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For decades, the term “rhizosphere fauna” has been used as a synonym to denote agricultural pests among root herbivores, mainly nematodes and insect larvae. We want to break with this constrictive view, since the connection between plants and rhizosphere fauna is far more complex than simply that of resource and consumer. For example, plant roots have been shown to be neither defenceless victims of root feeders, nor passive recipients of nutrients, but instead play a much more active role in defending themselves and in attracting beneficial soil microorganisms and soil fauna. Most importantly, significant indirect feed-backs exist between consumers of rhizosphere microorganisms and plant roots. In fact, the majority of soil invertebrates have been shown to rely profoundly on the carbon inputs from roots, breaking with the dogma of soil food webs being mainly fueled by plant litter input from aboveground. In this review we will highlight areas of recent exciting progress and point out the black boxes that still need to be illuminated by rhizosphere zoologists and ecologists. [ABSTRACT FROM AUTHOR]

This review summarizes and discusses methodological approaches for studies on the impact of plant roots on the surrounding rhizosphere and for elucidation of the related mechanisms, covering a range from simple model experiments up to the field scale. A section on rhizosphere sampling describes tools and culture systems employed for analysis of root growth, root morphology, vitality testing and for monitoring of root activity with respect to nutrient uptake, water, ion and carbon flows in the rhizosphere. The second section on rhizosphere probing covers techniques to detect physicochemical changes in the rhizosphere as a consequence of root activity. This comprises compartment systems to obtain rhizosphere samples, visualisation techniques, reporter gene approaches and remote sensing technologies for monitoring the conditions in the rhizosphere. Approaches for the experimental manipulation of the rhizosphere by use of molecular and genetic methods as tools to study rhizosphere processes are discussed in a third section. Finally it is concluded that in spite of a wide array of methodological approaches developed in the recent past for studying processes and interactions in the rhizosphere mainly under simplified conditions in model experiments, there is still an obvious lack of methods to test the relevance of these findings under real field conditions or even on the scale of ecosystems. This also limits reliable data input and validation in current rhizosphere modelling approaches. Possible interactions between different environmental factors or plant-microbial interactions (e.g. mycorrhizae) are frequently not considered in model experiments. Moreover, most of the available knowledge arises from investigations with a very limited number of plant species, mainly crops and studies considering also intraspecific genotypic differences or the variability within wild plant species are just emerging. [ABSTRACT FROM AUTHOR]

Life on Earth is sustained by a small volume of soil surrounding roots, called the rhizosphere. The soil is where most of the biodiversity on Earth exists, and the rhizosphere probably represents the most dynamic habitat on Earth; and certainly is the most important zone in terms of defining the quality and quantity of the Human terrestrial food resource. Despite its central importance to all life, we know very little about rhizosphere functioning, and have an extraordinary ignorance about how best we can manipulate it to our advantage. A major issue in research on rhizosphere processes is the intimate connection between the biology, physics and chemistry of the system which exhibits astonishing spatial and temporal heterogeneities. This review considers the unique biophysical and biogeochemical properties of the rhizosphere and draws some connections between them. Particular emphasis is put on how underlying processes affect rhizosphere ecology, to generate highly heterogeneous microenvironments. Rhizosphere ecology is driven by a combination of the physical architecture of the soil matrix, coupled with the spatial and temporal distribution of rhizodeposits, protons, gases, and the role of roots as sinks for water and nutrients. Consequences for plant growth and whole-system ecology are considered. The first sections address the physical architecture and soil strength of the rhizosphere, drawing their relationship with key functions such as the movement and storage of elements and water as well as the ability of roots to explore the soil and the definition of diverse habitats for soil microorganisms. The distribution of water and its accessibility in the rhizosphere is considered in detail, with a special emphasis on spatial and temporal dynamics and heterogeneities. The physical architecture and water content play a key role in determining the biogeochemical ambience of the rhizosphere, via their effect on partial pressures of O2 and CO2, and thereby on redox potential and pH of the rhizosphere, respectively. We address the various mechanisms by which roots and associated microorganisms alter these major drivers of soil biogeochemistry. Finally, we consider the distribution of nutrients, their accessibility in the rhizosphere, and their functional relevance for plant and microbial ecology. Gradients of nutrients in the rhizosphere, and their spatial patterns or temporal dynamics are discussed in the light of current knowledge of rhizosphere biophysics and biogeochemistry. Priorities for future research are identified as well as new methodological developments which might help to advance a comprehensive understanding of the co-occurring processes in the rhizosphere. [ABSTRACT FROM AUTHOR]

