Fact-checked by Grok 2 weeks ago

Plant root exudates

Plant root exudates are organic compounds, primarily low-molecular-weight carbon-rich substances such as sugars, , organic acids, and , that are actively secreted or passively released by living roots into the surrounding environment known as the . These exudates represent a significant portion of photosynthates, accounting for 5–21% of the carbon fixed through , and serve as essential mediators for material cycling, energy exchange, and information transfer between plant roots and the ecosystem. The composition of root exudates is diverse and dynamic, encompassing primary metabolites like carbohydrates and amino acids, as well as secondary metabolites such as phenolics and terpenoids, with variations influenced by plant species, developmental stage, diurnal cycles, and environmental stressors like drought or nutrient availability. For instance, across phylogenetically distinct plants like Arabidopsis thaliana, Brachypodium distachyon, and Medicago truncatula, a core set of about 21% of exudate metabolites— including amino acids and organic acids— is conserved, suggesting a universal role in recruiting beneficial soil microbes. Exudation patterns can shift rapidly, with up to 32% of compounds showing diurnal fluctuations, highlighting the adaptive nature of this process. Functionally, root exudates play pivotal roles in plant-soil interactions, including mobilizing nutrients like and iron through and acidification by acids, thereby enhancing plant uptake in -poor . They also shape the by acting as nutrients and chemical signals that attract symbiotic microbes, such as nitrogen-fixing and arbuscular mycorrhizal fungi, while deterring pathogens via compounds. Additionally, exudates mediate plant-plant communication through allelopathic effects, where compounds like inhibit competing roots, and influence broader processes such as carbon turnover, where they can both stabilize carbon in mineral-associated pools and prime microbial , leading to counterintuitive net effects on storage depending on exudate quantity and quality. These multifaceted roles underscore the importance of root exudates in , , and responses to , as alterations in exudate profiles under stress conditions can affect microbial diversity and .

Definition and Composition

Definition

Plant root exudates are low-molecular-weight organic compounds that are actively or passively released from intact, living plant roots into the surrounding . These compounds, which include sugars, , and organic acids, originate from metabolic processes within root cells and diffuse or are secreted across the root plasma membrane. The primary purpose of root exudates is to modify the environment near the roots, known as the , to facilitate acquisition and mediate interactions with such as and fungi. By altering chemistry and providing carbon sources, exudates promote the growth of beneficial microorganisms that enhance uptake, suppress pathogens, and improve resilience to environmental stresses. Root exudates are distinct from root lysates or sloughed cells, which represent passive debris released from dead or dying tissues through cell turnover or . Unlike these structural components, exudates are specifically mobilized from viable root cells and play an active role in dynamic soil-plant interactions rather than merely contributing to decomposition. The concept of root exudates was first described in the early through foundational studies on root-soil interactions, with significant recognition of their biological roles emerging in via observations of microbial stimulation around roots. Key advancements occurred in the with the application of radioisotope labeling techniques, which enabled precise tracking of exudate release and composition from living roots, revolutionizing understanding of rhizosphere dynamics.

Chemical Composition

Plant root exudates comprise a diverse array of low-molecular-weight compounds, primarily categorized into sugars, , organic acids, , and secondary metabolites. Sugars such as glucose and serve as primary carbon sources, while like glutamate provide nitrogenous compounds essential for microbial nutrition. Organic acids, including citrate and malate, function as chelators for nutrients, and act as signaling molecules. Secondary metabolites, exemplified by strigolactones, contribute to specialized interactions in the environment. The chemical composition of root exudates exhibits significant variability influenced by plant developmental stage, , nutrient availability, and environmental stresses. For instance, during early growth phases, exudates tend to be richer in simple sugars, whereas mature roots may increase secretion of secondary metabolites. affects the solubility and release of organic acids, with acidic conditions promoting their exudation. Nutrient deficiencies, such as limitation, trigger enhanced release of organic acids like citrate and malate to mobilize insoluble phosphates from the . Quantitatively, plants allocate approximately 5-20% of their annual photosynthate carbon to root exudates, representing a substantial in belowground interactions. Specific exudation rates vary by and conditions; for example, in (Zea mays), sugars can constitute up to 1.5 mg per g root dry weight. These allocations underscore the ecological cost and benefit of exudate production for . The identification of root exudate components has relied on advanced analytical techniques since the 1970s, with (HPLC) and gas chromatography-mass spectrometry (GC-MS) becoming standard for separating and quantifying low-molecular-weight organics like sugars, , and acids. HPLC excels in analyzing polar compounds without derivatization, while GC-MS provides detailed structural elucidation through mass spectra, enabling comprehensive profiling of exudate diversity. These methods, pivotal in seminal studies from the era, continue to underpin modern approaches for exudates.

