Sugars constitute a significant fraction of exudates, and are a main carbon source for microbes. Interestingly, many more sugar uptake than release systems have been described. Sugar Transport Proteins utilize high extracellular proton levels to import sugars, and mutation of STPs leads to higher external sugar levels. Sugars Will Eventually Be Exported Transporters are sugar uniporters, and all root expressed members localize to the vacuole. Due to an alteration of root sugar homeostasis, SWEET mutant plants exhibited higher sugar export from roots compared with wild type plants, and were more susceptible to disease. Intriguingly, no transporters directly exporting sugars into the rhizosphere have been characterized so far, and it is debated whether sugar exudation is a transport driven process at all. Potential evidence for passive sugar efflux was supported by the observation of higher sucrose concentrations around young, permeable root tissue than around older, less permeable root tissue. However, because sugars are synthesized in leaves, they still need to be unloaded either from phloem or from root cells to be exuded into the rhizosphere, a process likely depending on transporters due to the hydrophilic nature of sugars. A further indication of the presence of elusive transporters is the differential sugar exudation in various environments, as shown, for example, for maize grown in potassium , phosphate , or iron deficient conditions.Sugar alcohols are imported by secondary active proteins with broad substrate specificity , whereas the modes of export are enigmatic. Sugar phosphates are involved in intracellular carbohydrate metabolism,hydroponic vertical farming and plastid localized sugar–phosphate co transporters have been reported in several species.
Although sugar phosphates are detected in exudates, neither import nor export mechanisms are currently characterized.Amino acids are recognized by microbial chemoreceptors crucial for the early steps of root colonization, making amino acids an important fraction of exudates. Modulation of amino acid transport could be either a means of communication with microbes, or a response to microbial presence. Amino acid uptake is mediated by several transporter families with broad substrate specificity. Amino acid exudation is affected by several transporters expressed in vascular tissue: mutation of phloem localized UmamiTs resulted in lower amino acid exudation, whereas mutation of xylem localized Glutamine Dumpers caused increased exudation. Although no plasma membrane localized amino acid exporters have been characterized so far, several lines of evidence suggest their presence. First, higher tryptophan exudation from older root zones than younger parts suggests the involvement of transport proteins in exudation, due to the fully formed Casparian strips and thick cell walls in mature root parts interfering with diffusion. Second, concentration differences between amino acids in root exudates and root extracts are not the same for all the amino acids, suggesting the selective transport of at least some amino acids. Third, various transporter families exhibit bidirectional amino acid transport characteristics in heterologous systems , and could be involved in amino acid exudation.Organic acids constitute a large fraction of exudates, and are microbial nutrients. No importers have been characterized so far, but the release of malate and citrate by Aluminium Activated Malate Transporters and Multi drug and Toxic Compound Extrusion families are among the few well understood examples of transporters involved in exudation .
Activity of members of both families is often modulated by metal ions and microbes . Uncharacterized ALMT and MATE family members are primary candidates for exporters of other organic acids due to their similarity to already characterized members, their plasma membrane localization, and their function as proton antiporters.Nucleotides are imported by secondary active transporters, but their exudation remains elusive. It is well established that extracellular ATP has a signaling function, and ABC transporters were proposed to mediate cellular export. Peptide uptake is transporter mediated in heterologous systems, and a role of ABC transporters in peptide exudation has been suggested .Fatty acid transport is necessary for mycorrhizal symbiosis: mycorrhizal fungi depended on their hosts for the synthesis of certain fatty acids, and the current model includes transport of lipids by ABCG proteins in the symbiotic membrane. One ABCG member, STR, was previously shown to be required for mycorrhization. Interestingly, arabidopsis ABCG transporters were similarly shown to export fatty acids for cutin synthesis in above ground tissues . Lipid transport was required not only for symbiotic interactions, but also for pathogen colonization. Fatty acids are detected in root exudates , but the mode of lipid exudation into the rhizosphere has yet to be discovered. A role in lipid exudation could be envisioned for root expressed ABCG members .Secondary metabolites are ubiquitous in root exudates, and ABC transporters are likely candidates for specialized metabolite transport into the rhizosphere. A distinct exudation profile was described for seven ABC mutants, and one mutant line displayed an altered microbial community. Although the causal metabolites could not be identified, the authors noted transport of the same compound by various proteins, and possible broad substrate specificity for some transporters. In a later study, exudates of arabidopsis ABCG37/PDR9 mutant lines were found to be deficient in several phenylpropanoids.