This paper reviews strategies for manipulating plants and their root-associated microorganisms to improve plant health and productivity. Some strategies directly target plant processes that impact on growth, while others are based on our knowledge of interactions among the components of the rhizosphere (roots, microorganisms and soil). For instance, plants can be engineered to modify the rhizosphere pH or to release compounds that improve nutrient availability, protect against biotic and abiotic stresses, or encourage the proliferation of beneficial microorganisms. Rhizobacteria that promote plant growth have been engineered to interfere with the synthesis of stress-induced hormones such as ethylene, which retards root growth, and to produce antibiotics and lytic enzymes active against soilborne root pathogens. Rhizosphere engineering also can involve the selection by plants of beneficial microbial populations. For example, some crop species or cultivars select for and support populations of antibiotic-producing strains that play a major role in soils naturally suppressive to soil-borne fungal pathogens. The fitness of root-associated bacterial communities also can be enhanced by soil amendment, a process that has allowed the selection of bacterial consortia that can interfere with bacterial pathogens. Plants also can be engineered specifically to influence their associated bacteria, as exemplified by quorum quenching strategies that suppress the virulence of pathogens of the genus Pectobacterium. New molecular tools and powerful biotechnological advances will continue to provide a more complete knowledge of the complex chemical and biological interactions that occur in the rhizosphere, ensuring that strategies to engineer the rhizosphere are safe, beneficial to productivity, and substantially improve the sustainability of agricultural systems. [ABSTRACT FROM AUTHOR]

Iron is an essential element for plants and microbes. However, in most cultivated soils, the concentration of iron available for these living organisms is very low because its solubility is controlled by stable hydroxides, oxyhydroxides and oxides. In the rhizosphere, there is a high demand of iron because of the iron uptake by plants, and microorganisms which density and activity are promoted by the release of root exudates. Plants and microbes have evolved active strategies of iron uptake. Iron incorporation by these organisms lead to complex interactions ranging from competition to mutualism. These complex interactions are under the control of physico-chemical properties of the soils in which they occur, and reciprocally iron uptake strategies of plants and microbes impact these soil properties. These iron-mediated interactions between soils, plants and microbes impact the plant growth and health and their analysis, together with that of the resulting iron dynamics, is of a major agronomic interest. Analysis of the complex interactions soils, plants and microbes represent also a unique opportunity to progress in our knowledge of the rhizosphere ecology. This progression requires merging complementary expertises and study strategies in soil science, plant biology and microbiology. This review provides information on (i) iron status in soil and rhizosphere, iron uptake by plants and microbes, and on (ii) the corresponding study strategies. Finally, illustrations of how integration of these approaches allows gaining knowledge in the complex interactions occurring in the rhizosphere are given. [ABSTRACT FROM AUTHOR]

It is the aim of this chapter to present an overview of new, molecular tools that have been developed over recent years to study individual, single cells and composite, complex communities of microorganisms in the rhizosphere. We have carefully focused on culture-independent assays and selected methodologies that have already been or will soon be applicable for rhizosphere microbiology. Emphasis is placed on rhizosphere bacteria and the review first describes a number of the new methodologies developed for detection and localization of specific bacterial populations using modern electron and fluorescence microscopy combined with specific tagging techniques. First half of the chapter further comprises a thorough treatise of the recent development of reporter gene technology, i.e. using specific reporter bacteria to detect microscale distributions of rhizosphere compounds such as nutrients, metals and organic exudates or contaminants. Second half of the chapter devoted to microbial community analysis contains a thorough treatise of nucleotide- and PCR-based technologies to study composition and diversity of indigenous bacteria in the natural rhizosphere. Also included are the most recent developments of functional gene and gene expression analyses in the rhizosphere based on specific mRNA transcript or transcriptome analysis, proteome analysis and construction of metagenomic libraries. [ABSTRACT FROM AUTHOR]

