Root-induced changes in radial oxygen loss, rhizosphere oxygen profile, and nitrification of two rice cultivars in Chinese red soil regions

Aims

To evaluate the external and internal morphological differences of roots that might influence rice root radial oxygen loss (ROL) and the corresponding rhizosphere nitrification activity, growth characteristics and nitrogen nutrition of rice.

Methods

The root ROL and rhizosphere oxygen profile were determined using a miniaturised Clark-type oxygen microelectrode system, and the rhizosphere nitrification activity was studied with a short-term nitrification activity assay.

Results

The rice biomass, nitrogen accumulation and nitrogen use efficiency (NUE) of ZH (high yield) were significantly higher than those of HS (low yield). The root biomass, number, diameter and porosity of ZH were also much greater than those of HS. The inner and surface oxygen concentrations of the root of ZH were significantly higher than those of HS. The order of paddy soil oxygen penetration depth was ZH?>?HS?>?CK, and the order of the oxygen concentrations detected in the water layer and rhizosphere soil was the same. The rhizosphere nitrification activity and nitrate concentration of ZH were significantly higher than those of HS.

Conclusions

More porous and thicker roots improved the individual root ROL, and more adventitious root numbers enhanced the entire plant ROL and correspondingly improved the rhizosphere nitrification activity, which might influence the growth and nitrogen nutrition of rice.     

Growth of tree roots in hostile soil: A comparison of root growth pressures of tree seedlings with peas

Background and Aims

As part of a study on growth of tree roots in hostile soil, we envisaged that establishment and survival of trees on hard, dry soil may depend on their ability to exert axial root growth pressures of similar magnitude to those of the roots of agricultural plants (with significant root thickening when roots grow across an air gap or cracks and biopores). We selected tree species originating from a range of different soil and climatic conditions to evaluate whether their relative success on harsh soil (in an evolutionary sense) might be related to the magnitude of root growth pressures they could exert, or how they performed in the very early stages of growth after germination.

Methods

We measured the maximum axial root growth force (F_max) on single lateral root axes of 3- to 4- month old seedlings of 6 small-seeded eucalypts from 2 different habitats and 2 contrasting soil types. Root growth rate, root diameter and F_max were also measured on the primary root axes of a large-seeded acacia and a domesticated annual (Pisum sativum) seedling for up to 10 days following germination.

Results

The lateral roots of the 6 eucalypts and the primary roots of the acacia were considerably smaller than the primary roots of P. sativum and they exerted average forces of similar magnitude to one another (0.198 to 0.312 N). The maximum axial root growth pressures were all in the range 150 to 250 kPa but E. leucoxylon, E. loxophleba and A. salicina exerted the greatest pressures among the trees, and comparable pressures to those exerted by the primary roots of 2-day-old P. sativum (211-252 kPa). Although the primary roots of acacia seedlings exerted increasing axial root growth pressures over a 10-day period following germination, the pressures were still only slightly greater than those of the domesticated plant, P. sativum.

Conclusions

The lack of any very large differences in axial root growth pressures between trees and domesticated plants suggests that trees that grow well in harsh soil don’t do so by exerting higher root growth pressures alone but by also exploring the network of cracks and pores more effectively than do other plants that are less successful.     

How does the root system inhibit windthrow in thinned black spruce sites in the boreal forest?

Key message

The root shape and the angle between roots play an important role to prevent windthrow occurrence.

Abstract

Partial cutting is frequently applied to increase the volume growth of residual stems. However, the opening of the forest increases the wind speed within the site, and consequently, the risk of windthrow. In the case of black spruce, uprooted trees are normally characterized by a lifting of the root plate. This research was conducted to compare the root systems of standing and uprooted black spruces, after commercial thinning, by looking at root architecture, volume and radial growth. For this purpose, data from a pool of 18 standing and 18 uprooted trees from three study areas were analyzed. The distribution of roots around the stump was compared between both types of trees, standing and uprooted. The radial growth was measured at 30 cm in the stem, 10 cm and 60 cm in the roots. The shape (I and T-beam) and volume were recorded for each root system. The structure of the roots was also mapped to obtain a spatial overview of the angle between roots. The root shape (at 10 and 60 cm) and the angle between roots combined with the diameter of the stem at stump height seem to determine the vulnerability of black spruce to windthrow. Uprooted trees developed fewer roots, with a large sector around the stump without lateral roots which suggests its major implication in the resistance to windthrow.     

