Understanding the impact of root morphology on overturning mechanisms: a modelling approach

BACKGROUND AND AIMS: The Finite Element Method (FEM) has been used in recent years to simulate overturning processes in trees. This study aimed at using FEM to determine the role of individual roots in tree anchorage with regard to different rooting patterns, and to estimate stress distribution in the soil and roots during overturning. METHODS: The FEM was used to carry out 2-D simulations of tree uprooting in saturated soft clay and loamy sand-like soil. The anchorage model consisted of a root system embedded in a soil block. Two root patterns were used and individual roots removed to determine their contribution to anchorage. KEY RESULTS: In clay-like soil the size of the root-soil plate formed during overturning was defined by the longest roots. Consequently, all other roots localized within this plate had no influence on anchorage strength. In sand-like soil, removing individual root elements altered anchorage resistance. This result was due to a modification of the shape and size of the root-soil plate, as well as the location of the rotation axis. The tap root and deeper roots had more influence on overturning resistance in sand-like soil compared with clay-like soil. Mechanical stresses were higher in the most superficial roots and also in leeward roots in sand-like soil. The relative difference in stresses between the upper and lower sides of lateral roots was sensitive to root insertion angle. Assuming that root eccentricity is a response to mechanical stresses, these results explain why eccentricity differs depending on root architecture. CONCLUSIONS: A simple 2-D Finite Element model was developed to better understand the mechanisms involved during tree overturning. It has been shown how root system morphology and soil mechanical properties can modify the shape of the root plate slip surface as well as the position of the rotation axis, which are major components of tree anchorage.     

Variation of wood strength in tree roots

In order to determine if the distribution of lateral root strengthis related to the shape of the system and the forces withstood,wood samples were taken from roots of various mature tree speciesand the strength tested. Root strength decreased along the root at different rates, dependingon the type of root system present. Slightly tapered lateralroots in plate root systems were relatively stronger furtheraway from the stem than the highly tapered laterals in heartand tap root systems. Wood strength in Norway spruce (platesystem) was found to increase along the lateral roots beforedecreasing again. The increase in strength may coincide withthe point of maximum bending of the root as the tree sways backwardsand forwards in the wind. Strength was also found to increaseon the underside of lateral roots in the plate systems of poplar.The undersides of these roots will experience high compressivestresses due to the weight of the tree pushing the root on tothe hard bearing surface of the soil. External loading forces in plate root systems will be transmittedinto the soil further away from the stem due to the lack ofbranches, therefore a high strength along the root will helpresist mechanical stress. The high rate of branching near thestem, or large, rigid, main tap root, found in heart and taproot systems, respectively, allows a faster dissipation of forcesnearer the stem, therefore a high investment in strength furtheralong the root is not necessary. Key words: Anchorage, stability, wind, stress, forces     

Effect of soil acidity, soil strength and macropores on root growth and morphology of perennial grass species differing in acid-soil resistance

It is unclear whether roots of acid-soil resistant plants have significant advantages, compared with acid-soil sensitive genotypes, when growing in high-strength, acid soils or in acid soils where macropores may allow the effects of soil acidity and strength to be avoided. The responses of root growth and morphology to soil acidity, soil strength and macropores by seedlings of five perennial grass genotypes differing in acid-soil resistance were determined, and the interaction of soil acidity and strength for growth and morphology of roots was investigated. Soil acidity and strength altered root length and architecture, root hair development, and deformed the root tip, especially in acid-soil sensitive genotypes. Root length was restricted to some extent by soil acidity in all genotypes, but the adverse impact of soil acidity on root growth by acid-soil resistant genotypes was greater at high levels of soil strength. Roots reacted to soil acidity when growing in macropores, but elongation through high-strength soil was improved. Soil strength can confound the effect of acidity on root growth, with the sensitivity of acid-resistant genotypes being greater in high-strength soils. This highlights the need to select for genotypes that resist both acidity and high soil strength.     