The Rhizosphere Environment

Overview of the Rhizosphere

The refers to the narrow zone of soil surrounding plant , typically extending 1–5 mm from the surface, where activities profoundly alter the physical, chemical, and biological properties of the soil compared to bulk soil. This dynamic interface is characterized by elevated microbial densities, often 10–100 times higher than in surrounding bulk soil, fostering a of . The term "" was first coined in 1904 by German agronomist Lorenz Hiltner, who described it as the "root zone of soil enrichment," emphasizing its role in enhancing nutrient availability through microbial influences on . Key features of the include steep gradients in availability, with higher concentrations of carbon and near the roots due to root-mediated inputs and microbial turnover. Oxygen levels deplete rapidly close to the root surface as a result of root and microbial metabolism, creating microaerobic or conditions that favor certain microbial groups. Additionally, can fluctuate by up to two units across the , influenced by root uptake, proton extrusion, and microbial processes, which in turn affect solubility and microbial community structure. The 's microbial is dominated by , followed by fungi and , with total microbial abundances reaching 10^9 to 10^11 cells per gram of rhizosphere , sustained largely by root-derived carbon inputs that promote proliferation. This elevated density and underpin the rhizosphere's function as a biogeochemical , where microbial communities drive processes like and nutrient mobilization essential for plant growth.

Role of Exudates in Rhizosphere Formation

Plant root exudates play a pivotal role in shaping the by altering its chemical, biological, and physical properties, thereby creating a dynamic microenvironment that supports plant growth and microbial interactions. These low-molecular-weight compounds, including sugars, , and organic acids, diffuse from roots into the surrounding , establishing gradients that influence soil chemistry and microbial recruitment. Chemically, root exudates modify the rhizosphere through the release of organic acids such as citric and malic acid, which lower the soil pH and facilitate the solubilization of immobile minerals like phosphorus. For instance, citric acid at concentrations typical of root exudates can enhance phosphorus availability by chelating metal ions bound to phosphate. This acidification process not only mobilizes nutrients but also influences metal speciation, such as iron and aluminum, thereby regulating their bioavailability in the soil solution. Biologically, exudates act as chemoattractants that selectively recruit beneficial microbes to the , promoting the formation of biofilms and shifting microbial community composition. Compounds like benzoxazinoids in root exudates specifically attract , enriching its abundance in the rhizosphere. This selective structuring can increase beneficial bacterial densities by 10- to 100-fold compared to bulk soil, stabilizing microbial consortia that aid in nutrient cycling and stress tolerance. Physically, polysaccharide-rich in root exudates binds soil particles, improving aggregation and water retention in the . from species like increases soil aggregate stability by acting as a biological glue, enhancing water-holding capacity under drying conditions and facilitating penetration through compacted soils. This structural modification creates a more porous , which supports prolonged hydration and microbial activity near the surface. Quantitatively, root exudates represent up to 40% of the photosynthetically fixed carbon allocated to the , driving significant carbon flux that influences oxygen and gradients. This substantial input fuels microbial respiration, leading to local oxygen depletion and the establishment of microsites with reduced potentials, which in turn affect nutrient transformations like .

Mechanisms of Exudate Release

Physiological Processes

Plant root exudates are released through a combination of passive and active physiological mechanisms that facilitate the transport of diverse compounds across root cell membranes into the . Non-polar, lipophilic compounds, such as certain and hydrocarbons, primarily exit via through the lipid bilayers of membranes, driven by concentration gradients without energy expenditure. In contrast, polar compounds like sugars, , and require facilitated or ; for instance, polar molecules and often move through ion channels or permeases via , while active involves ATP-binding cassette () transporters that pump metabolites against gradients. A well-characterized example is the exudation of malate, an that chelates aluminum in acidic soils, mediated by the aluminum-activated malate transporter 1 (ALMT1) protein, which functions as an anion channel in the root membrane. The primary cellular sites of exudate release are the root epidermis and , where metabolites are loaded symplastically—through plasmodesmata connecting living cells—before crossing into the , the outside the plasma membrane. Root hairs, as specialized extensions of epidermal cells, play a prominent role in this process by increasing surface area and actively secreting exudates, thereby enhancing local concentrations in the . This symplastic-to-apoplastic transition ensures efficient delivery of compounds to the soil interface, with the flux predominantly occurring at undifferentiated zones like the root apex. Exudation rates vary with root developmental stages, peaking during periods of active such as the of lateral , where epidermal disruption and increased metabolic activity promote higher release. At these sites, the process supports branching by altering local chemistry, though the exact mechanisms linking development to flux remain under study. Many release pathways, particularly via ABC transporters and proton-coupled systems, are ATP-dependent, representing a metabolic investment that draws from respiration to sustain exudate production and . Models of carbon allocation indicate that exudation can account for a notable fraction of belowground carbon , though precise contributions to total energy budgets vary by and conditions. Recent as of 2025 has also highlighted the role of exuded proteins in these mechanisms, contributing to acquisition, microbial , and defense.

