3.3.3. Belowground competition for water and nutrientsBelowground competition occurs when plants decrease thegrowth, survival or fecundity of neighbours by reducing availablesoil resources. Contrary to aboveground competition,which primarily involves a single resource, light, plants competefor a broad range of soil resources including water andat least 20 mineral nutrients that differ in molecular size, valence,oxidation state and mobility within the soil (Casper andJackson, 1997). The components of the mixture may be complementaryin a spatial sense by exploiting different layers ofthe soil with their root systems. Components of a mixture maycomplement each other nutritionally (different needs in quantities,preferential use of different chemical forms). Mixturesof leguminous and non-leguminous species are well known inthat regard, and provide repeatable examples of overyieldingdue to nutritional complementation. To study those interactions,three aspects have to be taken into account that addressthe resource supply to the roots, the characteristics of the rootsystem, and the demand for water and carbon allocation, respectively(Fig. 5):(i) Resource supply to the roots involves four main processes:the distribution of resources in the soil and theiravailability, which depends on soil biophysical and chemicalproperties, interception by the roots (<10%), massflow, which affects water and mobile nutrients such asNO−3 , and diffusion, which affects nutrients such as P andK.(ii) Root system characteristics include morphological plasticity– root location in time (Caldwell and Richards,1986) and space (de Willingen and Van Noordwijk,1987), investment in root biomass, root length or surface– and physiological plasticity – rate of resource uptake inrelation to enzyme functioning.(iii) The demand for water. Water distribution depends on thepartitioning of evaporative demand between the species’components, and on soil evaporation (Ozier-Lafontaineet al., 1997, 1998).An analysis of the belowground processes and resource useby plants presents tremendous challenges as there are still generalmethodological difficulties despite the advances made intechniques and equipment design. For example, roots of thecomponent plants can intermingle (Gregory and Reddy, 1982),making the task of separating the respective root systems verycumbersome. Staining techniques generally fail to distinguishone root system from another. Other possibilities, such as54 E. Malézieux et al.isotopic discrimination of 13C between C3 legumes and C4cereals are efficient but require special equipment (Wong andOsmond, 1991; Lichtfouse, 1997).
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