Coarse and fine root plants affect pore size distributions differently

Aims

Small scale root-pore interactions require validation of their impact on effective hydraulic processes at the field scale. Our objective was to develop an interpretative framework linking root effects on macroscopic pore parameters with knowledge at the rhizosphere scale.

Methods

A field experiment with twelve species from different families was conducted. Parameters of Kosugi’s pore size distribution (PSD) model were determined inversely from tension infiltrometer data. Measured root traits were related to pore variables by regression analysis. A pore evolution model was used to analyze if observed pore dynamics followed a diffusion like process.

Results

Roots essentially conditioned soil properties at the field scale. Rooting densities higher than 0.5 % of pore space stabilized soil structure against pore loss. Coarse root systems increased macroporosity by 30 %. Species with dense fine root systems induced heterogenization of the pore space and higher micropore volume. We suggested particle re-orientation and aggregate coalescence as main underlying processes. The diffusion type pore evolution model could only partially capture the observed PSD dynamics.

Conclusions

Root systems differing in axes morphology induced distinctive pore dynamics. Scaling between these effective hydraulic impacts and processes at the root-pore interface is essential for plant based management of soil structure.     

Does biochar influence soil physical properties and soil water availability?

Aims

This study aims to (i) determine the effects of incorporating 47 Mg ha^?1 acacia green waste biochar on soil physical properties and water relations, and (ii) to explore the different mechanisms by which biochar influences soil porosity.

Methods

The pore size distribution of the biochar was determined by scanning electron microscope and mercury porosimetry. Soil physical properties and water relations were determined by in situ tension infiltrometers, desorption and evaporative flux on intact cores, pressure chamber analysis at ?1,500 kPa, and wet aggregate sieving.

Results

Thirty months after incorporation, biochar application had no significant effect on soil moisture content, drainable porosity between –1.0 and ?10 kPa, field capacity, plant available water capacity, the van Genuchten soil water retention parameters, aggregate stability, nor the permanent wilting point. However, the biochar-amended soil had significantly higher near-saturated hydraulic conductivity, soil water content at ?0.1 kPa, and significantly lower bulk density than the unamended control. Differences were attributed to the formation of large macropores (>1,200 μm) resulting from greater earthworm burrowing in the biochar-amended soil.

Conclusion

We found no evidence to suggest application of biochar influenced soil porosity by either direct pore contribution, creation of accommodation pores, or improved aggregate stability.     

The effect of phosphorus on cluster-root formation and functioning of Embothrium coccineum (R. et J. Forst.)

Background and aims

Embothrium coccineum (R. et J. Forst.) is a Proteaceae species from the southern part of South America. South-central Chilean soils are younger and contain more phosphorus (P) than soils in Australia and South Africa, where Proteaceae are common. Phosphorus deficiency is the main factor promoting cluster-root formation in Proteaceae. It is not known, however, whether this also applies to E. coccineum, which grows on soils with higher P content.

Methods

Four-month-old seedlings were grown for 4 weeks in hydroponic cultures with 1 μM P or 50 μM P. The number of cluster roots, relative height increment, biomass distribution, cluster root/total root biomass ratio, foliar P concentration, root acid phosphatase activity and root carboxylate-exudation rates were determined.

Results

Seedlings growing at 50 μM P showed a 10?, 1.3? and 3.3-fold greater increase in relative height, total dry mass and foliar P concentration, respectively, compared with those grown at1 μM P. However, seedlings grown at 1 μM P showed a 5?, 16?, 1.7? and 1.3-fold greater number of cluster roots, cluster root/total root biomass ratio, phosphatase activity and total carboxylate exudation, respectively, as compared with those grown at 50 μM P.

Conclusions

A low P supply promotes the initiation, growth and metabolic activity of cluster roots which is in accordance with reports on Proteaceae species occurring in ancient and highly weathered soils.     

Root growth in biopores—evaluation with in situ endoscopy

Background and aims

The significance of biopores for nutrient acquisition from the subsoil depends on root-soil contact, which in turn is influenced by root architecture. The aim of this study was to detect differences regarding the architecture and root-soil contact of homorhizous barley roots (Hordeum vulgare L.) and allorhizous oilseed rape roots (Brassica napus L.) growing in biopores.