Regulation Factors

The release of root exudates is tightly regulated by a combination of internal physiological signals and external environmental cues, ensuring adaptive responses to developmental needs and stresses. Internally, phytohormones play a central role in modulating production and composition. For instance, enhances root exudation by upregulating involved in synthesis, such as and organic acids, thereby influencing interactions. Developmental signals, including circadian rhythms, also drive rhythmic patterns in exudate secretion; in , root exudation of sugars and peaks during the day, synchronized with the plant's internal clock to optimize microbial recruitment. External factors, particularly nutrient availability, significantly alter exudate profiles to facilitate acquisition. Under , graminaceous plants like upregulate the secretion of phytosiderophores, such as mugineic acid, which chelate iron in the soil, with exudation rates increasing at least 10-fold compared to iron-sufficient conditions. Abiotic stresses exert contrasting effects: typically reduces overall exudate release by limiting root metabolic activity and carbon allocation, as observed in seedlings where total organic carbon exudation was significantly reduced, with rates up to threefold lower under deficit. In contrast, biotic stresses like attack stimulate compound exudation; for example, foliar infection in leads to increased root secretion of coumarins and , enhancing defense signaling in the . At the genetic level, specific transporters govern exudate release, with the multidrug and toxic compound extrusion () family being prominent since their identification in the early 2000s. In , the AtMATE encodes a citrate transporter activated under aluminum , enabling targeted efflux to detoxify the , a conserved across for metal tolerance. Feedback loops involving microbes further refine regulation; bacterial signals, such as N-acyl homoserine lactones, can be perceived by roots, prompting amplified exudation of compatible compounds that reinforce beneficial associations, as seen in legume-rhizobial symbioses. These interactions create dynamic feedbacks where initial exudate cues shape microbial communities, which in turn modulate plant to sustain exudation.

Functions and Interactions

Kin Recognition

Plants recognize kin through chemical cues in root exudates, enabling them to adjust root foraging behavior to minimize competition with siblings while aggressively exploring resources near unrelated individuals. This phenomenon involves reduced root proliferation and exudation in response to kin signals, promoting cooperative resource partitioning among related plants. The first empirical demonstration of kin recognition occurred in the annual sea rocket (Cakile edentula), where plants grown with siblings allocated less biomass to roots and exhibited greater separation in root placement compared to those paired with strangers, suggesting root interactions as the primary cue. Subsequent studies in Arabidopsis thaliana confirmed that root exudates mediate this process in hydroponic systems, with exposure to stranger exudates inducing significantly more lateral root formation than sibling exudates, and active root secretion required for discrimination. While evidence supports kin recognition in several species, the precise mechanisms and evolutionary drivers remain subjects of debate. Signaling molecules within root exudates, such as , serve as key cues for kin detection, influencing downstream and root architecture. , including , are secreted in varying profiles that differ between kin and non-kin, triggering differential root responses; for instance, A. thaliana roots exposed to non-kin exudates show enhanced branching to preempt resources. These signals integrate with broader dynamics, occasionally intersecting with microbial communities that amplify or modify exudate effects. Experimental evidence underscores the adaptive value of , rooted in theory, where favor relatives to enhance by curbing . In (Triticum aestivum), kin-paired exhibit reduced root mass overlap and total root compared to stranger pairs, leading to more efficient acquisition and higher overall yields. Genetic studies in A. thaliana using T-DNA insertion of ABC transporters (e.g., AtPGP1, AtATH1, AtATH10) abolish kin-specific responses, confirming exudate transport dependency, as roots fail to distinguish siblings and show indiscriminate proliferation. Similar patterns occur in C. edentula, where kin reduce competitive root investment, supporting the evolutionary conservation of this trait across species.

