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.     

Fine root morphological and functional traits in Fagus sylvatica and Fraxinus excelsior saplings as dependent on species, root order and competition

Aims

The mechanisms of belowground competition are not well understood. Addressing literature reports on competition-induced changes in tree fine root morphology, we conducted a growth experiment with tree saplings to investigate competition effects on important root morphological and functional traits in a root order-focused analysis.

Methods

European beech and European ash saplings were grown for 34 months in containers under greenhouse conditions in monoculture (2 conspecific plants), in mixture (1 beech and 1 ash) or as single plants. The root system was fractionated according to root orders and eight morphological and functional properties were determined.

Results

Root order was the most influential factor affecting the fine root traits (except for root diameter and δ¹³C); a significant species identity effect was found for root diameter, tissue density, N concentration and δ¹³C. Ash fine roots were thicker, but had lower tissue densities, contained more N and had systematically higher δ¹³C values than beech roots. The competition treatments had no significant effect on morphological root traits but altered δ¹³C in the 2nd root order.

Conclusion

Neither intra- nor interspecific root competition affected fine root morphology significantly suggesting that competition-induced root modification may not be a universal phenomenon in temperate trees.     

Application of X-ray computed tomography to quantify fresh root decomposition in situ

Background and aims

Much of our understanding of plant root decomposition and related carbon cycling come from mass loss rates calculated from roots buried in litter bags. However, this may not reflect what actually happens in the soil, where the interactions between root and soil structure presents a more complex physico-chemical environment compared to organic matter isolated in a porous bag buried in disturbed soil. This work investigates the potential of using X-ray micro-computed tomography (CT) to measure root decomposition in situ.

Methods

Roots of Vicia faba L. were excised from freshly germinated seeds, buried in re-packed soil cores and cores incubated for 60 days. Changes in root volume and surface area were measured using repeated scans. Additional samples were destructively harvested and roots weighed to correlate root mass with root volume. The method was further applied to an experiment to investigate the effects of soil bulk density and soil moisture on root decomposition.

Results

Root volume (X-ray CT) and root mass (destructive harvest) decreased by 90 % over the 60 day incubation period, by which stage, root volume and mass had stabilised. There was a strong correlation (R ²?=?0.97) between root volume and root mass.

Conclusions

X-ray CT visualization and analysis provides a unique toolbox to understand root decomposition in situ.     

Spatial root distribution of plants growing in vertical media for use in living walls

Background and Aims

For plants growing in living walls, the growth potential is correlated to the roots ability to utilize resources in all parts of the growing medium and thereby to the spatial root distribution. The aim of the study was to test how spatial root distribution was affected by growing medium, planting position and competition from other plants.

Methods

Five species (Campanula poscharskyana cv. ‘Stella’, Fragaria vesca cv. ‘Småland’, Geranium sanguineum cv. ‘Max Frei’, Sesleria heufleriana and Veronica officinalis cv. ‘Allgrün’) were grown in three growing media (coir and two of rockwool) in transparent boxes under greenhouse conditions. Root frequency was registered and the activity of individual root systems was studied via ¹⁵N uptake and plant dry weight was measured.

Results

Plants in coir had stronger root growth in all parts of the medium than plants in rockwool. Upwards root growth was limited for plants in the middle or lower parts of the medium and ¹⁵N measurements confirmed that only plants in the bottom of the box had active roots in the bottom of the medium. The species differed in root architecture and spatial root distribution.

Conclusions

The choice of growing medium, plant species and planting position is important for a living wall as it affects the spatial root growth of the plants.     

Wheat root diversity and root functional characterization

Background and Aims

Under limited moisture conditions, roots can play an outstanding role with respect to yield stability by effective absorption of water from soil. A targeted integration of root traits into plant breeding programs requires knowledge on the existing root diversity and access to easy and cost-effective methods. This study aimed to assess wheat root diversity, root properties in relation to water regime, and the efficiency of root capacitance for in situ screening.

Methods

Root morphological, anatomical properties and root capacitance of wheat species from different ploidy levels were studied under field conditions in 2 years contrasting in water regime. Soil water content was weekly measured.

Results

Significant genotypic differences were observed for most root traits. The investigated genotypes exploited different strategies to maximize soil water depletion, e.g. high topsoil root length density, low tissue mass density, high specific root length, deep rooting and looser xylem vessels. Multivariate statistics of root traits revealed an acceptable genotypic differentiation according to regional origin, genetics and capacity to extract soil water.

