A. Horbett and J. L. Brash, eds.),

pp. 321-333. Am. Chem. Soc.,Washington, D.C.

Quispel, A. (1983). Dinitrogen-fixing symbioses with legumes, non-legumes angiosperms, and associative symbioses. In “Inorganic Plant Nutrition. Encyclopedia of Plant Physiology.” (A. Lauchli and R. L. Bieleski, eds.), New Series, Vol. 15A, pp. 286-329. Springer-Verlag. Berlin.

Riley, D., and Barber, S. A. (I97 I). Effect of ammonium and nitrate fertilization on phosphorus uptake

as related to root-induced pH changes at the root-soil interface. Soil Sci. SOC. Am. froc. 35,


Robert, M., and Berthelin. J. (1986). Role of biological and biochemical factors in soil mineral weathering. In “Interactions of Soil Minerals with Natural Organics and Microbes,” (P.M. Huang and

M. Schnitzer, eds.), Special publication 17, pp. 453-495. Soil Sci. Soc.Am., Madison, W1.

Romheld, V. (1986). pH-Veranderungen in der Rhizosphiire verschiedener Kulturpflanzenarten in Abhangigkeit vom Nhstoffangebot. Potash Rev. 55, 1-8.

Romheld, V. (1991). The role of phytosiderophores in acquisition of iron and other micronutrients in

graminaceous species: An ecological approach. Plant Soil 130, 127-1 34.

Romheld, V.. and Marschner, H. (1983). Mechanism of iron uptake by peanut plants. I. Fe(II1) reduction, chelate splitting, and release of phenolics. Plant Physiol. 71,949-954.

Romheld, V., and Marschner. H.(1986a). Mobilization of iron in the rhizosphere of different plant

species. Adv. Plant Nutr: 2, 155-204.

Romheld, V.. and Marschner, H. (1986b). Evidence for a specific uptake system for iron phytosiderophores in roots of grasses. Plant Physiol. 80, 175-180.

Romheld, V., and Marschner, H. (1990). Genotypical differences among graminaceous species in release of phytosiderophores and uptake of iron phytosiderophores. Plant Soil 123, 147-153.

Romheld, V., Miiller, C., and Marschner, H. (1984). Localization and capacity of proton pumps in roots

of intact sunflower plants. Plant Physiol. 76,603-606.

Ruiz, L. (1992). Mobilisation du phosphore des apatites dans la rhizosphbre. Rble de I’excrttion de

protons par le racines. Ph.D. thesis, Universitt Montpellier II, France.

Ruiz, L.. and Arvieu, J. C. (1992). Solubilization of phosphate minerals by plant roots. In “Proc. 4th

Intl. IMPHOS Conf.” Ghent, Belgium.

Saleque, M. A., and Kirk, G. J. D. (1995). Root-induced solubilization of phosphate in the rhizosphere

of lowland rice. New Phytol. 129,325-336.

Scherer, H. W., and Ahrens, G. (1996). Depletion of non-exchangeable NH4-N in the soil-root interface in relation to clay mineral composition and plant species. Eur: J. Agron. 5, 1-7.

Schwertmann, U. (1991). Solubility and dissolution of iron oxides. Plant Soil 130, 1-25.

Sinha, B. K., and Singh, N. T. (1974). Effect of transpiration rate on salt accumulation around corn

roots in a saline soil. Agron. J. 66,557-560.

Sinha, B. K., and Singh, N. T. (1976). Chloride accumulation near corn root under different transpiration, soil moisture, and soil salinity regimes. Agron. J. 68,346348.

Sparks, D. L. (1987). Potassium dynamics in soils. Adv. Soil Sci. 6,1-63.

Staunton, S., and Leprince, F. (1996). Effect of pH and some organic anions on the solubility of soil

phosphate: Implications for P bioavailability. Eur: J. Soil Sci. 47,231-239.

Steffens, D. ( 1987). Einfluss langjahriger Diingung mit verschiedenen Phosphatdungerformen aud die

Phosphatverfiigbarkeit in der Rhizosphiire von Raps. Z. fjanzenern. Bodenk. 150,75-80.

Stirzaker, R. J., and Passioura, J. B. (1996). The water relations of the root-soil interface. Plant Cell

Envi,: 19,201-208.

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