Interactions with Soil Microorganisms

Plant root exudates play a pivotal role in recruiting microorganisms to the by providing carbon sources such as sugars and , which serve as nutrients to attract and sustain bacterial populations like . In , exuded from roots act as signaling molecules that specifically induce in compatible , such as Sinorhizobium meliloti in (Medicago sativa), promoting and nodule formation for symbiotic . For instance, like and in (Glycine max) root exudates activate NodD proteins, leading to production and infection thread development, with knockdown of synthase genes reducing these by up to 100% and completely inhibiting nodulation. This recruitment process is modulated by environmental factors, including low availability, which enhances biosynthesis and exudation to favor symbiotic partners. Exudates also modulate microbial communities through secondary metabolites that either inhibit pathogens or promote beneficial symbionts. In maize (Zea mays), benzoxazinoids such as 2,4-dihydroxy-7-methoxy-1,4-benzoxazin-3-one (DIMBOA) in root exudates exhibit antimicrobial activity, suppressing fungal pathogens like Fusarium species by altering rhizosphere microbiota and favoring antagonistic bacteria such as Pseudomonas putida. These compounds reduce populations of proteobacteria and actinobacteria associated with pathogenesis, thereby enriching beneficial microbes that produce antibiotics like phenazines for biocontrol. Conversely, strigolactones in root exudates stimulate arbuscular mycorrhizal (AM) fungi by inducing hyphal branching at concentrations as low as 10⁻¹³ M, activating mitochondrial metabolism to increase respiration and lipid catabolism within 1 hour, which enhances fungal proliferation and root colonization across species like Gigaspora rosea and Glomus intraradices. Root exudates drive significant shifts in soil microbial community structure, accounting for substantial portions of diversity changes in the , as revealed by metagenomic studies. For example, high-sugar treatments reduce bacterial richness and evenness (P < 0.05), with dominating up to 59% of the community, while organic acid-rich exudates alter overall composition (PERMANOVA, P = 0.001) over 20 days, enriching functional genes for phytohormone production like that suppress pathogens. These shifts promote beneficial that compete with pathogens, as seen in benzoxazinoid-mediated enrichment of antibiotic-producing taxa. Metagenome-assembled genomes from such studies identify diverse phyla (20 total) responsive to variations, underscoring exudates' role in assembling disease-suppressive microbiomes. Symbiotic interactions facilitated by exudates lead to enhanced and crop yield improvements. In legume-rhizobia symbioses, flavonoid-induced nodulation increases N₂ fixation efficiency, with systems like faba bean () and showing 35% higher biomass and 61% greater grain yield without fertilizer, attributed to elevated exudation (up 73%) and upregulated nodulation genes. These outcomes can boost yields by 28–61% depending on inputs, demonstrating exudates' quantitative impact on symbiotic in agricultural settings.

Ecological Significance

Nutrient Cycling

Plant root exudates play a crucial role in nutrient cycling by mobilizing insoluble soil nutrients, particularly phosphorus (P) and iron (Fe), through the action of low-molecular-weight organic acids such as citrate, malate, and oxalate. These compounds chelate metal ions, forming soluble complexes that enhance nutrient availability in the rhizosphere. For instance, citrate exudation under P or Fe deficiency conditions increases the solubility of sparingly soluble phosphates and iron oxides by lowering soil pH and competing for binding sites on mineral surfaces. In acidic soils, citrate can elevate P concentrations in the soil solution by 10- to 1000-fold, facilitating direct uptake by plant roots and reducing dependency on fertilizer inputs. Exudates also mediate nutrient cycling indirectly by stimulating soil microbial communities involved in organic nutrient mineralization. Sugars and amino acids in root exudates serve as carbon sources, promoting the growth of phosphatase-producing bacteria that hydrolyze organic P compounds into bioavailable inorganic forms, thereby recycling P within the rhizosphere. Similarly, for nitrogen (N) cycling, exudate-derived amino acids enhance microbial mineralization rates, converting organic N into ammonium that plants can assimilate, with higher exudate C:N ratios shifting microbial strategies toward greater N release. This microbial stimulation can increase phosphatase activity by up to sixfold in response to exudate addition, amplifying P recycling efficiency. The quantitative impacts of exudate-mediated cycling are substantial, particularly for acquisition in grown on nutrient-poor soils. In many agricultural systems, exudate-induced solubilization accounts for a major proportion of uptake, with organic acids contributing to enhanced that supports productivity; for example, in phosphorus-limited environments, citrate exudation has been shown to boost root uptake rates significantly through targeted modeling of processes. Studies in cultivation demonstrate that improved nutrient mobilization via exudates can lead to yield increases under flooded conditions by optimizing and dynamics. These effects underscore the role of exudates in bridging soil nutrient limitations and plant demands. Over longer timescales, root exudates influence (SOM) turnover by providing labile carbon inputs that accelerate microbial and stabilize residues into . Exudates stimulate microbial activity, promoting the incorporation of plant-derived carbon into stable SOM pools and enhancing overall across growing seasons. This process alters rates, with exudates contributing to formation by fostering aggregate stability and nutrient retention in the matrix.

