The rhizosphere encompasses the millimeters of soil surrounding a plant root where complex biological and ecological processes occur. This review describes recent advances in elucidating the role of root exudates in interactions between plant roots and other plants, microbes, and nematodes present in the rhizosphere. Evidence indicating that root exudates may take part in the signaling events that initiate the execution of these interactions is also presented. Various positive and negative plant-plant and plant-microbe interactions are highlighted and described from the molecular to the ecosystem scale. Furthermore, methodologies to address these interactions under laboratory conditions are presented.

Here we present evidence that Centaurea maculosa (spotted knapweed), an invasive species in the western United States, displaces native plant species by exuding the phytotoxin (-)-catechin from its roots. Our results show inhibition of native species' growth and germination in field soils at natural concentrations of (-)-catechin. In susceptible species such as Arabidopsis thaliana, the allelochemical triggers a wave of reactive oxygen species (ROS) initiated at the root meristem, which leads to a Ca2+ signaling cascade triggering genome-wide changes in gene expression and, ultimately, death of the root system. Our results support a "novel weapons hypothesis" for invasive success.

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Relatively little is known about the exact mechanisms used by Bacillus subtilis in its behavior as a biocontrol agent on plants.Here, we report the development of a sensitive plant infection model demonstrating that the bacterial pathogen Pseudomonas syringae pv tomato DC3000 is capable of infecting Arabidopsis roots both in vitro and in soil. Using this infection model, we demonstrated the biocontrol ability of a wild-type B. subtilis strain 6051 against P. syringae. Arabidopsis root surfaces treated with B. subtilis were analyzed with confocal scanning laser microscopy to reveal a three-dimensional B. subtilis biofilm. It is known that formation of biofilms by B. subtilis is a complex process that includes secretion of surfactin, a lipopeptide antimicrobial agent. To determine the role of surfactin in biocontrol by B. subtilis, we tested a mutant strain, M1, with a deletion in a surfactin synthase gene and, thus, deficient in surfactin production. B. subtilis M1 was ineffective as a biocontrol agent against P. syringae infectivity in Arabidopsis and also failed to form robust biofilms on either roots or inert surfaces. The antibacterial activity of surfactin against P. syringae was determined in both broth and agar cultures and also by live-dead staining methods. Although the minimum inhibitory concentrations determined were relatively high (25 g mL Ϫ1 ), the levels of the lipopeptide in roots colonized by B. subtilis are likely to be sufficient to kill P. syringae. Our results collectively indicate that upon root colonization, B. subtilis 6051 forms a stable, extensive biofilm and secretes surfactin, which act together to protect plants against attack by pathogenic bacteria.Beneficial plant rhizobacteria (PR) are associated with the surfaces of plant roots and may increase plant yield by mechanisms that impart improved mineral nutrient uptake, disease suppression, or phytohormone production (Kloepper et al., 1991;Lutenberg et al., 1991;Costacurta and Vanderleyden, 1995;Defago and Keel, 1995). An important trait of PR is their ability to effectively colonize the rhizosphere and maintain a stable relationship with the surface of plant roots (Lutenberg and Dekkers, 1999). PR may also interact with a variety of soil microorganisms that are normally present in the rhizosphere, in some cases acting as a biocontrol agent against pathogenic bacteria (Pinton et al., 2001). Interestingly, poor root colonization by PR may result in decreased biocontrol activity (Schippers et al., 1987). One beneficial rhizobacterium is Bacillus subtilis, which is ubiquitous in soil, can promote plant growth, protect against fungal pathogen attack ( Utkhede and Smith, 1992;Asaka and Shoda, 1996;Emmert and Handelsman, 1999), and play a role in the degradation of organic polymers in the soil (Emmert and Handelsman, 1999). Among the first successful biocontrol agents used against insects and pathogens were members of the genus Bacillus (Powell and Jutsum, 1993). Commercial strains of B. subtilis have been marketed as biocontrol agents for f...

Beneficial soil bacteria confer immunity against a wide range of foliar diseases by activating plant defenses, thereby reducing a plant's susceptibility to pathogen attack. Although bacterial signals have been identified that activate these plant defenses, plant metabolites that elicit rhizobacterial responses have not been demonstrated. Here, we provide biochemical evidence that the tricarboxylic acid cycle intermediate L-malic acid (MA) secreted from roots of Arabidopsis (Arabidopsis thaliana) selectively signals and recruits the beneficial rhizobacterium Bacillus subtilis FB17 in a dose-dependent manner. Root secretions of L-MA are induced by the foliar pathogen Pseudomonas syringae pv tomato (Pst DC3000) and elevated levels of L-MA promote binding and biofilm formation of FB17 on Arabidopsis roots. The demonstration that roots selectively secrete L-MA and effectively signal beneficial rhizobacteria establishes a regulatory role of root metabolites in recruitment of beneficial microbes, as well as underscores the breadth and sophistication of plant-microbial interactions.