Soils are the product of the activities of plants, which supply organic matter and play a pivotal role in weathering rocks and minerals. Many plant species have a distinct ecological amplitude that shows restriction to specific soil types. In the numerous interactions between plants and soil, microorganisms also play a key role. Here we review the existing literature on interactions between plants, microorganisms and soils, and include considerations of evolutionary time scales, where possible. Some of these interactions involve intricate systems of communication, which in the case of symbioses such as the arbuscular mycorrhizal symbiosis are several hundreds of millions years old; others involve the release of exudates from roots, and other products of rhizodeposition that are used as substrates for soil microorganisms. The possible reasons for the survival value of this loss of carbon over tens or hundreds of millions of years of evolution of higher plants are discussed, taking a cost-benefit approach. Co-evolution of plants and rhizosphere microorganisms is discussed, in the light of known ecological interactions between various partners in terrestrial ecosystems. Finally, the role of higher plants, especially deep-rooted plants and associated microorganisms in the weathering of rocks and minerals, ultimately contributing to pedogenesis, is addressed. We show that rhizosphere processes in the long run are central to biogeochemical cycles, soil formation and Earth history. Major anticipated discoveries will enhance our basic understanding and allow applications of new knowledge to deal with nutrient deficiencies, pests and diseases, and the challenges of increasing global food production and agroecosystem productivity in an environmentally responsible manner. [ABSTRACT FROM AUTHOR]

Diversity, structure and productivity of above-ground compartment of terrestrial ecosystems have been generally considered as the main drivers of the relationships between diversity and ecosystem functioning. More recently it has been suggested that plant population dynamics may be linked with the development of the below-ground community. The biologically active soil zone where root-root and root-microbe communications occur is named “Rhizosphere” where root exudates play active roles in regulating rhizosphere interactions. Root exudation can regulate the soil microbial community, withstand herbivory, facilitate beneficial symbioses, modify the chemical and physical soil properties and inhibit the growth of competing plant species. In this review, we explore the current knowledge assessing the importance of root exudates in plant interactions, in communications between parasitic plants and their hosts and how some soil microbial components could regulate plant species coexistence and change relationships between plants. This review will be focussed on several well documented biological processes regulating plant-plant communications such as exotic plant species invasions, negative root-root communication (allelopathy) and parasitic plant / host plant interactions and how some soil microbial components can interfere with signal traffic between roots. The reported data show that the overall effect of one plant to another results from multiple interacting mechanisms where soil microbiota can be considered as a key component. [ABSTRACT FROM AUTHOR]

The article discusses some aspects of the taxonomical and functional microbial diversity of the rhizosphere. Rhizospheres are responsible for making a transition zone between the bulk soil and the plant root surface. According to botanists, bacteria residing in the rhizosphere most likely originate from the surrounding bulk soil and thrive under conditions prevailing in the neighbourhood of plant roots. Studies have concluded that microorganisms present in the rhizosphere play important roles in the growth and in ecological fitness of their plant host.

This paper will exemplify molecular communications in the rhizosphere, especially between plants and bacteria, and between bacteria and bacteria. More specifically, we describe signalling pathways that allow bacteria to sense a wide diversity of plant signals, plants to respond to bacterial infection, and bacteria to coordinate gene expression at population and community level. Thereafter, we focus on mechanisms evolved by bacteria and plants to disturb bacterial signalling, and by bacteria to modulate hormonal signalling in plants. Finally, the dynamics of signal exchange and its biological significance we elaborate on the cases of Rhizobium symbiosis and Agrobacterium pathogenesis. [ABSTRACT FROM AUTHOR]

The rhizosphere is a hot spot of microbial interactions as exudates released by plant roots are a main food source for microorganisms and a driving force of their population density and activities. The rhizosphere harbors many organisms that have a neutral effect on the plant, but also attracts organisms that exert deleterious or beneficial effects on the plant. Microorganisms that adversely affect plant growth and health are the pathogenic fungi, oomycetes, bacteria and nematodes. Most of the soilborne pathogens are adapted to grow and survive in the bulk soil, but the rhizosphere is the playground and infection court where the pathogen establishes a parasitic relationship with the plant. The rhizosphere is also a battlefield where the complex rhizosphere community, both microflora and microfauna, interact with pathogens and influence the outcome of pathogen infection. A wide range of microorganisms are beneficial to the plant and include nitrogen-fixing bacteria, endo- and ectomycorrhizal fungi, and plant growth-promoting bacteria and fungi. This review focuses on the population dynamics and activity of soilborne pathogens and beneficial microorganisms. Specific attention is given to mechanisms involved in the tripartite interactions between beneficial microorganisms, pathogens and the plant. We also discuss how agricultural practices affect pathogen and antagonist populations and how these practices can be adopted to promote plant growth and health. [ABSTRACT FROM AUTHOR]