Mechanical interactions between neighbouring roots during pullout tests

Background and Aims

The quantification of root reinforcement function is important for landscape managers and engineers. The estimation of root mechanical reinforcement is often based on models that do not consider the potential interaction between neighbouring roots. Root-soil mechanical interactions related to the root spacing and bundle geometry remain unclear including potential effects on the reliability of the current models. The objective of this study is to quantify the mechanical interactions among neighbouring roots or roots networks using modelling approaches and pullout laboratory experiments.

Methods

Based on simple geometrical characterization of individual root geometry, we calculated dissipation patterns of frictional root-soil interfacial stresses in radial and longitudinal directions. Considering simple superposition of shear stresses within the soil matrix, we quantified characteristic root densities at which the radial mechanical interactions influence global pullout behaviour of the root bundle both for branched and unbranched roots. Laboratory pullout tests on root bundles were carried out at root spacings of 15, 35 and 105 mm. In addition, we tested effects of non-parallel (crossing) root bundle geometry.

Results

We found no significant statistical differences in root pullout force for the different root spacing in parallel alignment of roots. Branches increase pullout force by 1.5 times. Moreover, the mean displacement at the pullout peak-force was 7.2 % of length for unbranched roots and about 4.1 % of length for branched roots. The model shows its potential comparing it with empirical results concerning the holes leaved by roots, according with the branch pattern.

Conclusion

The study quantifies the influence of root spacing and arrangement geometry within a root bundle on its mechanical behaviour. The assumption of “non-interacting” neighbouring roots in root reinforcement methods is no longer valid for root spacing less than 15 mm and root reinforcement methods. Moreover crossing roots shown a statistical difference. This information is important for improved understanding root reinforcement mechanisms in steep hill slope and the interplay between anchoring /failure and root bundle pullout vs root breakage.     

Quantifying the effect of soil compaction on three varieties of wheat (Triticum aestivum L.) using X-ray Micro Computed Tomography (CT)

Aims

X-ray Micro Computed Tomography (CT) enables interactions between roots and soil to be visualised without disturbance. This study examined responses of root growth in three Triticum aestivum L. (wheat) cultivars to different levels of soil compaction (1.1 and 1.5?g?cm^?3).

Methods

Seedlings were scanned 2, 5 and 12?days after germination (DAG) and the images were analysed using novel root tracking software, RootViz3D?, to provide accurate visualisation of root architecture. RootViz3D? proved more successful in segmenting roots from the greyscale images than semi-automated segmentation, especially for finer roots, by combining measurements of pixel greyscale values with a probability approach to identify roots.

Results

Root density was greater in soil compacted at 1.5?g?cm^?3 than at 1.1?g?cm^?3 (P?=?0.04). This effect may have resulted from improved contact between roots and surrounding soil. Root diameter was greater in soil at a high bulk density (P?=?0.006) but overall root length was reduced (P?=?0.20). Soil porosity increased with time (P?<?0.001) in the uncompacted treatment.

Conclusions

RootViz3D? root tracking software in X-ray CT studies provided accurate, non-destructive and automated three dimensional quantification of root systems that has many applications for improving understanding on root-soil interactions.     

Effect of species, root branching order and season on the root traits of 13 perennial grass species

Aims

Inter-specific comparisons of plant traits may vary depending on intra-specific variation. Here we examine the impact of root branching order and season on key functional root traits for grass species. We also compare root traits among co-existing grass species as a step towards defining root trait syndromes.