Arabidopsis PDR9 was previously characterized as auxin precursor transporter, which suggests a broad substrate specificity for PDR9. Interestingly, a PDR9 homolog was highly expressed in cluster roots of white lupin devoid of phosphate, illustrating PDR9 involvement in response to various abiotic stresses. These studies illustrate the potential for the discovery of novel transporter functions in the ABC family, an excellent target for future studies investigating root exudation. In addition, MATE proteins transport secondary metabolites into the vacuole,vertical hydroponic garden and plasma membrane localized members could also be involved in secondary metabolite exudation. In summary, more transport proteins involved in metabolite import into roots than in export from roots have been reported so far . The characterization of additional transport families involved in exudation will enable the generation of mutant lines that are devoid of the exudation of specific metabolites. Such lines could be used to investigate the correlation of exudation profiles and microbial communities.Plant derived transporters and exometabolites are intrinsic to plant–mycorrhizal and rhizobial symbioses . We speculate that, although there is paucity of evidence, plants analogously select for a beneficial rhizobiome. Given that plants evolved in the presence of microbes, a subset of which benefits plant growth, we hypothesize that, over millennia, plant exudation via active transport processes evolved with the substrate specificity of plant associated bacteria. In Box 2, we discuss exudates and other steps involved in root microbiome assembly, analogously to the establishment of plant–mycorrhizal and rhizobial symbioses. However, intense future research is needed to reveal the precise mechanisms governing plant microbiome assembly, and the possible beneficial functions of the microbial community. The major mechanisms by which plants are thought to modulate microbial interactions currently include: modulation of their exudate profiles ; root morphology ; and regulation of immune system activities . In turn, mechanisms for successful rhizosphere colonization by soil microbes require that they: are metabolically active ; sense the plant ; move towards the root and successfully compete with other microbes for root niches . In addition, for successful colonization of the rhizoplane or root tissue, microbes must be able to attach to the surface or enter root tissue . Despite apparent parallels between plant microbiomes and the aforementioned symbioses, plant microbiomes have some specific characteristics. First, microbiomes are detected in all environmental conditions, whereas mycorrhizal and rhizobial symbioses are induced in specific circumstances. Second, microbiomes occur on various tissues, whereas rhizobia and mycorrhiza interface with roots only. Third, microbiomes comprise many members, whereas the aforementioned symbioses persist between two predominant partners. Fourth, although most members of the microbiome originate from the environment similar to rhizobia and mycorrhiza, there is evidence that some endophytes may be vertically transmitted via seeds.
Future research should focus on the factors involved in microbiome assembly, the relative contribution of epi and endophytes to microbiomes, and the signaling crosstalk between plants and microbial communities.Rhizobiome assembly and the involvement of the plant in this process are currently enigmatic. Here, we have discussed multiple factors shaping the rhizobiome, including host genotype and development, root morphology, border cells and mucilage, and root exudates. Root exudation is a dynamic process, likely dependent on a plethora or transporters that are mostly uncharacterized. Spatially defined exudation likely results in distinct microbial communities that are observed to be associated with specific root parts. The success of microbial colonization of the rhizosphere depends on several aspects, such as chemotaxis, substrate specificity, competitiveness, and cooperativeness. Furthermore, endophytes likely form biofilms on the root surface, and encounter the plant immune system. Although some factors shaping root microbiomes emerge, many open questions remain .Due to the chemical complexity of soil, exudation is traditionally analyzed in hydroponic culture, an environment distant from the more natural settings of plant microbiome studies. Furthermore, novel technologies enabling high throughput screening of putative transporters against possible substrates are needed to reveal the impact of the respective substrates on the rhizobiome and, in turn, on plant health. An increased understanding of root morphology, exudation, and involved transporters will likely enable the engineering or breeding of plants with altered abilities to interact with specific beneficial microbes or pathogens. This needs to be complemented with an improved understanding of the substrate preferences of plant associated microbes, their interactions, and the mechanisms through which they benefit the plant. A holistic understanding of the functions of a healthy plant rhizobiome would enable the directed design of customized microbial communities. With this, specific plants in a given environment could be tailored to a specific purpose, such as phytoremediation, stress resistance, altered plant development, or increased yield.Interactions between plants and microbes are an integral part of our terrestrial ecosystem. There are several types of plant microbe interactions: competition, commensalism, mutualism, and parasitism. The more common interactions are commensalism or mutualism, where either one or both species benefit from the relationship, respectively . There are several excellent reviews reporting current research on lifestyles and molecular interactions of plant associated bacteria , rhizosphere interactions , plant responses to bacterial quorum sensing signals , endophyte applications , and rhizosphere bacteria responses to transgenic plants . Examination of these interactions helps us to understand natural phenomena that affect our daily lives and could lead to applications resulting in sustainable resources, less impact on the environment, cleanup of pollution and influence on atmospheric gases on a global scale. Advantages of using these interactions for biotechnological applications are many fold. The use of naturally existing plant microbe symbiosis for plant growth and bio control reduces synthetic fertilizer and pesticide treatments leading to cost effectiveness and less impact by nutrients and pesticides on surrounding fauna and flora. The production of useful compounds with pharmaceutical and industrial relevance using plant bacteria symbiosis is energy efficient and diminishes the need to add expensive precursors and catalysts. Remediation through conventional method, such as excavate and treat, is expensive and labor intensive. Conversely, plantmicrobial remediation strategies can be less intrusive and much more economical .Carbon sequestration through plant rhizosphere processes is a potentially sustainable method to lowering atmospheric carbon . This review focuses on recent progress in the fields of plant growth promotion, plant disease control, production of bio active compounds and bio materials, remediation of contaminated sites, and carbon sequestration. The potential of applying these new developments are discussed. Figure 1 summarizes applications resulting from microbe shoot and microbe root interactions and techniques used. Table 1 is a glossary of the techniques mentioned in this review. Plant microbe interactions have been utilized to improve plant growth for the production of food, fiber, bio fuels and key metabolites. The mutualistic interaction can be beneficial in directly providing nutrients to the plant or increasing the availability of compounds such as iron or phosphate. Free living plant growth promoting bacteria also produce compounds that directly affect plant metabolism or modulate phytohormone production or degradation. The phytohormones: auxins, cytokinins, gibberellic acid , abscisic acid and ethylene are signaling molecules essential for growth which mediate a range of developmental processes in plants. Recent studies on each of these areas are presented in the following section. As chemical fertilizers are costly both to the agricultural businesses and the environment, development of biofertilizers is an important and exciting area.