Plant-Plant Communication

Plant root exudates facilitate inter-plant communication by releasing chemical signals into the soil that influence the growth, defense, and competitive interactions of neighboring plants, distinct from kin-specific cues. These signals, including low-molecular-weight compounds like phenolics and terpenoids, diffuse through the rhizosphere to modulate the physiology of adjacent individuals, promoting or inhibiting development based on environmental pressures. Allelopathy represents a key mechanism of plant-plant communication via root exudates, where phenolic compounds and terpenoids are released to suppress the growth of competitors. For instance, juglone, a naphthoquinone exuded by black walnut (Juglans nigra) roots, inhibits seed germination and root elongation in susceptible species like tomatoes and corn by disrupting cellular respiration and nutrient uptake. Similarly, sorgoleone, a lipid benzoxazinoid from sorghum (Sorghum bicolor) root exudates, reduces weed seed germination by up to 50% through interference with membrane integrity and enzyme activity. Field studies since the 1980s have quantified the diffusion of such allelochemicals up to 10 cm from the root surface, enabling effective inhibition over short distances in dense plantings. Beyond inhibition, root exudates mediate stress signaling among plants, particularly in response to herbivory, by releasing volatile organic compounds that prime neighboring plants for defense activation. In (Solanum lycopersicum) systems, root-emitted green leaf volatiles (GLVs), such as (Z)-3-hexenyl acetate, increase upon insect attack and diffuse to adjacent plants, enhancing pathway expression and reducing subsequent damage. These volatiles warn non-attacked neighbors, inducing systemic resistance without direct contact. Evolutionarily, root exudate-mediated communication promotes optimal plant spacing in dense populations by balancing competition and facilitation, with genetic underpinnings identified through (QTL) mapping in crops like and . QTL studies reveal heritable variation in exudate composition that enhances allelopathic effects, contributing to weed suppression and improved stand establishment under crowding. This evolution likely arose to optimize partitioning in natural and agricultural settings.