Methods

In situ endoscopy was used as a technique that allows non-destructive display of pore wall characteristics and root morphology inside large biopores under field conditions.

Results

For both crops, about 85 % of all roots did establish contact to the pore wall. However, according to their different root architecture, the two crops varied in their strategy of resource acquisition: While barley was characterized by thin vertical or ingrowing roots, most of them in direct contact to the pore wall, oilseed rape established contact to the pore wall predominantly via lateral roots.

Conclusions

Root morphological and pore wall assessment with in situ endoscopy in combination with detailed studies of soil biochemical and soil physical parameters of the pore wall is considered an essential prerequisite for more precise future modelling of nutrient acquisition and uptake.     

Effects of different plant root exudates and their organic acid components on chemotaxis, biofilm formation and colonization by beneficial rhizosphere-associated bacterial strains

Aim

It is necessary to understand the roles of root exudates involved in plant-microbe interactions to inform practical application of beneficial rhizosphere microbial strains.

Methods

Colonization of Bacillus amyloliquefaciens SQR9 (isolated from cucumber rhizosphere) and Bacillus subtilis N11 (isolated from banana rhizosphere) of their original host was found to be more effective as compared to the colonization of the non-host plant. Organic acids in the root exudates of the two plants were identified by High performance liquid chromatography (HPLC). The chemotactic response and effects on biofilm formation were assessed for SQR9 and N11 in response to cucumber and banana root exudates, as well as their organic acids components.

Results

Citric acid detected exclusively in cucumber exudates could both attract SQR9 and induce its biofilm formation, whereas only chemotactic response but not biofilm formation was induced in N11. Fumaric acid that was only detected in banana root exudates revealed both significant roles on chemotaxis and biofilm formation of N11, while showing only effects on biofilm formation but not chemotaxis of SQR9.

Conclusion

The relationship between PGPR strain and root exudates components of its original host might contribute to preferential colonization. This study advances a clearer understanding of the mechanisms relevant to application of PGPR strains in agricultural production.     

Phenotypic seedling responses of a metal-tolerant mutant line of sunflower growing on a Cu-contaminated soil series: potential uses for biomonitoring of Cu exposure and phytoremediation

Background and aims

The potential use of a metal-tolerant sunflower mutant line for both biomonitoring and phytoremediating a Cu-contaminated soil series was investigated.

Methods

The soil series (21–1,170 mg Cu kg^?1) was sampled in field plots at control and wood preservation sites. Sunflowers were cultivated 1 month in potted soils under controlled conditions.

Results

pH and dissolved organic matter influenced Cu concentration in the soil pore water. Leaf chlorophyll content and root growth decreased as Cu exposure rose. Their EC₁₀ values corresponded to 104 and 118 μg Cu L^?1 in the soil pore water, 138 and 155 mg Cu kg^?1 for total soil Cu, and 16–18 mg Cu kg^?1 DW shoot. Biomass of plant organs as well as leaf area, length and asymmetry were well correlated with Cu exposure, contrary to the maximum stem height and leaf water content.

Conclusions

Physiological parameters were more sensitive to soil Cu exposure than the morphological ones. Bioconcentration and translocation factors and distribution of mineral masses for Cu highlighted this mutant as a secondary Cu accumulator. Free Cu²⁺ concentration in soil pore water best predicted Cu phytoavailability. The usefulness of this sunflower mutant line for biomonitoring and Cu phytoextraction was discussed.     

Microdistribution of major to trace elements between roots of Halimione portulacoides and host sediments (Tagus estuary marsh,Portugal)

Aim

This study presents a micrometre-scale map of the elemental distribution within roots and surrounding sediment of Halimione portulacoides of a contaminated salt marsh in the Tagus estuary.

Methods

Microprobe particle induced X-ray emission analysis was performed in sediment slices containing roots with tubular rhizoconcretions attached to host sediments.