Douglas-fir root biomass and rooting profile in relation to soils in a mid-elevation area (Beaujolais Mounts,France)

Douglas-fir is the main reforestation species in the French Massif Central area (14 000 ha), but little is known about its rooting strategy in different soil conditions. This information has important implications for the choice of better soils for settling Douglas-fir, and consequently limiting risks of failure, pests or diseases. As a result, the influence of edaphic conditions on rooting patterns of dominant Douglas-fir was studied over a large range of ecological conditions in a mid-elevation area of the French Massif Central (Beaujolais Mounts). Root systems were studied extensively using the trench profile wall technique and the sector method in 74 pure and evenly aged Douglas-fir stands. The stands were chosen as being representative of soil conditions among 165 stands in an auto-ecological study. The rooting patterns were related to seven typical soil profiles, and to root profile groups. Results stressed that edaphic constraints due to substratum and soil structures have a strong influence on root system morphology. Important variations in root biomass and vertical distribution were highlighted among soils. Small fine root biomass is maximal for soils with no major edaphic constraints. The vertical distribution of fine root biomass is positively correlated for some soil types with organic C, total N, and most cations. For some types it was negatively correlated with the amount of exchangeable aluminum and coarse fragments, and with constraining rock facies. Harsher soils however, showed no correlation between soil chemical variables and fine-root biomass. A practical implication is that Douglas-fir seems to be a pliable and adaptive species: variation in habit and root system biomass are considerable within a study area which was presumed uniform.     

Biomechanics of nodal, seminal and lateral roots of barley: effects of diameter, waterlogging and mechanical impedance

Background and aims

Biomechanical properties of cereal root systems largely control both resistance to root lodging and their ability to stabilise soil. Abiotic stresses can greatly modify root system growth and form. In this paper the effect of waterlogging and moderate mechanical impedance on root biomechanics is studied for both lateral roots and the main axes of barley.

Methods

Barley (Hordeum vulgare) plants were subjected to transient water-logging and moderate mechanical impedance in repacked soil columns. Roots were excavated, separated into types (nodal, seminal or lateral) and tested in tension to measure strength and elastic modulus.

Results

Water-logging and mechanical impedance substantially changed root system growth whilst root biomechanical properties were affected by waterlogging. Root strength was generally greater in thin roots and depended on root type. For example, seminal roots 0.4–0.6 mm in diameter were approximately seven times stronger and five times stiffer than lateral roots of the same diameter when mechanically impeded. Root sample populations typically exhibited negative power-law relationships between root strength and diameter for all root types. Mechanical impedance slowed seminal root elongation by approximately 50 % and resulted in a 15 % and 11 % increase in the diameter of in nodal and seminal roots respectively. Power-law relationships between root diameter and root biomechanical properties corresponded to the different root types. Coefficients for between root diameter, strength and elastic modulus improved when separated by root type, with R² values increasing in some roots from 0.05 to 0.71 for root strength and 0.08 to 0.74 for elastic modulus.

Conclusions

Moderate mechanical impedance did not influence the tensile strength of roots, but, waterlogging diminished the relationship between root strength and diameter. Separation of root type improved predictions of root strength and elastic modulus using power-law regressions.     

Effect of different soil types,pH and phosphorus levels on the wheat seedling and the incidence ofPythium graminicolum

Summary In the present study, the effect of different types of soil, pH and phosphorus levels on root, shoot development and browning extent on shoot of wheat seedlings growth in soil inoculated withPythium graminicolum; was studied. Out of the four soil types the development of root and shoot was better in Domatta soil where as the disease development was more in black clay soil. The pH values below 5.5 and above 8.0 are favourable for the root and shoot development whereas these values are unfavourable for the development of the Pythium and consequently for infection by it. The percentage of browning of the collar region was more in soils near neutral (pH 6.5–7.1) and it was less in acidic (pH 5.0) or alkaline (pH 8.6) soils. The development of wheat roots and shoot in soil with 30 lb/acre phosphorus was good as compared to that in soil with P₀, P_15,45 and P₆₀ lb/acre in inoculated soil. The percentage of extent of browning on collar region was less at 15 lb and 30 lb/acre P, due to less development of the disease.     