Methods

Monocultures of 13 grass species, grown under field conditions and subjected to intensive management, were used to record root trait values for coarse roots (1st order, >0.3?mm), fine roots (2nd and 3rd orders, <0.2?mm) and mixed root samples over three growing seasons.

Results

Branching order and species had a significant effect on root trait values, whereas season showed a marginal effect. The diameter of coarse roots was more variable than that of fine roots and, as expected, coarse roots had higher tissue density and lower specific root length values than fine roots. Principal component analysis run on eight root traits provided evidence for two trait syndromes related to resource acquisition and conservation strategies across grass species.

Conclusions

Our data show that root branching order is the main determinant of root trait variation among species. This highlights the necessity to include the proportion of fine vs coarse roots when measuring traits of mixed root samples.     

Effects of copper on growth, radial oxygen loss and root permeability of seedlings of the mangroves Bruguiera gymnorrhiza and Rhizophora stylosa

Purpose

Mangrove wetlands have experienced significant contaminant input such as copper (Cu), aggravated by rapid urban development. This study aimed to investigate the possible function of root permeability in metal detoxification.

Methods

Pot trials were conducted to evaluate the responses of root permeability in relation to metal (Cu) exposure in seedlings of two mangroves: Bruguiera gymnorrhiza and Rhizophora stylosa.

Results

Copper inhibited plant growth and root permeability of the two species significantly (due to decreases in root porosity, thickening of exodermis and increases in lignification), leading to a significant reduction in radial oxygen loss (ROL). A negative correlation between soil Cu and ROL from root tip was also observed. The observed metal uptake by excised roots further indicated that increased lignification would directly prevent excessive Cu from further entering into the roots.

Conclusions

In summary, the two mangroves reacted to Cu by producing an impermeable barrier in roots. Such an inducible barrier to ROL is likely to be an adaptive strategy against Cu toxicity. This study reveals new evidence of a structural adaptive strategy for metal tolerance by mangrove plants.     

Soil penetration resistance, a suitable soil property to account for variations in root elongation and branching

Background and Aims

Root system development is affected by soil conditions. The effects of bulk density, water content and penetration resistance on root development processes were investigated in peach trees.

Methods

Peach tree rootstocks were grown in various soil conditions, combining two bulk densities (1.2 and 1.5?g soil.cm^-3) and three water contents (0.14, 0.17 and 0.20?g.g^-1soil). Root parameters (tip diameter, length of apical unbranched zone, branching density and diameters of main and lateral roots) and plant growth (leaves, branches, trunk, root dry mass) were measured. Root growth processes (elongation, branching) were studied using relationships between root parameters.

Results

The proportion of biomass allocated to each plant compartment was similar whatever the soil conditions. Variations in root development were best explained by the variation in penetration resistance, rather than other soil properties. Increased soil penetration resistance reduced the root elongation rate, especially for thick roots. In addition, the branching pattern was affected. In soil with a high penetration resistance, the root system shape differs from a typical herringbone pattern.

Conclusions

These results allow quantification of the root system plasticity, and improve our understanding of the interactions between root development and soil properties.     

A randomization method for efficiently and accurately processing fine roots, and separating them from debris, in the laboratory

Background and Aims

We developed a method for processing roots from soil cores and monoliths in the laboratory to reduce the time and cost devoted to separating roots from debris and improve the accuracy of root variable estimates. The method was tested on soil cores from a California oak savanna, with roots from trees, Quercus douglasii, and annual grasses.

Methods

In the randomized sampling method, one isolates the sample fraction consisting of roots and organic debris?<?= 1 cm in length, and randomizes it through immersion in water and vigorous mixing. Sub-samples from the mixture are then used to estimate the percentage of roots in this fraction, thereby enabling an estimate of total sample biomass.

Results

We found that root biomass estimates, determined through the randomization method, differed from total root biomass established by meticulously picking every root from a sample with an error of 3.0 % +/? 0.6 %?s.e.