Results

Strong concentration gradients were found particularly in the inner part of rhizoconcretions adjacent to the root wall. Local enrichment was observed in sediment interstices with Fe precipitates and other associated elements. A maximum of 55 % of Fe was measured near the concretion–root interface, with a decrease to <5 % in the host sediment. Maximum concentrations of P (3 %), As (1,200 μg g^?1) and Zn (3,000 μg g^?1) were registered in concretions, one order of magnitude above the values of the host sediment. The elemental concentration profiles across roots showed that the epidermis was an efficient selective barrier to the entrance of elements. Fe and As were retained in the epidermis. The highest Cu and Zn concentrations were also observed in the epidermis. However, the concentrations of Mn, Cu and Zn increased in the inner root.

Conclusions

As and Fe were mostly retained in the concretion, whereas P, Mn, Cu and Zn were mobilised by the root.     

Estimating the absorptive root area in Norway spruce by using the common direct and indirect earth impedance methods

Aims

Estimates of root absorption magnitude are needed for the balanced management of forest ecosystems, but no methods able to work on the whole tree and stand level were available. Modified earth impedance method was developed recently and here it was tested, by comparing the results with those obtained by combination of several classical methods.

Methods

We used direct (soil cores, scanning and microscopy) and indirect (sap flow patterns and modified earth impedance) methods in an attempt to estimate the absorptive root area indexes (RAI) at two sites of about 25 and 40-years-old Norway spruce. We considered the geometric surfaces of all scanned fine roots to be equal to the fine root absorptive area (RAI _scan ). To estimate the potentially physically permeable area of fine roots, we microscopically evaluated the point of secondary xylem appearance and calculated the geometric area of root portions with primary structure (RAI _micro ). We termed the area of electrically conductive root surface as the active (ion) absorptive area (RAI _mei ) and measured its extent by the modified earth impedance (MEI) method.

Results

The highest values for absorptive root areas at the two experimental sites we obtained with the scanning method (RAI _scan was considered to be 100%), followed by the RAI _micro (51%) and RAI _mei (32%). RAImei reached about 2/3 of RAImicro. The surface area of the ectomycorrhizal hyphae was an order of magnitude larger than that of all fine roots, but the MEI did not measure such increase.

Conclusions

We showed that the absorptive root area, indirectly estimated by the MEI, provides consistent results that approach the values obtained for fine roots with a primary structure estimated by traditional direct methods. The similar range of the values for the absorptive root surface area obtained by microscopy and by the MEI method indicates that this method is feasible and that it could be used to determine the extent of active absorptive root surface areas in forests.     

Pattern of antioxidant enzyme activities and hydrogen peroxide content during developmental stages of rhizogenesis from hypocotyl explants of Mesembryanthemum crystallinum L.

Key message

H ₂ O ₂ is necessary to elicit rhizogenic action of auxin. Activities of specific catalase and manganese superoxide dismutase forms mark roots development.

Abstract

Hypocotyl explants of Mesembryanthemum crystallinum regenerated roots on medium containing 2,4-dichlorophenoxyacetic acid. Explants became competent to respond to the rhizogenic action of auxin on day 3 of culture, when hydrogen peroxide content in cultured tissue was the highest. l-Ascorbic acid added to the medium at 5 μM lowered the H₂O₂ level, inhibited rhizogenesis and induced non-regenerative callus, suggesting that certain level of H₂O₂ is required to promote root initiation. Coincident with the onset of rhizogenic determination, meristemoids formed at the periphery of the hypocotyl stele and the activity of the manganese form of superoxide dismutase, MnSOD-2 was induced. Once induced, MnSOD-2 activity was maintained through the post-determination phase of rooting, involving root growth. MnSOD-2 activity was not found in non-rhizogenic explants maintained in the presence of AA. Analyses of the maximum photochemical efficiency of photosystem II and the oxygen uptake rate revealed that the explants were metabolically arrested during the predetermination stage of rhizogenesis. Respiratory and photosynthetic rates were high during root elongation and maturation. Changes in catalase and peroxidase activities correlated with fluctuations of endogenous H₂O₂ content throughout rhizogenic culture. Expression of a specific CAT-2 form accompanied the post-determination stage of rooting and a high rate of carbohydrate metabolism during root growth. On the other hand, the occurrence of MnSOD-2 activity did not depend on the metabolic status of explants. The expression of MnSOD-2 activity throughout root development seems to relate it specifically to root metabolism and indicates it as a molecular marker of rhizogenesis in M. crystallinum.     