重金属对土壤中萝卜种子发芽与根伸长抑制的生态毒性

高等植物是生态系统中的基本组成部分。一个平衡、稳定的生态系统生产健康、优良的高等植物。反之 ,一个不稳定或受到外来污染的生态系统 ,对高等植物的生长可带来不利和可见的负面影响。因此 ,利用高等植物的生长状况监测土壤污染程度 ,是从生态学角度衡量土壤健康状况 ,评价土壤质量的重要方法之一[4 ,6] 1) 。土壤生态毒理学评价方法是对化学分析方法的重要补充。目前已建立的高等植物毒理试验有三种方法 ,即 1根伸长试验 ;2种子发芽试验 ;3早期植物幼苗生长试验[3 ,5,6,10 ] 。最初 ,这类试验主要用于纯化学品的毒性检验 ,但随着对土壤…     

Volume Contents

Distribution of root systems through soils and recolonization of root channels by successive crops are fundamental, though difficult to study, processes of soil ecology. This article reports a minirhizotron (MR) study of corn and alfalfa root systems throughout the soil profile of Kalamazoo loam (fine-loamy, mixed, mesic Typic Hapludalf) monolith lysimeters for a three-year succession of corn, alfalfa and corn. Multiple-date comparisons within and between years were conducted to estimate total root densities in each soil horizon. Root recolonization was assessed by comparing every video frame of paired minirhizotrons, from recordings conducted one growing season apart. Distributions of corn root systems were modified by tillage practices. In 1994, root populations of corn in the Bt1 horizon peaked 75–90 days after planting (DAP). Numbers of corn roots per m2 in the Bt1 horizon were consistently higher for no-tillage (NT) than for conventional tillage (CT) lysimeters, in 1994 and 1996. Distribution of alfalfa roots within the soil profile was not significantly modified by tillage. However, alfalfa root decomposition rates responded to conventional and no-tillage practices and were specific for each soil horizon. Corn root systems growing in soils previously cropped with alfalfa presented similar patterns of root distribution by horizons as that of the previous alfalfa crop. Successive corn root systems did not display similar distribution patterns throughout the soil profile from one growing season to the next. Proportions of roots of the current crop recolonizing root induced macropores (RIMs) of the previous crop averaged 18% for corn after corn, 22% for alfalfa after corn and 41% for corn after alfalfa, across Bt horizons and tillage treatments. In conclusion, distribution of corn root systems appeared to be modified by tillage practices and root recolonization of RIMs was controlled by the preceding crop.     

The role of plant species and soil condition in the structural development of the rhizosphere

Roots naturally exert axial and radial pressures during growth, which alter the structural arrangement of soil at the root–soil interface. However, empirical models suggest soil densification, which can have negative impacts on water and nutrient uptake, occurs at the immediate root surface with decreasing distance from the root. Here, we spatially map structural gradients in the soil surrounding roots using non‐invasive imaging, to ascertain the role of root growth in early stage formation of soil structure. X‐ray computed tomography provided a means not only to visualize a root system in situ and in 3‐D but also to assess the precise root‐induced alterations to soil structure close to, and at selected distances away from the root–soil interface. We spatially quantified the changes in soil structure generated by three common but contrasting plant species (pea, tomato, and wheat) under different soil texture and compaction treatments. Across the three plant types, significant increases in porosity at the immediate root surface were found in both clay loam and loamy sand soils and not soil densification, the currently assumed norm. Densification of the soil was recorded, at some distance away from the root, dependent on soil texture and plant type. There was a significant soil texture × bulk density × plant species interaction for the root convex hull, a measure of the extent to which root systems explore the soil, which suggested pea and wheat grew better in the clay soil when at a high bulk density, compared with tomato, which preferred lower bulk density soils. These results, only revealed by high resolution non‐destructive imagery, show that although the root penetration mechanisms can lead to soil densification (which could have a negative impact on growth), the immediate root–soil interface is actually a zone of high porosity, which is very important for several key rhizosphere processes occurring at this scale including water and nutrient uptake and gaseous diffusion.     