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Pseudomonas aeruginosa is an opportunistic human pathogen capable of forming a biofilm under physiological conditions that contributes to its persistence despite long-term treatment with antibiotics. Here, we report that pathogenic P. aeruginosa strains PAO1 and PA14 are capable of infecting the roots of Arabidopsis and sweet basil (Ocimum basilicum), in vitro and in the soil, and are capable of causing plant mortality 7 d postinoculation. Before plant mortality, PAO1 and PA14 colonize the roots of Arabidopsis and sweet basil and form a biofilm as observed by scanning electron microscopy, phase contrast microscopy, and confocal scanning laser microscopy. Upon P. aeruginosa infection, sweet basil roots secrete rosmarinic acid (RA), a multifunctional caffeic acid ester that exhibits in vitro antibacterial activity against planktonic cells of both P. aeruginosa strains with a minimum inhibitory concentration of 3 g mL Ϫ1 . However, in our studies RA did not attain minimum inhibitory concentration levels in sweet basil's root exudates before P. aeruginosa formed a biofilm that resisted the microbicidal effects of RA and ultimately caused plant mortality. We further demonstrated that P. aeruginosa biofilms were resistant to RA treatment under in vivo and in vitro conditions. In contrast, induction of RA secretion by sweet basil roots and exogenous supplementation of Arabidopsis root exudates with RA before infection conferred resistance to P. aeruginosa. Under the latter conditions, confocal scanning laser microscopy revealed large clusters of dead P. aeruginosa on the root surface of Arabidopsis and sweet basil, and biofilm formation was not observed. Studies with quorum-sensing mutants PAO210 (⌬rhlI), PAO214 (⌬lasI), and PAO216 (⌬lasI ⌬rhlI) demonstrated that all of the strains were pathogenic to Arabidopsis, which does not naturally secrete RA as a root exudate. However, PAO214 was the only pathogenic strain toward sweet basil, and PAO214 biofilm appeared comparable with biofilms formed by wild-type strains of P. aeruginosa. Our results collectively suggest that upon root colonization, P. aeruginosa forms a biofilm that confers resistance against root-secreted antibiotics.Pseudomonas aeruginosa, a gram-negative bacterium commonly isolated from soil and water, is renowned for its nutritional and ecological versatility. As an opportunistic human pathogen, P. aeruginosa is a common cause of nosocomial infections and is responsible for persistent infections in immunocompromised individuals and for the chronic lung infections of patients with cystic fibrosis (Govan and Deretic, 1996). P. aeruginosa is also capable of causing serious infections in nonmammalian host species such as insects (Jander et al., 2000), nematodes (Mahajan-Miklos et al., 1999), and plants (Rahme et al., 1995;Silo-Suh et al., 2002). The effectiveness of this organism in causing infection is likely due to a suite of well-regulated virulence factors and defense mechanisms such as multidrug resistance pumps (Chuanchuen et al., 2001) and biofilm forma...

Though recent work has demonstrated that plants can recognize species, kin versus strangers, and self/non-self roots, no mechanism for identity recognition in plants has yet been found. Here we examined the role of soluble chemicals in signaling among roots. Utilizing Arabidopsis thaliana, we exposed young seedlings to liquid media containing exudates from siblings, strangers (non-siblings), or only their own exudates. In one experiment, root secretions were inhibited by sodium orthovanadate and root length and number of lateral roots were measured. In a second experiment, responses to siblings, strangers, and their own exudates were measured for several accessions (genotypes), and the traits of length of the longest lateral root and hypocotyl length were also measured. The exposure of plants to the root exudates of strangers induced greater lateral root formation than exposure of plants to sibling exudates. Stranger recognition was abolished upon treatment with the secretion inhibitor. In one experiment, plants exposed to sibling or stranger exudates have shorter roots than plants only exposed to their own exudates. This self/non-self recognition response was not affected by the secretion inhibitor. The results demonstrate that that kin recognition and self/non-self are two separate identity recognition systems involving soluble chemicals. Kin recognition requires active secretion by roots.