Macropore effects on phosphorus acquisition by wheat roots – a rhizotron study

Background and aims

Macropores may be preferential root pathways into the subsoil. We hypothesised that the presence of macropores promotes P-uptake from subsoil, particularly at limited water supply in surface soil. We tested this hypothesis in a rhizotron experiment with spring wheat (Triticum aestivum cv. Scirocco) under variation of fertilisation and irrigation.

Methods

Rhizotrons were filled with compacted subsoil (bulk density 1.4 g cm^?3), underneath a P-depleted topsoil. In half of these rhizotrons the subsoil contained artificial macropores. Spring wheat was grown for 41 days with and without irrigation and ³¹P–addition. Also, a ³³P–tracer was added at the soil surface to trace P-distribution in plants using liquid scintillation counting and radioactive imaging.

Results

Fertilisation and irrigation promoted biomass production and plant P-uptake. Improved growing conditions resulted in a higher proportion of subsoil roots, indicating that the topsoil root system additionally promoted subsoil nutrient acquisition. The presence of macropores did not improve plant growth but tended to increase translocation of ³³P into both above- and belowground biomass. ³³P–imaging confirmed that this plant-internal transport of topsoil-P extended into subsoil roots.

Conclusions

The lack of penetration resistance in macropores did not increase plant growth and nutrient uptake from subsoil here; however, wheat specifically re-allocated topsoil-P for subsoil root growth.

     

A new model for root growth in soil with macropores

Background and aims

The use of standard dynamic root architecture models to simulate root growth in soil containing macropores failed to reproduce experimentally observed root growth patterns. We thus developed a new, more mechanistic model approach for the simulation of root growth in structured soil.

Methods

In our alternative modelling approach, we distinguish between, firstly, the driving force for root growth, which is determined by the orientation of the previous root segment and the influence of gravitropism and, secondly, soil mechanical resistance to root growth. The latter is expressed by its inverse, soil mechanical conductance, and treated similarly to hydraulic conductivity in Darcy’s law. At the presence of macropores, soil mechanical conductance is anisotropic, which leads to a difference between the direction of the driving force and the direction of the root tip movement.

Results

The model was tested using data from the literature, at pot scale, at macropore scale, and in a series of simulations where sensitivity to gravity and macropore orientation was evaluated.

Conclusions

Qualitative and quantitative comparisons between simulated and experimentally observed root systems showed good agreement, suggesting that the drawn analogy between soil water flow and root growth is a useful one.

     

Changes in root size and distribution in relation to nitrogen accumulation during maize breeding in China

Background and aims

Modern maize breeding has increased maize yields worldwide. The changes in above-ground traits accompanying yield improvement are well-known, but less information is available as to the effect of modern plant breeding on changes in maize root traits.

Methods

Root growth, nitrogen uptake, dry matter accumulation and yield formation of six maize hybrids released from 1973 to 2000 in China were compared. Experiments were conducted under low and high nitrogen supply in a black soil in Northeast China in 2010 and 2011.

Results

While nitrogen accumulation, dry matter production and yield formation have been increased, modern maize breeding in China since 1990 has reduced root length density in the topsoil without much effect on root growth in the deeper soil. The efficiency of roots in acquiring N has increased so as to match the requirement of N accumulation for plant growth and yield formation. The responses of root growth, nitrogen and dry matter accumulation, and grain yield to low-N stress were similar in the more modern hybrids as in the older ones.

Conclusions

Modern maize breeding has constitutively changed root and shoot growth and plant productivity without producing any specific enhancement in root responsiveness to soil N availability.     

Oxidative stress is associated with aluminum toxicity recovery in apex of pea root

Aims

Although many studies on the mechanism of Al toxicity and tolerance have been conducted independently, events occurring during the recovery process from Al injury is limited. This study was to investigate Al toxicity recovery mechanism focusing in morphological and physiological aspect.

Methods

We investigated the mechanisms underlying Al toxicity recovery in terms of oxidative stress using the pea root apex as a model system.