A generic 3D finite element model of tree anchorage integrating soil mechanics and real root system architecture

Understanding the mechanism of tree anchorage in a forest is a priority because of the increase in wind storms in recent years and their projected recurrence as a consequence of global warming. To characterize anchorage mechanisms during tree uprooting, we developed a generic finite element model where real three-dimensional (3D) root system architectures were represented in a 3D soil. The model was used to simulate tree overturning during wind loading, and results compared with real data from two poplar species (Populus trichocarpa and P. deltoides). These trees were winched sideways until failure, and uprooting force and root architecture measured. The uprooting force was higher for P. deltoides than P. trichocarpa, probably due to its higher root volume and thicker lateral roots. Results from the model showed that soil type influences failure modes. In frictional soils, e.g., sandy soils, plastic failure of the soil occurred mainly on the windward side of the tree. In cohesive soils, e.g., clay soils, a more symmetrical slip surface was formed. Root systems were more resistant to uprooting in cohesive soil than in frictional soil. Applications of this generic model include virtual uprooting experiments, where each component of anchorage can be tested individually.     

黄土丘陵区不同植被根系数量特征及离散程度

根系的空间分布与植物自身的生物学特性、土壤资源、土壤结构等因素有密切的关系。对黄土丘陵区3种植被类型根系的数量特征及离散程度进行了研究,结果表明:山杨、灌丛和油松林地的根系数量随着根系径级的增加而减少;山杨林地根系最丰富,油松林次之,灌丛地根系数量最少。3种植被根系离散程度随深度的增加有明显的差异,其中灌丛和山杨林根系的离散指数在15cm深处达到最大值,随后又降低,而油松林的变化没有明显规律。无论是表层或是深层,3种植被根系离散指数均小于1,说明植被根系在土壤中呈聚集分布。在土壤相对均质的黄土地区,根系的空间聚集分布可能是根系的固有特性。     

Effect of soil type and plant species on the fluorescent pseudomonads nitrate dissimilating community

The distribution of nitrogen dissimilative abilities among 618 isolates of fluorescent pseudomonads was studied. These strains were isolated from two uncultivated soils (C and D; collected at Chateaurenard and Dijon, France, respectively) and from rhizosphere, rhizoplane and root tissue of two plant species (flax and tomato) cultivated on these two soils. According to their ability to dissimilate nitrogen, the isolates have been distributed into three metabolic types: non-dissimilators, NO₂ ^- accumulators and denitrifiers. While the three metabolic types were recovered in all the compartments of soil D experiments, only two (non-dissimilators and denitrifiers) were recovered in all the compartments of soil C experiments. Even under the contrasting conditions of the two soil types, both plants were able to select the nitrate dissimilating community among the total community of fluorescent Pseudomonas, but the mode of this selection seems to be dependent on both plant and soil type. The soil type appears to be unable to significantly modulate the strong selective effect of tomato. Indeed, similar dissimilator to non-dissimilator ratios were found in the root tissue of this plant species cultivated in both soils. In contrast, the different dissimilator to non-dissimilator ratios observed in flax roots between soils C and D suggest that the selective effect of flax was modulated by the soil type. Taxonomic identifications showed that the 618 isolates were distributed among three species (P. chlororaphis, P. fluorescens, P. putida) plus an intermediate type between P. fluorescens and P. putida. However, no clear relationship between the distribution of the metabolic types (functional diversity) and the distribution of bacterial species has been found. This revised version was published online in June 2006 with corrections to the Cover Date.     