Selection of modern varieties has typically been performed in standardized, high fertility systems with a primary focus on yield. This could have contributed to the loss of plant genes associated with efficient nutrient acquisition strategies and adaptation to soil-related biotic and abiotic stresses if such adaptive strategies incurred a cost to the plant that compromised yield. Furthermore, beneficial interactions between plants and associated soil organisms may have been made obsolete by the provision of nutrients in high quantity and in readily plant available forms. A review of evidence from studies comparing older traditional varieties to modern high yielding varieties indeed showed that this has been the case. Given the necessity to use scarce and increasingly costly fertilizer inputs more efficiently while also raising productivity on poorer soils, it will be crucial to reintroduce desirable rhizosphere-related traits into elite cultivars. Traits that offer possibilities for improving nutrient acquisition capacity, plant–microbe interaction s and tolerance to abiotic and biotic soil stresses in modern varieties were reviewed. Despite the considerable effort devoted to the identification of suitable donors and of genetic factors associated with these beneficial traits, progress in developing improved varieties has been slow and has so far largely been confined to modifications of traditional breeding procedures. Modern molecular tools have only very recently started to play a rather small role. The few successful cases reviewed in this paper have shown that novel breeding approaches using molecular tools do work in principle. When successful, they involved close collaboration between breeders and scientists conducting basic research, and confirmation of phenotypes in field tests as a ‘reality check’. We concluded that for novel molecular approaches to make a significant contribution to breeding for rhizosphere related traits it will be essential to narrow the gap between basic sciences and applied breeding through more interdisciplinary research that addresses rather than avoids the complexity of plant–soil interactions. [ABSTRACT FROM AUTHOR]

Plant-assisted bioremediation or phytoremediation holds promise for in situ treatment of polluted soils. Enhancement of phytoremediation processes requires a sound understanding of the complex interactions in the rhizosphere. Evaluation of the current literature suggests that pollutant bioavailability in the rhizosphere of phytoremediation crops is decisive for designing phytoremediation technologies with improved, predictable remedial success. For phytoextraction, emphasis should be put on improved characterisation of the bioavailable metal pools and the kinetics of resupply from less available fractions to support decision making on the applicability of this technology to a given site. Limited pollutant bioavailability may be overcome by the design of plant–microbial consortia that are capable of mobilising metals/metalloids by modification of rhizosphere pH (e.g. by using Alnus sp. as co-cropping component) and ligand exudation, or enhancing bioavailability of organic pollutants by the release of biosurfactants. Apart from limited pollutant bioavailability, the lack of competitiveness of inoculated microbial strains (in particular degraders) in field conditions appears to be another major obstacle. Selecting/engineering of plant–microbial pairs where the competitiveness of the microbial partner is enhanced through a “nutritional bias” caused by exudates exclusively or primarily available to this partner (as known from the “opine concept”) may open new horizons for rhizodegradation of organically polluted soils. The complexity and heterogeneity of multiply polluted “real world” soils will require the design of integrated approaches of rhizosphere management, e.g. by combining co-cropping of phytoextraction and rhizodegradation crops, inoculation of microorganisms and soil management. An improved understanding of the rhizosphere will help to translate the results of simplified bench scale and pot experiments to the full complexity and heterogeneity of field applications. [ABSTRACT FROM AUTHOR]