Conclusions

Under supply-driven environments, dehydration avoidance via water uptake maximization can be achieved through high topsoil rooting density. In this regard, root capacitance can be useful for in situ screening.     

Root dynamics of Carex stricta-dominated tussock meadows

Background and aims

The roots of tussock-forming plants contribute to the formation of microtopographic features in many ecosystems, but the dynamics of such roots are poorly understood. We examined the spatial heterogeneity of tussock fine root dynamics to investigate allocation patterns and the role of root productivity in the persistence of tussock structures.

Methods

We compared the spatial variability of fine root (<1 mm, 1–2 mm) density, biomass, % live, allocation, turnover rate (using bomb 14C), and productivity of four Carex stricta Lam.-dominated tussock meadows in the upper Midwest, USA (3 reference, 1 restored site).

Results

Relative to underlying microsites, tussocks were warm, dry, and high in root density, productivity, % live biomass, and turnover. Root productivity averaged 649 g?m^?2 yr^?1 (±208) in reference sites, comprised 57 % (±10) of total net production, and was concentrated in tussocks (70 %?±?4). Root turnover rate averaged 0.63 yr^?1 (±0.08), but tussocks had ~50 % faster root turnover than the underlying soil, and <1 mm roots turned over ~40 % faster than 1–2 mm roots.

Conclusions

Our detailed analysis of the spatial heterogeneity of tussock root dynamics suggests that high allocation and elevated turnover of tussock roots facilitates organic matter accumulation and tussock persistence over time.     

Estimating the parameters of a 3-D root distribution function from root observations with the trench profile method: case study with simulated and field-observed root data

Background and aims

Root length density (RLD) is a parameter that is difficult to measure, but crucial to estimate water and nutrient uptake by plants. In this study a novel approach is presented to characterize the 3-D root length distribution by supplementing data of the 3-D distribution of root intersections with data of root length density from a limited number of soil cores.

Methods

The method was evaluated in a virtual experiment using the RootTyp model and a field experiment with cauliflower (Brassica oleracea L. botrytis) and leek (Allium porrum, L.).

Results

The virtual experiment shows that total root length and root length distribution can be accurately estimated using the novel approach. Implementation of the method in a field experiment was successful for characterizing the growth of the root distribution with time both for cauliflower and leek. In contrast with the virtual experiment, total root length could not be estimated based upon root intersection measurements in the field.

Conclusions

The novel method of combining root intersection data with root length density data from core samples is a powerful tool to supply root water uptake models with root system information.     

Longleaf and loblolly pine seedlings respond differently to soil compaction,water content,and fertilization

Aims

Longleaf pine (Pinus palustris Mill.) is being restored across the U.S. South for a multitude of ecological and economic reasons, but our understanding of longleaf pine’s response to soil physical conditions is poor. On the contrary, our understanding of loblolly pine (Pinus taeda L.) root and shoot growth response to soil conditions is well established.

Methods

We performed a comparative greenhouse study which modeled root length density, total seedling biomass, and the ratio of aboveground:belowground mass as functions of volumetric water content, bulk density and soil fertility (fertilized or not).

Results

Root length density was about 35 % greater in longleaf pine seedlings compared to loblolly pine seedlings, and was reasonably well modeled (R ²?=?0.54) for longleaf pine by bulk density (linear), volumetric water content (quadratic), soil fertility, and the interactions of bulk density, volumetric water content, species, and soil fertility. The aboveground:belowground mass ratio (ABR) increased at both extremes of water content.

Conclusions

This research indicates that young longleaf pine seedling root systems respond more negatively to extremes of soil physical conditions than loblolly pine, and compacted or dry loamy soils should be ameliorated in addition to normal competition control, especially on soils degraded by past management.     

Relation of fine root distribution to soil C in a Cunninghamia lanceolata plantation in subtropical China

Background and aims

Growth and distribution of fine roots closely depend on soil resource availability and affect soil C distribution in return. Understanding of relationships between fine root distribution and soil C can help to predict the contribution of fine root turnover to soil C accumulation.

