Soil organisms are recognized as ecosystem engineers and key for aggregation in soil due to bioturbation, organic matter (OM) decomposition, and excretion of biogenic OM. The activity of soil organisms is beneficial for soil quality, functions, and nutrient cycling. These attributions are based on field-scale observations that link the presence and activity of organisms to spatiotemporal changes in soil properties and can be traced back to the formation of biogenic aggregates. This biogenic formation pathway encompasses a cascade of processes so far not discussed comprehensively. A more general approach needs to consider the activity and feedback loops between soil biota, the active release of biogenic OM by excretion, the interaction of biogenic OM with soil constituents, the formation of organo-mineral associations, and how these become incorporated in aggregated structures. Especially the function of biogenically excreted OM, which is quite complex in composition, is controversial as it permits or inhibits aggregation. This review analyzes the various roles of biogenically excreted OM may take as an aggregation agent. We will show that its function depends on the interplay of numerous factors, including environmental conditions, variety of OM producers, composition and availability of biogenically excreted OM, and type of interacting mineral phase. We consider biogenically excreted OM to affect aggregate formation in three different ways: (I) as a bridging agent which promotes the aggregation due to surface modifications and attraction, (II) as a separation agent which favors the formation, mobility, and transport of organo-mineral associations and inhibits their further inclusion into aggregates, and (III) as a gluing agent which mediates aggregate stability, after an external force provokes a close approach of soil particles. We conclude that biogenically excreted OM takes these functional roles simultaneously and to a varying extent across spatiotemporal scales. Hence, biogenically excreted OM is involved in the surface modification of soil particles, in the enmeshment and gluing of particles into soil aggregates, in the (im-)mobilization, and in facilitating the transport of particles. All that depends on the interplay of a hierarchy of factors comprising the local soil community's composition, the properties of biogenically excreted OM, and the conditions of the immediate environment. • Cross-scale consideration of the influence of biogenic organic matter on aggregation. • Mineral interactions with mucus, extracellular polymeric substances, and root exudates. • Critical discussion of biogenic organic matter as aggregation promoting and inhibiting agent. • Function of biogenic organic matter as bridging, separation, and gluing agent. • Biogenically induced aggregation is governed by the interplay of different species. 

Pathways of biogenically excreted organic matter into soil aggregates

Soil organisms are recognized as ecosystem engineers and key for aggregation in soil due to bioturbation, organic matter (OM) decomposition, and excretion of biogenic OM. The activity of soil organisms is beneficial for soil quality, functions, and nutrient cycling. These attributions are based on field-scale observations that link the presence and activity of organisms to spatiotemporal changes in soil properties and can be traced back to the formation of biogenic aggregates. This biogenic formation pathway encompasses a cascade of processes so far discussed not comprehensively. A more general approach needs to consider the activity and feedback loops between soil biota, the active release of biogenic OM by excretion, the interaction of biogenic OM with soil constituents, the formation of organo-mineral associations, and how these become incorporated in aggregated structures. Especially the function of biogenically excreted OM, which is quite complex in composition, is controversial as it permits or inhibits aggregation. This review analyzes the various roles of biogenically excreted OM may take as an aggregation agent. We will show that its function depends on the interplay of numerous factors, including environmental conditions, variety of OM producers, composition and availability of biogenically excreted OM, and type of interacting mineral phase. We consider biogenically excreted OM to affect aggregate formation in three different ways: (I) as a bridging agent which promotes the aggregation due to surface modifications and attraction, (II) as a separation agent which favors the formation, mobility, and transport of organo-mineral associations and inhibits their further inclusion into aggregates, and (III) as a gluing agent which mediates aggregate stability, after an external force provokes a close approach of soil particles. We conclude that biogenically excreted OM takes these functional roles simultaneously and to a varying extent across spatiotemporal scales. Hence, biogenically excreted OM is involved in the surface modification of soil particles, in the enmeshment and gluing of particles into soil aggregates, in the (im-)mobilization, and in facilitating the transport of particles. All that depends on the interplay of a hierarchy of factors comprising the local soil community's composition, the properties of biogenically excreted OM, and the conditions of the immediate environment.

https://digital.csic.es/bitstream/10261/244975/1/1-s2.0-S2214158821000179-main.pdf

Analytical challenges and solutions for performing metabolomic analysis of root exudates

Plants are in constant interaction with a myriad of soil microorganisms in the rhizosphere, an area of soil in close contact with plant roots. Recent research has highlighted the importance of plant-specialized metabolites (PSMs) in shaping and modulating the rhizosphere microbiota; however, the molecular mechanisms underlying the establishment and function of the microbiota mostly remain unaddressed. Flavonoids and saponins are a group of PSMs whose biosynthetic pathways have largely been revealed. Although these PSMs are abundantly secreted into the rhizosphere and exert various functions, the secretion mechanisms have not been clarified. This review summarizes the roles of flavonoids and saponins in the rhizosphere with a special focus on interactions between plants and the rhizosphere microbiota. Furthermore, this review introduces recent advancements in the dynamics of these metabolites in the rhizosphere and indicates potential applications of PSMs for crop production and discusses perspectives in this emerging research field.