We review methods and models that help to assess how root activity changes soil properties and affects the fluxes of matter in the soil. Subsections discuss (1) experimental systems including plant treatments in artificial media, studying the interaction of model root and microbial exudates with soil constituents, and microcosms to distinguish between soil compartments differing in root influence, (2) the sampling and characterization of rhizosphere soil and solution, focusing on the separation of soil at different distances from roots and the spatially resolved sampling of soil solution, (3) cutting-edge methodologies to study chemical effects in soil, including the estimation of bioavailable element or ion contents (biosensors, diffusive gradients in thin-films), studying the ultrastructure of soil components, localizing elements and determining their chemical form (microscopy, diffractometry, spectroscopy), tracing the compartmentalization of substances in soils (isotope probing, autoradiography), and imaging gradients in-situ with micro electrodes or gels or filter papers containing dye indicators, (4) spectroscopic and geophysical methods to study the plants influence on the distribution of water in soils, and (5) the modeling of rhizosphere processes. Macroscopic models with a rudimentary depiction of rhizosphere processes are used to predict water or nutrient requirements by crops and forests, to estimate biogeochemical element cycles, to calculate soil water transport on a profile scale, or to simulate the development of root systems. Microscopic or explanatory models are based on mechanistic or empirical relations that describe processes on a single root or root system scale and/or chemical reactions in soil solution. We conclude that in general we have the tools at hand to assess individual processes on the microscale under rather artificial conditions. Microscopic, spectroscopic and tracer methods to look at processes in small “aliquots” of naturally structured soil seem to step out of their infancy and have become promising tools to better understand the complex interactions between plant roots, soil and microorganisms. On the field scale, while there are promising first results on using non-invasive geophysical methods to assess the plant’s influence on soil moisture, there are no such tools in the pipeline to assess the spatial heterogeneity of chemical properties and processes in the field. Here, macroscopic models have to be used, or model results on the microscopic level have to be scaled up to the whole plant or plot scale. Upscaling is recognized as a major challenge. [ABSTRACT FROM AUTHOR]

Changes in chemical properties in soil around plant roots influence many microbial processes, including those having an impact on greenhouse gas emissions. To potentially mitigate these emissions according to the Kyoto protocol, knowledge about how and where these gases are produced and consumed in soils is required. In this review, we focus on the greenhouse gases nitrous oxide and methane, which are produced by nitrifying and denitrifying prokaryotes and methanogenic archaea, respectively. After describing the microbial processes involved in production and consumption of nitrous oxide and methane and how they can be affected in the rhizosphere, we give an overview of nitrous oxide and methane emissions from the rhizosphere and soils and sediments with plants. We also discuss strategies to mitigate emissions from the rhizosphere and consider possibilities for carbon sequestration. [ABSTRACT FROM AUTHOR]

The rhizodeposition of plants dramatically influence the surrounding soil and its microflora. Root exudates have pronounced selective and promoting effects on specific microbial populations which are able to respond with chemotaxis and fast growth responses, such that only a rather small subset of the whole soil microbial diversity is finally colonizing roots successfully. The exudates carbon compounds provide readily available nutrient and energy sources for heterotrophic organisms but also contribute e.g. complexing agents, such as carboxylates, phenols or siderophores for the mobilization and acquisition of rather insoluble minerals. Root exudation can also quite dramatically alter the pH- and redox-milieu in the rhizosphere. In addition, not only specific stimulatory compounds, but also antimicrobials have considerable discriminatory effect on the rhizosphere microflora. In the “biased rhizosphere” concept, specific root associated microbial populations are favored based on modification of the root exudation profile. Rhizosphere microbes may exert specific plant growth promoting or biocontrol effects, which could be of great advantage for the plant host. Since most of the plant roots have symbiotic fungi, either arbuscular or ectomycorrhizal fungi, the impact of plants towards the rhizosphere extends also to the mycorrhizosphere. The selective effect of the roots towards the selection of microbes also extends towards the root associated and symbiotic fungi. While microbes are known to colonize plant roots endophytically, also mycorrhiza are now known to harbor closely associated bacterial populations even within their hyphae. The general part of the manuscript is followed by the more detailed presentation of specific examples for the selection and interaction of roots and microbes, such as in the rhizosphere of strawberry, potato and oilseed rape, where the soil-borne plant pathogen Verticillium dahliae can cause high yield losses; the potential of biocontrol by specific constituents of the rhizosphere microbial community is demonstrated. Furthermore, plant cultivar specificity of microbial communities is described in different potato lines including the case of transgenic lines. Finally, also the specific selective effect of different Medicago species on the selection of several arbuscular mycorrhizal taxa is presented. [ABSTRACT FROM AUTHOR]