Results

The accumulation of reactive oxygen species was remarkably high in the root under continued Al treatment but decreased in the recovering root. The superoxide anion exuded in the presence of nicotinamide adenine dinucleotide phosphate (NADPH) showed a similar tendency with respect to the accumulation of reactive oxygen species. A similar pattern of lignin content and superoxide dismutase activity was observed among the treatments, while the increased peroxidation in the root under continued Al treatment did not decline with recovery treatment. A longitudinal section of the root under continued Al treatment showed the accumulation of superoxide anion, lignin and peroxide (H₂O₂) at the epidermal and outer cortex region where the Al induced injuries, including ruptures, are detected.

Conclusions

Oxidative stress is associated with the mechanism of Al toxicity recovery. The recovery process might include the elongation of the central cylinder as a consequence of the oxidative stress-induced formation of the zonal region (ZR). The results further suggest a plausible role for the ZR in the programmed cell death-like function involved in Al toxicity recovery.     

The root microbiota—a fingerprint in the soil?

Background

The root system of a plant is known to host a wide diversity of microbes that can be essential or detrimental to the plant. Microbial ecologists have long struggled to understand what factors structure the composition of these communities. An overlooked part of the microbial community succession in root systems has been the potential for individual variation among plants shaped by early colonisation events such as microbial exposure of the seed inside the parent plant and during dispersal.

Scope

In this review we outline life events of the plant that can affect the composition of its root microbiota and relate ecological theory of community assembly to the formation of the root microbiota.

Conclusion

All plants are exposed to environmental conditions and events throughout their lifetime that shape their phenotype. The microbial community associated with the plant is ultimately an extension of this phenotype. Therefore, only by following a plant from its origin inside the flower to senescence, can we fully understand how the associated microbial community was assembled and what determined its composition.     

The plasticity of the growth and proliferation of wheat root system under elevated CO2

Background and Aims

Understanding crop responses to increasing atmospheric CO₂ requires knowledge of how their root systems grow, proliferate and function. The effect of elevated CO₂ on the growth and proliferation of wheat root system (Triticum aestivum L.), was examined.

Methods

Two pairs of sister lines of wheat contrasting in vigour (CV97 and CV207) and tillering (7750N and 7750PF) were grown in rhizo-boxes under ambient (380 μl L^?1) and elevated CO₂ (700 μl L^?1), and the root growth and proliferation mapped.

Results

Elevated CO₂ effects on shoot and root biomass were observed in the lines contrasting for vigour, but not in the lines contrasting for tillering. Root biomass was reduced by 67 % in the high vigour line CV97, reducing total plant biomass by 26 % compared to the low vigour line, CV207. This was due to a reduction in root length down the 1 m soil profile and root proliferation in the top 0.2 m layer. The reduction in root biomass was not compensated by an increase in shoot biomass.

Conclusions

The reduction in root biomass under elevated CO₂ in the vigour line CV97 can be explained through its inability to increase the sink strength due to the failure to increase tiller number to which the plant presumably responded by increasing losses of the newly assimilated carbon by respiration.     

Superoxide production induced by short-term exposure of barley roots to cadmium, auxin, alloxan and sodium dodecyl sulfate

Key message

Abiotic stress-induced superoxide generation depending on its localization, level, duration and presumably also on the action of other signals may lead to different stress responses.

Abstract

The purpose of this study was to analyze the alterations in superoxide generation and morphogenesis following short-term Cd, IAA and alloxan treatments, during stress and recovery period in barley root tips. At low Cd concentration the transient accumulation of superoxide in the epidermal cells was accompanied by root growth inhibition and radial expansion of cortical cells in the elongation zone of root tips. These morphological changes were very similar to the externally applied IAA-induced responses. However, the role of superoxide generated in the epidermal cells by low concentration of Cd and IAA is probably alone not sufficient for the induction of these processes. SDS as an activator of NOX activity caused a strong accumulation of superoxide in the epidermal cells along the whole root apex but without any changes in root morphology and growth. On the other hand, higher Cd concentrations as well as alloxan stimulated the generation of superoxide in the cortical tissue of the elongation zone of root tip, which was accompanied by the induction of cell death. Our results suggest that enhanced superoxide generation, depending on its localization, level, duration and presumably also on the action of other signals, may lead to altered root morphology (15?μM Cd or IAA), root growth inhibition (alloxan), transient root growth cessation (30?μM Cd) or to the death of cells/root at higher (60?μM) Cd concentrations.     