Methods

A study was conducted in a subtropical Cunninghamia lanceolata plantation to assess the fine root mass density (FRMD), fine root C density (FRCD) of different fine root groups as well as their relations with soil C.

Results

The FRMD and FRCD of short-lived roots, dead roots and herb roots peaked in the 0–10 cm soil layer and decreased with soil depth, while FRMD, FRCD of long-lived roots peaked in the 10–20 cm soil layer. Soil C was positively related to FRMD and FRCD of total fine roots (across all three soil layers), dead roots (0–10 cm) and herb roots (10–20 cm) as well as FRCD of short-lived roots (20–40 cm) (P <0.05).

Conclusions

Soil C was mainly affected by herb roots in upper soil layers and by woody plant roots in deeper soil layers.     

N- and P-mediated seminal root elongation response in rice seedlings

Aims

In rice, seminal root elongation plays an important role in acquisition of nutrients such as N and P, but the extent to which different N forms and P concentrations affect root growth is poorly understood. This study aimed to examine N- and P-mediated seminal root elongation response and to identify putative QTLs associated with seminal root elongation.

Methods

Seminal root elongation was evaluated in 15 diverse wild and cultivated accessions of rice, along with 48 chromosome segment substitution lines (CSSLs) derived from a cross between the rice variety ‘Curinga’ and Oryza rufipogon (IRGC 105491). Root elongation in response to different forms of N (NH₄ ⁺, NO₃ ^? and NH₄NO₃) and concentrations of P was evaluated under hydroponic conditions, and associated putative QTL regions were identified.

Results

The CSSL parents had contrasting root responses to N and P. Root elongation in O. rufipogon was insensitive to N source and concentration, whereas Curinga was responsive. In contrast to N, seminal root elongation and P concentration was positively correlated. Three putative QTLs for seminal root elongation in response to N were detected on chromosome 1, and one QTL on chromosome 3 was associated with low P concentration.

Conclusions

Genetic variation in seminal root elongation and plasticity of nutrient response may be appropriate targets for marker-assisted selection to improve rice nutrient acquisition efficiency.     

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.     

Root elongation rate is correlated with the length of the bare root apex of maize and lupin roots despite contrasting responses of root growth to compact and dry soils

Background

We investigated interacting effects of matric potential and soil strength on root elongation of maize and lupin, and relations between root elongation rates and the length of bare (hairless) root apex.

Methods

Root elongation rates and the length of bare root apex were determined for maize and lupin seedlings in sandy loam soil of various matric potentials (?0.01 to ?1.6 MPa) and bulk densities (0.9 to 1.5 Mg m^?3).

Results

Root elongation rates slowed with both decreasing matric potential and increasing penetrometer resistance. Root elongation of maize slowed to 10 % of the unimpeded rate when penetrometer resistance increased to 2 MPa, whereas lupin elongated at about 40 % of the unimpeded rate. Maize root elongation rate was more sensitive to changes in matric potential in loosely packed soil (penetrometer resistances <1 MPa) than lupin. Despite these differing responses, root elongation rate of both species was linearly correlated with length of the bare root apex (r² 0.69 to 0.97).

Conclusion

Maize root elongation was more sensitive to changes in matric potential and mechanical impedance than lupin. Robust linear relationships between elongation rate and length of bare apex suggest good potential for estimating root elongation rates for excavated roots.     

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.     

Spatial variation in root system activity of tomato (Solanum lycopersicum L.) in response to short and long-term waterlogging as determined by 15N uptake

Background and aim

Root system activity is affected by abiotic stresses, which often creates spatial differences in root conditions. This is expected to influence plants ability to cope with suboptimal conditions.

Methods

Changes in root system activity were determined as ¹⁵N root uptake in top and bottom layers of potted tomato plants (Solanum lycopersicum L.), after waterlogging the bottom layer for 24 h or 5 d. The plants were grown in peat-based media; non-compacted or highly-compacted, resulting in differences in gas diffusion, air permeability and oxygen availability.

Results

The roots were affected by short-term waterlogging (24 h) by decreasing uptake in the bottom layer and increasing uptake in the pot top layer. Long-term waterlogging (5 d) decreased the ¹⁵N root uptake more in both layers. Root uptake recovered fast (within 6 h) after short-term waterlogging, whereas recovery of long-term waterlogged roots took more than 24 h, suggesting production of new root biomass. Despite affecting physical properties, medium compaction did not affect root uptake. Aboveground biomass was affected by waterlogging by increasing the dry matter percentage, decreasing nitrogen (N) percentage and increasing starch content.