Diurnal and seasonal variation in root xylem embolism in neotropical savanna woody species: impact on stomatal control of plant water status

Vulnerability to water-stress-induced embolism and variation in the degree of native embolism were measured in lateral roots of four co-occurring neotropical savanna tree species. Root embolism varied diurnally and seasonally. Late in the dry season, loss of root xylem conductivity reached 80% in the afternoon when root water potential (psi root) was about -2.6 MPa, and recovered to 25-40% loss of conductivity in the morning when psi root was about -1.0 MPa. Daily variation in psi root decreased, and root xylem vulnerability and capacitance increased with rooting depth. However, all species experienced seasonal minimum psi root close to complete hydraulic failure independent of their rooting depth or resistance to embolism. Predawn psi root was lower than psi soil when psi soil was relatively high (> -0.7 MPa) but became less negative than psi soil, later in the dry season, consistent with a transition from a disequilibrium between plant and soil psi induced by nocturnal transpiration to one induced by hydraulic redistribution of water from deeper soil layers. Shallow longitudinal root incisions external to the xylem prevented reversal of embolism overnight, suggesting that root mechanical integrity was necessary for recovery, consistent with the hypothesis that if embolism is a function of tension, refilling may be a function of internal pressure imbalances. All species shared a common relationship in which maximum daily stomatal conductance declined linearly with increasing afternoon loss of root conductivity over the course of the dry season. Daily embolism and refilling in roots is a common occurrence and thus may be an inherent component of a hydraulic signaling mechanism enabling stomata to maintain the integrity of the hydraulic pipeline in long-lived structures such as stems.     

Penetrometer resistance,root penetration resistance and root elongation rate in two sandy loam soils

Root penetration resistance and elongation of maize seedling roots were measured directly in undisturbed cores of two sandy loam soils. Root elongation rate was negatively correlated with root penetration resistance, and was reduced to about 50 to 60% of that of unimpeded controls by a resistance of between 0.26 and 0.47 MPa. Resistance to a 30° semiangle, 1 mm diameter penetrometer was between about 4.5 and 7.5 times greater than the measured root penetration resistance. However, resistance to a 5° semiangle, 1 mm diameter probe was approximately the same as the resistnace to root penetration after subtracting the frictional component of resistance. The diameter of roots grown in the undisturbed cores was greater than that of roots grown in loose soil, probably as a direct result of the larger mechanical impedance in the cores.     

Mechanics of root growth

Summary A model is developed for the rate of elongation of a root tip in terms of the balance of pressures acting on the root. Differentials of this equation give expressions for the changes in root elongation rate with respect to soil water potential and soil mechanical resistance. The model predicts that root cells osmoregulate against both water stress and soil mechanical resistance with predicts that root cells osmoregulate against both water stress and soil mechanical resistance with similar efficiencies which are less than 100%. Analysis of published data leads to the conclusion that root tips of pea osmoregulate with 70% efficiency. A working equation is developed for the elongation rate of roots in conditions of combined water stress and mechanical resistance.     

Vertical distribution of maize roots in relation to permanent soil characteristics

The vertical distribution of maize roots was studied in four contrasting soils, (arenosols, luvisols, planosols and vertisols) by using in-situ root mapping on vertical planes. The relationship between root contact frequency and depth was different for each soil, with a relatively low field-to-field variability within each soil type. The general aspect of this relationship did not change appreciably for three years in arenosols, with a low colonization in sandy layers probably being due to mechanical barriers. The relationship was consistently non-monotonic in luvisols and planosols, because of the sparse colonization of sandy layers. In planosols, these layers were traversed by some primary roots, which were essentially clustered in animal burrows. The distribution of root contact frequency was closer to an exponential function in vertisols. In these soils rooting depth and colonization of deep soil layers showed a marked increase during two dry years compared with a wet year. This was probably due to a denser net of shrinkage cracks and slickensides, where roots were essentially located in dry years. These results raise the possibility of modelling the decrease in root distribution with depth using soil information and climatic characteristics.     