Flavonoids and saponins in plant rhizospheres: roles, dynamics, and the potential for agriculture.

Global agriculture is challenged with achieving sustainable food security while the climate changes and the threat of drought increases. Much of the research attention has focused on above-ground plant responses with an aim to improve drought resistance. The hidden half, that is, the root system belowground, is receiving increasing attention as the interface of the plant with the soil. Because roots are a sensing organ for nutrients and moisture, we speculate that crop root system traits can be managed using smart nutrient application in order to increase drought resistance. Roots are known to be influenced both by their underlying genetics and also by responses to the environment, termed root plasticity. Though very little is known about the combined effect of water and nutrients on root plasticity, we explore the possibilities of root system manipulation by nutrient application. We compare the effects of different water or nutrient levels on root plasticity and its genetic regulation, with a focus on how this may affect drought resistance. We propose four primary mechanisms through which smart nutrient management can optimize root traits for drought resistance: (1) overall plant vigor, (2) increased root allocation, (3) influence specific root traits, and (4) use smart placement and timing to encourage deep rooting. In the longer term, we envision that beneficial root traits, including plasticity, could be bred into efficient varieties and combined with advanced precision management of water and nutrients to achieve agricultural sustainability.

Can smart nutrient applications optimize the plant's hidden half to improve drought resistance?

Root morphology and exudation define a plants' sphere of influence in soils. In turn, soil characteristics influence plant growth, morphology, root microbiome, and rhizosphere chemistry. Collectively, all these parameters have significant implications on the major biogeochemical cycles, crop yield, and ecosystem health. However, how plants are shaped by the physiochemistry of soil particles is still not well understood. We explored how particle size and chemistry of growth substrates affect root morphology and exudation of a model grass. We grew Brachypodium distachyon in glass beads with various sizes (0.5, 1, 2, 3 mm), as well as in sand (0.005, 0.25, 4 mm) and in clay (4 mm) particles and in particle-free hydroponic medium. Plant morphology, root weight, and shoot weight were measured. We found that particle size significantly influenced root fresh weight and root length, whereas root number and shoot weight remained constant. Next, plant exudation profiles were analyzed with mass spectrometry imaging and liquid chromatography-mass spectrometry. Mass spectrometry imaging suggested that both, root length and number shape root exudation. Exudate profiles were comparable for plants growing in glass beads or sand with various particles sizes, but distinct for plants growing in clay for in situ exudate collection. Clay particles were found to sorb 20% of compounds exuded by clay-grown plants, and 70% of compounds from a defined exudate medium. The sorbed compounds belonged to a range of chemical classes, among them nucleosides, organic acids, sugars, and amino acids. Some of the sorbed compounds could be desorbed by a rhizobacterium (Pseudomonas fluorescens WCS415), supporting its growth. This study demonstrates the effect of different characteristics of particles on root morphology, plant exudation and availability of nutrients to microorganisms. These findings further support the critical importance of the physiochemical properties of soils when investigating plant morphology, plant chemistry, and plant-microbe interactions.

/pdf/root-morphology-and-exudate-availability-are-shaped-by-56w7kwpn55.pdf

Root morphology and exudate availability are shaped by particle size and chemistry in Brachypodium distachyon.

Root morphology and exudation define a plants sphere of influence in soils, and are in turn shaped by the physiochemical characteristics of soil. We explored how particle size and chemistry of growth substrates affect root morphology and exudation of the model grass Brachypodium distachyon. Root fresh weight and root lengths were correlated with particle size, whereas root number and shoot weight remained constant. Mass spectrometry imaging suggested that both, root length and number shape root exudation. Exudate metabolite profiles detected with liquid chromatography / mass spectrometry were comparable for plants growing in glass beads or sand with various particles sizes, but distinct for plants growing in clay. However, when exudates of clay-grown plants were collected by removing the plants from the substrate, their exudate profile was similar to sand- or glass beads-grown plants. Clay particles sorbed 20% of compounds exuded by clay-grown plants, and 70% of compounds of a defined exudate medium. The sorbed compounds belonged to a range of chemical classes, among them nucleosides/nucleotides, organic acids, sugars, and amino acids. Some of the sorbed compounds could be de-sorbed by a rhizobacterium (Pseudomonas fluorescens WCS415), supporting its growth. We show that root morphology is affected by substrate size, and that root exudation in contrast is not affected by substrate size or chemistry. The availability of exuded compounds, however, depends on the substrate present. These findings further support the critical importance of the physiochemical properties of soils are crucial to consider when investigating plant morphology, exudation, and plant-microbe interactions.

https://biorxiv.org/content/biorxiv/early/2019/05/27/651570.full-text.pdf

Katherine Whiting

Papers

Root morphology and exudate availability are shaped by particle size and chemistry in Brachypodium distachyon.

Root morphology and exudate availability is shaped by particle size and chemistry in Brachypodium distachyon