References

  1. [1]
    The core metabolome and root exudation dynamics of three ... - Nature
    Mar 24, 2023 · Root exudates are plant-derived, exported metabolites likely shaping root-associated microbiomes by acting as nutrients and signals.
  2. [2]
    Root exudates contribute to belowground ecosystem hotspots
    Oct 5, 2022 · Root exudates are an essential carrier for material cycling, energy exchange, and information transfer between the belowground parts of plants and the soil.
  3. [3]
    Rhizosphere - Soil Ecology Wiki
    May 7, 2021 · Root exudation is the process of chemical excretion from the roots of plants as a means of interaction with the other organisms in soil. ... “Root ...Missing: definition | Show results with:definition
  4. [4]
  5. [5]
  6. [6]
    The Rhizosphere - Roots, Soil and Everything In Between - Nature
    Root exudates include both secretions (including mucilage) which are actively released from the root and diffusates which are passively released due to osmotic ...The Rhizosphere Defined · Root Exudates And Mineral... · Mycorrhizal Fungi And...
  7. [7]
    Are root exudates more important than other sources of ...
    At root apices, rhizodeposits originate from mucilages, exudates, and sloughed-off root cells/tissues that may or may not lyse their contents in the vicinity.
  8. [8]
    [PDF] PEA-ROOT EXUDATES AND THEIR EFFECT UPON ... - WUR eDepot
    A specific effect caused by root exudates was first demonstrated by O'BRIEN. andPRENTICE(1930). ... The amounts, identity and mutual proportion of compounds in ...
  9. [9]
    Plant root exudates | The Botanical Review
    Use of radioisotopes to investigate the sites of exudation along roots and the role of lateral roots in the exudation process should enable us to determine what ...
  10. [10]
    Root Exudates: Mechanistic Insight of Plant Growth Promoting ...
    The components of the root exudates are associated with the microbial population, notably, plant growth-promoting rhizobacteria (PGPR).
  11. [11]
    Plant exudates-driven microbiome recruitment and assembly ...
    Root tip exudates are rich in organic acids (such as citric acid and malic acid), plant hormones, and proteins, facilitating the regulation of rhizosphere ...
  12. [12]
    Root exudate stoichiometry is a key driver of soil N cycling
    Exudates consist of sugars, amino acids, organic acids (Smith, 1976) and numerous secondary metabolites (Salem et al., 2022), which are released both passively ...
  13. [13]
    Root Exudates Mediate the Processes of Soil Organic Carbon Input ...
    Jan 31, 2023 · Gross rhizodeposition carbon accounts for 5–20% of photosynthates ... Therefore, root functional traits determine the carbon allocation of root ...
  14. [14]
    Root exudation of sugars, amino acids, and organic acids by maize ...
    Dec 22, 2010 · Root exudation of sugars, amino acids, and organic acids by maize as affected by nitrogen, phosphorus, potassium, and iron deficiency.Missing: quantity | Show results with:quantity
  15. [15]
    Analytical challenges and solutions for performing metabolomic ...
    This paper critically reviews the most relevant sampling strategies, analytical methodologies, and data-mining approaches to study root exudates.
  16. [16]
    A standardized method for the sampling of rhizosphere and ...
    Jun 21, 2012 · 2007) where the bacterial density was 10 to 100 fold higher in rhizosphere as compared to bulk soil.Missing: thickness | Show results with:thickness
  17. [17]
    Rhizosphere - an overview | ScienceDirect Topics
    Hiltner coined the term 'rhizosphere' in 1904 to describe the zone of soil under the influence of plant roots. His observations were initially based on the ...
  18. [18]
    The role of the rhizobiome recruited by root exudates in plant ...
    Nov 16, 2024 · Root exudates serve as a bridge connecting plant roots and rhizosphere microbes, playing a key role in influencing the assembly and function of the rhizobiome.
  19. [19]
    Organic Acids Regulation of Chemical–Microbial Phosphorus ...
    Citric and oxalic acids, common root exudates, induced a strong mobilization of soil P, but only at doses considered to be much greater than those found in most ...Materials And Methods · Soil Samples And... · Ph And Phosphorus...
  20. [20]
    Root exudate-derived compounds stimulate the phosphorus ...
    Mar 10, 2023 · However, root exudates supplemented to the different bacterial appeared to enhance P solubilizing activity and overall P availability. Threonine ...
  21. [21]
    Benzoxazinoids in Root Exudates of Maize Attract Pseudomonas ...
    This so-called rhizosphere effect supports bacterial cell densities in the root vicinity up to 100-fold greater than in surrounding soil [2]. The chemical ...
  22. [22]
    Root exudates: The rhizospheric frontier for advancing sustainable ...
    ISR is a plant-wide defense response triggered by beneficial rhizosphere microbes, conferring protection against diverse pathogens and pests. REs contribute to ...
  23. [23]
    Living and decaying roots as regulators of soil aggregation and ...