Conclusions

This method greatly reduces the time and resources required for root processing from soil cores and monoliths, and improves the accuracy of root variable estimates compared to standard methods. This gives researchers the ability to increase sample frequency and reduce the error associated with studying roots at the landscape and plant scales.     

A displacement-based model for estimating the shear resistance of root-permeated soils

Aims

This paper presents a displacement-based model for predicting the relationship between the increase in shear resistance and shear displacement for soils permeated with an entire plant root system.

Methods

The root force in the root system is estimated based on the shear deformation developed in the soil. This displacement-based model takes a number of factors into account, including the distribution of the shear deformation in the soil, the root orientation, the mobilized root forces, and the root properties.

Results

The proposed model reasonably captures the relationship between the increase in the shear resistance (ΔS) and the shear displacement, as shown by a comparison of the predicted results with data from in situ shear tests.

Conclusions

Major findings are the following: (1) the ΔS value increases considerably with increasing b coefficients, which are used to describe the deformed shape of the shear zone, and Young’s moduli of roots at the early stage of shearing; (2) the ΔS value increases significantly with the τ value at large shear deformations; (3) short roots play an important role in the contribution of root systems to the shear resistance of the soil. However, the success of the model relies on the appropriate estimate of the deformation characteristics on the shear zone and the soil-root bond strength.     

The influence of stem guying on radial growth,stem form and internal resin features in radiata pine

Key message

Stem guying to prevent wind-induced swaying of radiata pine trees resulted in significant changes in radial growth, but did not affect the frequency of compression wood or resin features.

Abstract

Mechanical stress resulting from wind forces acting on trees can cause a number of direct and indirect effects ranging from microscopic changes in cambial activity through to stem breakage and uprooting. To better understand these effects on radial stem growth and wood properties, an experiment was established in a 13-year-old radiata pine (Pinus radiata D Don) stand in which 20 trees were guyed to prevent them from swaying. Radial growth was monitored in these trees and 20 matched controls at monthly intervals for 5 years. The trees were then felled and radial growth, resin features and compression wood were assessed on cross-sectional discs taken at fixed locations up the stem. There was a significant reduction in radial growth at breast height (1.4 m above the ground) in the guyed trees, but an increase in growth immediately above the guying point. A total of 277 resin features were observed in the growth rings formed following guying. The overall frequency of such features was related to height within the stem and annual ring number. No effect of stem guying was found on the incidence of compression wood. Interestingly, the distribution of resin features also did not differ between guyed and un-guyed trees. There was no evidence of a link between stem restraint as a result of guying and the incidence of resin features, suggesting that other factors, such as soil moisture may be more influential.     

Aluminum resistance mechanisms in oat (Avena sativa L.)

Background and aims

Enhanced aluminum (Al) resistance has been observed in dicots over-expressing enzymes involved in organic acid synthesis; however, this approach for improving Al resistance has not been investigated in monocots. Among the cereals, oat (Avena sativa L.) is considered to be Al resistant, but the basis of resistance is not known.

Methods

A hydroponic assay and hematoxylin staining for Al accumulation in roots were used to evaluate Al resistance in 15 oat cultivars. Malate and citrate release from roots was measured over a 24?h period. A malate dehydrogenase gene, neMDH, from alfalfa (Medicago sativa L.) was used to transform oat.

Results

Oat seedlings were highly resistant to Al, as a concentration of 325?μM AlK(SO₄)₂ was needed to cause a 50% decrease in root growth. Most oat cultivars tested are naturally resistant to high concentrations of Al and effectively excluded Al from roots. Al-dependent release of malate and Al-independent release of citrate was observed. Al resistance was enhanced in a transgenic oat line with the highest accumulation of neMDH protein. However, overall root growth of this line was reduced and expression of neMDH in transgenic oat did not enhance malate secretion.

Conclusions

Release of malate from oat roots was associated with Al resistance, which suggests that malate plays a role in Al resistance of oat. Over-expression of alfalfa neMDH enhanced Al resistance in some lines but was not effective alone for crop improvement.     

