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4.2.1 - The rhizosphere

The rhizosphere is the narrow zone of soil surrounding plant roots that is characterised by root exudation and an abundance of micro-oganisms which can be beneficial or harmful to plants, or have no effect on root growth and function. These microbes are saprophytic, pathogenic or symbiotic bacteria and fungi, including rhizobia forming nodules and arbuscular mycorrhizal fungi (Figure 4.14).


Figure 4.14 The rhizosphere is the narrow zone of soil surrounding plant roots that is characterised by root exudation and an abundance of saprophytic, pathogenic and symbiotic bacteria and fungi. These include rhizobia that form nodules, and arbuscular mycorrhizal fungi (AMF). The rhizoplane describes the root surface in contact with the soil. Root cap and root border cells near the root tip provide lubrication as the root expands into the soil. (Reproduced by permission from Macmillan Publishers Ltd from L. Philippot et al. Nature Rev Microbiol 11: 789-799, 2013)

Many root phenomena suggest specific roles for the rhizosphere. For example, roots have long been thought to find a relatively frictionless path through soils because of exudation of organic substances and cell sloughing, but the chemical and physical processes that underpin this phenomenon are still quite unclear (McKenzie et al. 2012). Production of new roots around local zones of enrichment (Section 4.1) is made far more effective through rhizosphere activity associated with these young roots. Phosphate availability is particularly likely to be improved by the presence of a rhizosphere. Potential mechanisms will be discussed below.

Enhancement of root growth under conditions which favour high root:shoot ratios and the attendant rhizosphere surrounding those roots (rhizosheath) require a substantial input of organic carbon from shoots. Some is used in structural roles, while roots and microbes also require large amounts of carbon to sustain respiration. Even in plants growing in nutrient-adequate, moist soils, 30–60% of net photosynthate finds its way to roots (Marschner 1995). Carbon allocation to roots can be even greater in poor soils or during drought. The rhizosphere accounts for a large amount of the root carbon consumption (Jones et al. 2009). Barber and Martin (1976) showed that 7–13% of net photosynthate was released by wheat roots over three weeks under sterile conditions while 18–25% was released when roots were not sterile. This difference might be considered carbon released because of microbially-induced demand in the rhizosphere, and therefore made unavailable for plant growth.


Figure 4.15 Concentration of Enterobacter cloacae (RP8) around wheat roots when the bacterium was introduced by inoculating seeds (circles) or soil (triangles). Uninoculated controls are shown as diamonds. Approximately 3 mm of the soil around roots supports an elevated bacterial population (A.F. Dijkstra et al. Soil Biol Biochem 19: 351-352, 1987)

Rhizosphere chemistry and physics differ from the adjacent soil matrix and root tissues. Gradients in solutes, water and gases combine with microbial activity to produce a unique compartment through which roots perceive bulk soil. This zone of influence typically extends not more than 3 mm from the root axis (Figure 4.15), partly due to the low diffusion coefficients of most solutes that move through the rhizosphere (10–12 to 10–15 m2 s–1 for ions such as orthophosphates). Even a relatively mobile ion such as nitrate, with a diffusion coefficient (D) of around 10–9 m2 s–1 in soil solution, diffuses through about 1 cm of soil in a day. Because the time required (t) for diffusion of ions is a function of the square of distance traversed (l), where t = l2/D a nitrate ion would take four days to travel 2 cm, nine days to travel 3 cm and so on. Similarly, organic carbon diffuses away from roots only slowly, sustaining a microbial population as it is consumed in the rhizosphere.

Roots advancing through soil perceive a wide range of chemical and biological environments: a rhizosphere simultaneously fulfils buffering, extraction and defence roles allowing roots to exploit soils. A rhizosphere is thus a dynamic space, responding to biological and environmental conditions and often improving acquisition of soil resources. New roots develop an active rhizosphere which matures rapidly as the root axis differentiates.