    In addition, root exudates (e.g., root mucilage or polysaccharide exudates) can either adhere directly to mineral surfaces or function as glue-like binding ...
  24. [24]
    [PDF] The Dynamic Effect of Root Exudates on Soil Structure - EGUsphere
    Nov 10, 2023 · Mucilage is known to promote a hysteretic behavior of soil water retention during WDC, resulting in higher water content in the rhizosphere ...
  25. [25]
    Harnessing root exudates for plant microbiome engineering and ...
    Plant roots exude up to 40% of their photosynthetically derived carbon, with these compounds ranging from small (e.g., carbohydrates, amino acids, organic ...
  26. [26]
    A pulse of simulated root exudation alters the composition and ...
    Root exudation might have caused local oxygen depletion and anaerobe metabolism. Abstract. Root exudation increases the concentration of readily available ...
  27. [27]
    Root Exudation of Primary Metabolites: Mechanisms and Their ...
    Feb 20, 2019 · Root exudation is an important process determining plant interactions with the soil environment. Many studies have linked this process to soil nutrient ...
  28. [28]
    An ABC Transporter Mutation Alters Root Exudation of ... - NIH
    In this study, we showed that a single gene mutation (ABC transporter, abcg30) involved in root exudation influences the soil microbial community. We analyzed ...
  29. [29]
    Aluminum-activated root malate and citrate exudation is ... - NIH
    In Arabidopsis, aluminum (Al) exclusion from the root is mainly facilitated by Al-activated root malate and citrate exudation through the ALMT1 malate
  30. [30]
    Root Exudation of Primary Metabolites: Mechanisms and Their ...
    Feb 21, 2019 · The bottom panel represents passive transport mechanisms, which allow diffusion following electrochemical gradients. Red arrows represent ...
  31. [31]
    Root hairs increase rhizosphere extension and carbon input to soil
    Dec 18, 2017 · Root hairs increase exudation and spatial rhizosphere extension, which probably enhance rhizosphere interactions and nutrient cycling in larger soil volumes.
  32. [32]
    Root Exudation: An In-Depth Experimental Guide - Authorea
    Nov 15, 2024 · We discuss state-of-the art of spatially resolved exudate collection, collection in controlled vs field conditions, and plant growth setups ...
  33. [33]
    Root Exudates Regulate Soil Fungal Community Composition and ...
    This study establishes root exudates as a mechanism through which a plant is able to regulate soil fungal community composition.<|control11|><|separator|>
  34. [34]
    Estimating the global root exudate carbon flux | Biogeochemistry
    Jul 9, 2024 · We estimate a global root exudate flux of 13.4 (10.1–20.2) Pg C y −1 , or about 9% (7–14%) of global annual gross primary productivity.Missing: oxygen depletion
  35. [35]
    Regulation and function of root exudates
    ### Summary of Regulation of Root Exudates (Badri & Vivanco, 2009)
  36. [36]
    The plant circadian clock influences rhizosphere community ... - Nature
    Oct 20, 2017 · Our findings suggest the circadian clock of the plant host influences rhizosphere community structure and function.
  37. [37]
    Understanding the Mechanisms of Fe Deficiency in the Rhizosphere ...
    May 10, 2023 · Under conditions of severe Fe shortage, phytosiderophores may account for 50–90% of the exudates secreted at the root tip (Fan et al., 1997, as ...
  38. [38]
    Effect of drought on root exudates from Quercus petraea and ...
    May 10, 2022 · Experimental evidence has demonstrated decreased root exudation under dry conditions, which can lead to a change in enzyme activity.
  39. [39]
    Root exudates drive the soil-borne legacy of aboveground pathogen ...
    Sep 12, 2018 · Our data suggest that plants can recruit beneficial rhizosphere communities via modification of plant exudation patterns in response to exposure to aboveground ...
  40. [40]
    A multidrug and toxic compound extrusion (MATE) transporter ...
    Nov 1, 2019 · Exudation of citrate from Arabidopsis roots is a Al-tolerance response that involves the STOP1-mediated induction of AtMATE (J. Liu ...Abstract · INTRODUCTION · RESULTS · DISCUSSION
  41. [41]
    Plant health: feedback effect of root exudates-rhizobiome interactions
    Root exudates aid the rhizobiome, which is crucial for plant growth, nutrition, and health. The rhizobiome provides support for optimum plant growth and ...
  42. [42]
    Kin recognition in an annual plant | Biology Letters - Journals
    Jun 13, 2007 · Plants recognize kin by allocating less root mass when competing with kin, indicating root interactions are the cue for kin recognition.
  43. [43]
    Root exudates mediate kin recognition in plants - PMC - NIH
    The results demonstrate that that kin recognition and self/non-self are two separate identity recognition systems involving soluble chemicals.
  44. [44]
    Environmental strigolactone drives early growth responses to ...
    Aug 22, 2022 · We conclude that plants both exude SLs as signals to modulate neighbor growth and detect environmental SLs as a cue for neighbor presence; ...
  45. [45]
    Mechanisms, ecology and agricultural aspects of kin recognition in ...