Physiological and biochemical responses of the iron chlorosis tolerant grapevine rootstock 140 Ruggeri to iron deficiency and bicarbonate

Background and aims

Iron (Fe) deficiency chlorosis associated with high levels of soil bicarbonate is one of the main nutritional disorders observed in sensitive grapevine genotypes. The aim of the experiment was to assess both the independent and combined effects of Fe and bicarbonate nutrition in grapevine.

Methods

Plants of the Fe chlorosis tolerant 140 Ruggeri rootstock were grown with and without Fe(III)-EDTA and bicarbonate in the nutrient solution. SPAD index, plant growth, root enzyme (PEPC, MDH, CS, NADP⁺ ?IDH) activities, kinetic properties of root PEPC, organic acid concentrations in roots and xylem sap and xylem sap pH were determined. A factorial statistical design with two factors (Fe and BIC) and two levels of each factor was adopted: +Fe and ?Fe, and +BIC and ?BIC.

Results

This rootstock strongly reacted to Fe deficiency by activating several response mechanisms at different physiological levels. The presence of bicarbonate in the nutrient solution changed the activity of PEPC and TCA related enzymes (CS, NADP⁺-IDH) and the accumulation/translocation of organic acids in roots of Fe-deprived plants. Moreover, this genotype increased root biomass and root malic acid concentration in response to high bicarbonate levels in the substrate. Bicarbonate also enhanced leaf chlorophyll content.

Conclusions

Along with a clear independent effect on Fe nutrition, our data support a modulating role of bicarbonate on Fe deficiency response mechanisms at root level.     

Arbuscular mycorrhizal fungi shift competitive relationships among crop and weed species

Aims

Because plants cannot change their environmental circumstances by changing their location, they must instead adapt to a wide variety of environmental conditions, especially soil conditions. One of the most effective ways for a plant to adapt to a given soil condition is by modifying its root system architecture. We aim to identify the genetic factors controlling root growth angle, a trait that affects root system architecture.

Methods

The present study consisted of a genetic analysis of the seminal root growth angle in wheat; the parental varieties of the doubled haploid lines (DHLs) used in this study exhibited significantly different root growth directions. Using the ‘basket’ method, the ratio of deep roots (DRR; the proportion of total roots with GA > 45 degrees) was observed for evaluating deep rooting.

Results

We were able to identify novel quantitative trait loci (QTLs) controlling the gravitropic and hydrotropic responses of wheat roots. Moreover, we detected one QTL for seminal root number per seedling (RN) on chromosome 5A and two QTLs for seminal root elongation rate (ER) on chromosomes 5D and 7D.

Conclusions

Gravitropic and hydrotropic responses of wheat roots, which play a significant role in establishing root system architecture, are controlled by independent genetic factors.     

Capacity of biological soil crusts colonized by the lichen <Emphasis Type="Italic">Diploschistes</Emphasis> to metabolize simple phenols

Background and aims

Soil compaction strongly affects water uptake by roots. The aim of the work was to examine soil—plant interactions with focus on the impact of distribution of compacted soil layers on growth and water uptake by wheat roots.

Methods

The growth-chamber experiment was conducted on wheat growth in soil with compacted soil layers. The system for maintaining constant soil water potential and measurement of daily water uptake from variously compacted soil layers was used.

Results

Layered soil compaction differentiated vertical root distribution to higher extent for root length than root mass. The propagation rate of a water extraction front was the highest through layers of moderately compacted soil. The root water uptake rate was on average 67 % higher from moderately than heavily compacted soil layers. Correlations between water uptake and the length of thick roots were increasing with increasing level of soil compaction.

Conclusions

The study shows that root amount, water uptake, propagation of water extraction and shoot growth strongly depend on the existence of compacted layers within soil profile. The negative effects of heavily compacted subsoil layer on water uptake were partly compensated by increased uptake from looser top soil layers and significant contribution of thicker roots in water uptake.     

