Root function and overall plant performance can beneﬁt conspicuously from processes in the rhizosphere. Infection by rhizobia (Section 4.4) and mycorrhizal fungi (Section 4.3) improve the nutritional status of many species. Rhizobial strains have even been used to manipulate rhizosphere biology. A signiﬁcant proportion of photoassimilate is used to support a rhizosphere, reflecting the high cost of microbial activity and polymer exudation. This pattern is repeated in many species with up to 20% of plant carbon consistently lost by roots, however, this value can vary substantially with the biotic and abiotic conditions. Relative rates of microbial and root respiration are almost impossible to estimate in roots growing in undisturbed soils because of the intimacy of roots and microbes. In addition to consuming large amounts of plant carbon, some microbes can produce phytotoxins, which can impose further restrictions on root function. Some microbes also contribute to nutrient depletion in the rhizosphere, for example by converting usable forms of nitrogen, i.e. nitrate or ammonium, into unusable forms like N2.
Mechanisms describing how a rhizosphere beneﬁts its host are even more elusive because of the diversity of reactions in such a small space. Chelation is identiﬁed as a major influence on nutrient acquisition and might also help ameliorate ion toxicities. Physical properties of the rhizosphere are even less well understood, with questions such as root lubrication, root–mucilage shrinkage and interfacial water transport not yet resolved. Physical properties of mucilage do not suggest it is an ideal lubricant. Whether the dynamic properties of a rhizosphere bring constant beneﬁts to a plant or simply passively coexist with growing roots remains a critical question.
One demonstrated benefit of the rhizosphere microbiome is the protection of the plant from diseases. Several mechanisms have been suggested for this effect (Bakker et al. 2013; Berendsen et al. 2012): Disease suppressiveness, the ability of the microbial inhabits of the rhizosphere to suppress the infection of plants by soil-swelling pathogens, has been ascribed to the production of antimicrobial substances by bacteria, to competition between beneficial and pathogenic microbes and to the induction of systemic resistance by beneficial bacteria. An intriguing example is the colonization of plants by pathogens, which can lead to changes in the exudation of organic acids that then attract beneficial bacteria that induce systemic resistance in the plant, reducing pathogen infection. The induction of systemic resistance to pathogens can also be triggered by specific signaling molecules of bacteria, quorum sensing signals, which bacteria used to ‘talk’ to each other to coordinate multicellular-like behaviours of bacterial colonies. Perception of quorum sensing signals from rhizosphere bacteria by plants can increase systemic resistance to pathogens in the shoot, and can also enhance symbiosis with nitrogen-fixing bacteria. Quorum sensing signal perception also triggers the production of so called quorum sensing mimic compounds – signals that interfere with bacterial communication in the rhizosphere (Teplitski et al. 2011). While we still need to identify most of the signals, signal mimics and exudate components in the interaction of roots with their microbiome, it is clear that plants actively create the rhizosphere, and that this is likely to benefit the plant in its environment.