    Kin recognition thus likely has important implications for evolution of plant traits, diversity of plant populations, ecological networks and community ...
  46. [46]
    The role of ABC transporters in kin recognition in Arabidopsis thaliana
    This study investigated the role of ABC transporters in kin recognition in A. thaliana based on previous evidence that root secretions are involved in the kin ...Introduction · Method And Materials · Abc Transporter Inhibitor...<|separator|>
  47. [47]
    The Role of Flavonoids in Nodulation Host-Range Specificity
    Aug 11, 2016 · To act as nodulation signals flavonoids must be secreted from the roots into the rhizosphere, which includes the root surface and inside ...
  48. [48]
    The role of flavonoids in the establishment of plant roots ... - NIH
    Flavonoids play an essential role in rhizobium-legume symbiosis as chemoattractant and nod gene inducers. They are suggested to act on mycorrhization.
  49. [49]
    Strigolactones Stimulate Arbuscular Mycorrhizal Fungi by Activating ...
    Jun 27, 2006 · Strigolactones are important rhizospheric plant signals involved in stimulating both the pre-symbiotic growth of AM fungi and the germination of parasitic ...
  50. [50]
    Variation in Root Exudate Composition Influences Soil Microbiome ...
    Our experimental results demonstrate how soil microbial community and genomic diversity is influenced by root exudates of differing chemical compositions.<|control11|><|separator|>
  51. [51]
    Root exudates drive interspecific facilitation by enhancing ... - NIH
    May 23, 2016 · We found that faba bean/maize intercropping enhances productivity, nodulation, and N 2 fixation of faba bean through interspecific root interactions.
  52. [52]
    Phosphorus and iron deficiencies induce a metabolic ...
    Jul 17, 2015 · Highlight. Fe and P deficiencies affect strawberry root metabolism, inducing citrate release and proton extrusion.
  53. [53]
    Microbial gross organic phosphorus mineralization can be ...
    Phosphatase activity was increased due to addition of root exudates up to a factor of 6. •. P uptake by microorganisms was only weakly and not consistently ...
  54. [54]
    Quantifying citrate-enhanced phosphate root uptake using ...
    Dec 5, 2019 · Organic acid exudation by plant roots is thought to promote phosphate (P) solubilisation and bioavailability in soils with poorly available ...Materials And Methods · Modelling And Data Fitting · Discussion
  55. [55]
    Integrated Microbiome and Metabolomic Analysis Reveal ... - Rice
    Apr 11, 2023 · It is estimated that 5–21% of the carbon fixed by photosynthesis or 15–25% of the carbon allocated to the root system enters the soil as root ...
  56. [56]
    Soil Management for Increased Soil Organic Matter (G2283)
    Root growth contributes to the increase of SOM by producing organic exudates, stimulating microbial activity and turnover, and by adding carbon in root tissues.
  57. [57]
    Root exudate signals in plant-plant interactions - PubMed
    Oct 7, 2020 · Root exudates mediate root detection and behaviour, kin recognition, flowering and production, driving inter- and intra-specific facilitation in cropping ...
  58. [58]
    Allelopathy and its application as a weed management tool: A review
    Nov 27, 2022 · The allelochemicals are released from plant parts by leaching from leaves or litter on the ground, root exudation, volatilization from leaves, ...
  59. [59]
    Juglone and allelopathy | Journal of Chemical Education
    It is well established that the black walnut tree and the tomato plant are involved allelopathically and that juglone is the active principle.
  60. [60]
    (PDF) Allelopathy in black walnut (Juglans nigra L.) alley cropping. I ...
    Aug 6, 2025 · A study was conducted to quantify the spatial and temporal variation in soil juglone (5-hydroxy-1,4-naphthoquinone) in a 10-year-old black walnut (Juglans ...
  61. [61]
    Allelopathic potential of Sorghum bicolor L. Root Exudates on ...
    The growth of duckweed (Lemna minor) was inhibited by 50. μM sorgoleone in liquid culture [9]. As well as the seedling growth of Amaranthus retroflexus, ...
  62. [62]
    Primed to grow: a new role for green leaf volatiles in plant stress ...
    Green leaf volatiles (GLV) have been well described to prime plants against biotic and abiotic stresses resulting in an accelerated and/or enhanced protective ...
  63. [63]
    (PDF) Exploring the Volatiles Released from Roots of Wild and ...
    Apr 24, 2025 · Here, we investigated if tomato plants under insect herbivore attack exhibited a different root volatilome than non-stressed plants, and ...
  64. [64]
    Herbivory-induced green leaf volatiles increase plant performance ...
    May 1, 2025 · Here we demonstrate that volatiles that are released by herbivore-attacked leaves trigger plant-soil feedbacks, resulting in increased performance of different ...Missing: stress tomato
  65. [65]
    The genetics underlying natural variation of plant–plant interactions ...
    Dec 12, 2017 · Root exudates also play a key role in the interaction of crops with parasitic plants. Resistance to Striga parasitic plants in sorghum cultivars ...