Varietal Response and Estimates of Heritability of Resistance to Meloidogyne javanica in Carrots

With methods developed in this study, varietal responses to M. javanica were evaluated and heritability of resistance of two promising carrot cultivars was estimated. More egg masses were found on root systems inoculated with eggs added to the soil in three holes in 250 cm³ cups than by mixing the inoculum with soil in the cups. A resistant breeding line, CNPH 1437, was discriminated from susceptible cultivar Nova Kuroda with inoculum levels higher than 2,000 eggs per cup. Greenhouse and field results suggested that cultivars Nantes Superior and Shin Kuroda were susceptible, Kuronan was somewhat tolerant, and Brasilia and Tropical were resistant to M. javanica. Nantes Superior or Shin Kuroda yielded less in carbofuran-treated soil (3 kg a.i./ha) than Kuronan, Brasilia, and Tropical did in nontreated soil. However, incorporation of the nematicide greatly increased yields of Kuronan (32%), Brasilia (62%), and Tropical (91%). Primary root galling at the seedling stage was an adequate parameter for resistance evaluation. Estimated heritability were 0.48 ± 0.07 for primary root galling and 0.35 ± 0.08 for egg mass production in Brasilia, and 0.16 ± 0.11 for primary root galling and 0.31 ± 0.09 for egg mass production in Kuronan.     

Hydraulic properties of sphagnum peat moss and tuff (scoria) and their potential effects on water availability

The importance of macrostructure to root growth of ryegrass (L. perenne) seedlings sown on the soil surface was studied in two soils in which the macrostructure had resulted mainly from root growth and macro-faunal activity. Sets of paired soil cores were used, one of each pair undisturbed and the other ground and repacked to the field bulk density. Undisturbed and repacked soils were first compared at equal water potentials in the range −1.9 to −300 kPa. At equal water potential, the undisturbed soil always had the greater strength (penetration resistance), and root growth was always greater in the repacked soil with no macrostructure than it was in the soil with macrostructure intact. At equal high strength (low water potentials) it appeared that root growth was better when soils were structured. When strength was low (high water potentials), root growth was better in the unstructured soil. Soils were then compared during drying cycles over 21 days. The average rate at which roots grew to a depth of 60 mm, and also the final percentage of plants with a root reaching 60 mm depth, was greatest in repacked soils without macrostructure. The species of vegetation growing in the soil before the experiment affected root growth in undisturbed soil; growth was slower where annual grasses and white clover had grown compared with soil which had supported a perennial grass. It appears that relatively few roots locate and grow in the macrostructure. Other roots grow in the matrix, if it is soft enough to be deformed by roots. Roots in the matrix of a structured soil grow more slowly than roots in structureless soil of equal bulk density and water potential. The development of macrostructure in an otherwise structureless soil, of the type studied, is of no advantage to most roots. However, once a macrostructure has developed, the few roots locating suitable macropores are able to grow at low water potential when soil strength is high. The importance of macrostructure to establishing seedlings in the field lies in rapid penetration of at least a few roots to a depth that escapes surface drying during seasonal drought. ei]{gnB E}{fnClothier}     

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.     

Land use and climatic factors structure regional patterns in soil microbial communities

Aim  Although patterns are emerging for macroorganisms, we have limited understanding of the factors determining soil microbial community composition and productivity at large spatial extents. The overall objective of this study was to discern the drivers of microbial community composition at the extent of biogeographical provinces and regions. We hypothesized that factors associated with land use and climate would drive soil microbial community composition and biomass.
Location  Great Basin Province, Desert Province and California Floristic Province, California, USA.
Methods  Using phospholipid fatty acid analysis, we compared microbial communities across eight land-use types sampled throughout the State of California, USA ( n = 1117).
Results  The main factor driving composition and microbial biomass was land-use type, especially as related to water availability and disturbance. Dry soils were more enriched in Gram-negative bacteria and fungi, and wetter soils were more enriched in Gram-positive, anaerobic and sulphate-reducing bacteria. Microbial biomass was lowest in ecosystems with the wettest and driest soils. Disturbed soils had less fungal and more Gram-positive bacterial biomass than wildland soils. However, some factors known to influence microbial communities, such as soil pH and specific plant taxa, were not important here.
Main conclusions  Distinct microbial communities were associated with land-use types and disturbance at the regional extent. Overall, soil water availability was an important determinant of soil microbial community composition. However, because of the inclusion of managed and irrigated agricultural ecosystems, the effect of precipitation was not significant. Effects of environmental and management factors, such as flooding, tillage and irrigation, suggest that agricultural management can have larger effects on soil microbial communities than elevation and precipitation gradients.