Neutron imaging reveals internal plant water dynamics

Background and aims

Knowledge of plant water fluxes is critical for assessing mechanistic processes linked to biogeochemical cycles, yet resolving root water transport dynamics has been a particularly daunting task. Our objectives were to demonstrate the ability to non-invasively monitor individual root functionality and water fluxes within Zea mays L. (maize) and Panicum virgatum L. (switchgrass) seedlings using neutron imaging.

Methods

Seedlings were propagated for 1–3 weeks in aluminum chambers containing sand. Pulses of water or deuterium oxide were then tracked through the root systems by collecting consecutive radiographs during exposure to a cold-neutron source. Water flux was manipulated by cycling on a growth lamp to alter foliar demand for water.

Results

Neutron radiography readily illuminated root structure, root growth, and relative plant and soil water content. After irrigation there was rapid root water uptake from the newly wetted soil, followed by hydraulic redistribution of water through the root system to roots terminating in dry soil. Water flux within individual roots responded differentially to foliar illumination based on supply and demand of water within the root system.

Conclusions

Sub-millimeter scale image resolution revealed timing and magnitudes of root water uptake, redistribution within the roots, and root-shoot hydraulic linkages—relationships not well characterized by other techniques.     

N2 fixation of pea hypernodulating mutants is more tolerant to root pruning than that of wild type

Background and aims

As a legume, pea plant has the ability to symbiotically fix N₂. However, symbiotic N₂ fixation is very sensitive to environmental stresses that affect plant growth, and there is little knowledge on the impact of root pruning on N₂ fixation and plant growth.

Methods

In this study, we removed half of the nodulated roots of pea wild-type Frisson and hypernodulating mutants P64, P118, and P121. Dinitrogen fixation was measured using ¹⁵N labeling and carbon assimilation and partitioning between plant organs using ¹³C labeling.

Results

Root pruning decreased N₂ fixation by ?46 to ?79 % in wild-type and mutants. Pea mutant P118 had a lower decrease of specific activity of N₂ fixation (?17 %) than both wild-type and other mutants (?36 to ?62 %). For all genotypes, root pruning increased root and nodule sinks strengths for carbon. For P118 and for P121, this was associated to higher nodule growth than for control plants, as measured 8 days after root pruning.

Conclusion

This is the first analysis of N₂-fixing plant response to root pruning. Importantly, we showed that some hypernodulating mutant pea lines (P118 and to a lesser extent P121) withstood this stress better than wild-type did.     

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.     

Field application of a DNA-based assay to the measurement of roots of perennial grasses

Background and aims

DNA-based methods present new opportunities for overcoming the difficulties of accurately identifying and quantifying roots of different plant species in field soils. In order to quantify species-specific root biomass from measurements of DNA, consideration needs to be given to replication and ability to recover roots for calibration purposes in order to account for spatial, temporal and inter- and intra-species variation in DNA content of roots and distribution of roots within the soil profile.

Methods

This paper develops the field application of a DNA-based technique for direct quantification of roots in soils. The method was applied to a field experiment to investigate differences in root growth of acid-soil resistant and sensitive genotypes of perennial pasture grasses in an acid soil. DNA was extracted directly from soil and species-specific DNA was quantified using quantitative real-time PCR prior to estimation of root biomass.

Results

Root growth of the perennial grasses was quantified using the DNA-based technique, although separate calibration procedures were needed to convert DNA content to root mass for each species, soil layer and sampling date. Compared to acid-soil resistant genotypes, lesser root growth in acid soil layers and reduced above-ground dry matter production was observed for acid-soil sensitive genotypes.

Conclusions

The DNA-based method allowed genotypic differences in root growth to be assessed directly in soil and was advantageous for rapid processing of a large number of samples. However, high replication was still required to overcome spatial variability and separate calibrations were required for different species and soil depths across sampling times. The technique demonstrated greater root growth of acid-soil resistant perennial grasses which was beneficial for their establishment and persistence.