Relationship between constitutive root aerenchyma formation and flooding tolerance in Zea nicaraguensis

Background and aims

The teosinte Zea nicaraguensis, which is adapted to frequently flooded lowlands, is considered a valuable germplasm resource for the development of flooding-tolerant maize. This species can form constitutive root aerenchyma under well-drained conditions. The objectives of this study were to screen Z. nicaraguensis accessions for the capacity to form constitutive aerenchyma, to obtain progeny with differing degrees of aerenchyma formation, and to compare the flooding tolerance of these progeny.

Methods

We evaluated constitutive aerenchyma formation in the root cortex of seedlings of eight accessions and several segregating populations of Z. nicaraguensis. We also evaluated flooding tolerance in lines selected for high or low degrees of constitutive aerenchyma formation.

Results

Seedlings of the eight accessions showed an extremely wide and continuous range of variation in aerenchyma formation. By phenotypic selection within two accessions, we obtained lines with either high or low degrees of constitutive aerenchyma formation. The lines selected for a higher degree of formation showed relatively high flooding tolerance evaluated by shoot dry weight ratio (flooded:control) than those with a lower degree of formation.

Conclusions

A greater capacity to form constitutive aerenchyma can enhance flooding tolerance.     

Quantifying the effect of soil moisture content on segmenting root system architecture in X-ray computed tomography images

Aims

A commonly accepted challenge when visualising plant roots in X-ray micro Computed Tomography (μCT) images is the similar X-ray attenuation of plant roots and soil phases. Soil moisture content remains a recognised, yet currently uncharacterised source of segmentation error. This work sought to quantify the effect of soil moisture content on the ability to segment roots from soil in μCT images.

Methods

Rice (Oryza sativa) plants grown in contrasting soils (loamy sand and clay loam) were μCT scanned daily for nine days whilst drying from saturation. Root volumes were segmented from μCT images and compared with volumes derived by root washing.

Results

At saturation the overlapping attenuation values of root material, water-filled soil pores and soil organic matter significantly hindered segmentation. However, in dry soil (ca. six days of drying post-saturation) the air-filled pores increased image noise adjacent to roots and impeded accurate visualisation of root material. The root volume was most accurately segmented at field capacity.

Conclusions

Root volumes can be accurately segmented from μCT images of undisturbed soil without compromising the growth requirements of the plant providing soil moisture content is kept at field capacity. We propose all future studies in this area should consider the error associated with scanning at different soil moisture contents.     

PP2A mediates lateral root development under NaCl-induced osmotic stress throughout auxin redistribution in Arabidopsis thaliana

Aim

Auxin plays an important role in modulating root system architecture. The effect of salinity on root development has been extensively studied; however, evidence on how salinity affects lateral root development and its underlying molecular mechanism is scarce. Here, we analyzed the role of protein phosphatase PP2A activity in auxin redistribution during Arabidopsis root system adaptation under NaCl-induced osmotic stress.

Method

Arabidopsis Col-0 and DR5::UidA seedlings were grown in MS media containing NaCl alone or in combination with the auxin transport inhibitor naphthylphthalamic acid, the synthetic auxin α-Naphthaleneacetic acid or the phosphatase inhibitor Okadaic acid. After 8 days, primary root length and lateral root number in seedlings were quantified and the auxin distribution was analyzed.

Results

Promotion of primary root shortening and lateral root development induced by osmotic stress correlated with an increase in active auxin content and a >50 % reduction in protein phosphatase type 2A (PP2A) activity. Moreover, the observed effects on seedlings under osmotic stress are more pronounced with the PP2A inhibitor Okadaic acid.

Conclusion

Our data suggest PP2A is a positive regulator of osmotic stress-induced root system architecture modulation, involving auxin redistribution in Arabidopsis thaliana.