The loss of organic and inorganic carbon from roots into soil underpins nearly all the major changes that occur in the rhizosphere. In this review we explore the mechanistic basis of organic carbon and nitrogen flow in the rhizosphere. It is clear that C and N flow in the rhizosphere is extremely complex, being highly plant and environment dependent and varying both spatially and temporally along the root. Consequently, the amount and type of rhizodeposits (e.g. exudates, border cells, mucilage) remains highly context specific. This has severely limited our capacity to quantify and model the amount of rhizodeposition in ecosystem processes such as C sequestration and nutrient acquisition. It is now evident that C and N flow at the soil–root interface is bidirectional with C and N being lost from roots and taken up from the soil simultaneously. Here we present four alternative hypotheses to explain why high and low molecular weight organic compounds are actively cycled in the rhizosphere. These include: (1) indirect, fortuitous root exudate recapture as part of the root’s C and N distribution network, (2) direct re-uptake to enhance the plant’s C efficiency and to reduce rhizosphere microbial growth and pathogen attack, (3) direct uptake to recapture organic nutrients released from soil organic matter, and (4) for inter-root and root–microbial signal exchange. Due to severe flaws in the interpretation of commonly used isotopic labelling techniques, there is still great uncertainty surrounding the importance of these individual fluxes in the rhizosphere. Due to the importance of rhizodeposition in regulating ecosystem functioning, it is critical that future research focuses on resolving the quantitative importance of the different C and N fluxes operating in the rhizosphere and the ways in which these vary spatially and temporally. [ABSTRACT FROM AUTHOR]

The rhizosphere is a complex environment where roots interact with physical, chemical and biological properties of soil. Structural and functional characteristics of roots contribute to rhizosphere processes and both have significant influence on the capacity of roots to acquire nutrients. Roots also interact extensively with soil microorganisms which further impact on plant nutrition either directly, by influencing nutrient availability and uptake, or indirectly through plant (root) growth promotion. In this paper, features of the rhizosphere that are important for nutrient acquisition from soil are reviewed, with specific emphasis on the characteristics of roots that influence the availability and uptake of phosphorus and nitrogen. The interaction of roots with soil microorganisms, in particular with mycorrhizal fungi and non-symbiotic plant growth promoting rhizobacteria, is also considered in relation to nutrient availability and through the mechanisms that are associated with plant growth promotion. [ABSTRACT FROM AUTHOR]

The article reviews the book "The Rhizosphere: Biochemistry and Organic Substances at the Soil-Plant Interface," second edition, edited by R. Pinto, Z. Varanini and P. Nannipieri.

The article reviews the book "The Rhizosphere: An Ecological Perspective," edited by Z.G. Cardon and J.L. Whitbeck.

The article reviews the book "The Rhizosphere: An Ecological Perspective," edited by Zoe G. Cardon and Julie L. Whitbeck.

Reviews the book "The Rhizosphere. Wiley Series in Ecological and Applied Microbiology," edited by J.M. Lynch.

The article reviews the book "The Rhizosphere: Biochemistry and Organic Substances at the Soil-Plant Interface," 2nd edn., edited by R. Pinton, Z. Varanini and P. Nannipieri.

The article reviews the book “The Rhizosphere: Biochemistry and Organic Substances at the Soil-Plant Interface,” edited by Roberto Pinton, Zeno Varanini, and Paolo Nannipieri.

The article reviews the book "The Rhizosphere: An Ecological Perspective," edited by Z. G. Cardon and J. L. Whitbeck.

The article reviews the book "Nutrient cycling in terrestrial ecosystems," edited by Petra Marschner and Zdenko Rengel.

The article reviews the book "The Rhizosphere: An Ecological Perspective," by Zoe G. Cardon and Julie L. Whitbeck.

Reviews the book "The Rhizosphere," by Elroy Curl and Bryan Truelove.

Reviews the book "The Rhizosphere and Plant Growth," by Keister, D. L. and Cregan, P. B.

Reviews the book "The Rhizosphere," by E.A. Curl and B. Truelove.