Conclusions

The results confirmed that root uptake in different layers of small pots could be distinguished by the ¹⁵N technique, which was applicable under potentially denitrifying conditions. The results demonstrated that during short-term stress in part of the root system plants increased uptake from the non-affected parts of the root system, probably as compensation for suboptimal conditions.     

The measurement of woody root decomposition using two methodologies in a Sitka spruce forest ecosystem

Background and aims

The decomposition of roots is an important process in the loss of carbon (C) and the mineralization of nitrogen (N) in forest ecosystems. The early stage decomposition rate of Sitka spruce (Picea sitchensis (Bong.) Carr.) roots was determined using trenched plots and decomposition bags.

Methods

Stumps of known age were trenched and quadrants (50?cm by 50?cm) excavated from randomly selected stumps every 6?months over 4?years, while the mass loss from buried roots in decomposition bags, divided among four diameter categories (ranging from fine roots <2?mm to large roots >50?mm), was monitored for 27?months. The C and N concentrations of excavated samples at different time points were analysed.

Results

The change in total root necromass per quadrant showed a higher decomposition rate-constant (k) of 0.24?±?0.068?year^?1 than the k-value of roots in decomposition bags (0.07?±?0.005?year^?1). The C concentration (47.24?±?0.609?%) did not significantly change with decomposition. There was a significant increase in the C:N ratio of roots in all diameter categories (fine: 48.92?%, small: 38.53?%, medium 11.71?%, large: 76.25?%) after 4?years of decomposition, driven by N loss. Root diameter accounted for 78?% of the variation in the N concentration of roots as decomposition progressed.

Conclusion

Though the trenched plot approach offered an alternative to the more common decomposition bag method for estimating root decomposition, high spatial variation and sampling difficulties may lead to an overestimation of the mass loss from trenched roots, thus, the decomposition bag method gives a more reliable decomposition rate-constant.     

Effects of biochar amendment on root traits and contaminant availability of maize plants in a copper and arsenic impacted soil

Background and aims

Biochar has been proposed as a tool to enhance phytostabilisation of contaminated soils but little data are available to illustrate the direct effect on roots in contaminated soils. This work aimed to investigate specific root traits and to assess the effect of biochar amendment on contaminant availability.

Methods

Amendment with two different types of biochar, pine woodchip and olive tree pruning, was assessed in a rhizobox experiment with maize planted in a soil contaminated with significant levels of copper and arsenic.

Results

Amendment was found to significantly improve root traits compared to the control soil, particularly root mass density and root length density. Copper uptake to plants and ammonium sulphate extractable copper was significantly less in the biochar amended soils. Arsenic uptake and extractability varied with type of biochar used but was not considered to be the limiting factor affecting root and shoot development.

Conclusions

Root establishment in contaminated soils can be enhanced by biochar amendment but choice of biochar is key to maximising soil improvement and controlling contaminant availability.     

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.     

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.     

Fine root and litterfall dynamics of three Korean pine (Pinus koraiensis) forests along an altitudinal gradient

Background and aims

Fine root and aboveground litterfall, two large fluxes of nutrients and carbon in the forest ecosystems, are key processes to be considered in efforts of measuring, modeling and predicting soil carbon sequestration.

Methods

We used sequential coring and litter trap to measure seasonal dynamics of fine root and litterfall in three Korean pine dominated forests along an altitudinal gradient in the Changbai Mountain during the 2012 growing season.

Results

Fine root biomass decreased significantly while necromass increased remarkably with altitude. Patterns and amounts of fine root production and mortality varied among forest types. Litterfall decreased significantly with altitude, whereas forest floor mass increased. Carbon inputs through fine root mortality and litterfall decreased significantly with altitude while carbon storage of fine root mass did not differ among forest types and carbon storage of forest floor mass was significantly larger in higher altitudinal forests due to lower turnover rates.

Conclusions

This study provided an insight into the variations of fine root and litterfall dynamics among three Korean pine forests which were associated with different vegetation traits and environmental conditions, and also the quantification of carbon fluxes through fine root mortality and litterfall for estimating carbon budget of temperate forest.     

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.