Root length dynamics in agroforestry with Gliricidia sepium as compared to sole cropping in the semi-deciduous rainforest zone of West Africa

Tree root systems may improve soil fertility through carbon inputs, uptake of leachable nutrients and maintenance of soil biomass, but can at the same time reduce crop yields by competition for water and nutrients. Quantitative information about the positive and negative effects of tree roots and their changes in space and time are necessary for the optimization of agroforestry associations. An alley cropping experiment was layed out as a randomized complete block design on a Plinthic Lixisol/Ferralic Cambisol with Gliricidia sepium hedgerows at 5 m distance, including a sole cropping control. The development of root systems was monitored by sequential soil coring (eight samplings) during one year, with maize and groundnut as crops. Additional information is presented from a single sampling for rice during the foregoing year. Pronounced fluctuations of live root length density indicated an important variability in the nutrient and water uptake capacity of the vegetation. At low total root length density, the hedgerows affected the root development in the agroforestry plots directly by the presence of their root systems. At high root length density, they affected root development mainly by improving crop root growth and influencing the composition of the spontaneous vegetation. The root length density of the hedgerows was too low to compete with the crops for soil resources. The hedgerows tended to increase root length densities in the subsoil when few roots were present, thus possibly reducing the risk of nutrient leaching. However, the length density of the perennial root systems decreased during the cropping season, presumably as an effect of repeated pruning, and attained minimum values almost at the same time as the crops. Trees with denser root systems which are less frequently pruned may be more efficient in achieving closer nutrient cycles, though at the cost of higher root competition with crops.     

Fractal geometry of bean root systems: correlations between spatial and fractal dimension

An obstacle to the study of root architecture is the difficulty of measuring and quantifying the three-dimensional configuration of roots in soil. The objective of this work was to determine if fractal geometry might be useful in estimating the three-dimensional complexity of root architecture from more accessible measurements. A set of results called projection theorems predict that the fractal dimension (FD) of a projection of a root system should be identical to the FD of roots in three-dimensional space (three-dimensional FD). To test this prediction we employed SimRoot, an explicit geometric simulation model of root growth derived from empirical measurements of common bean (Phaseolus vulgaris L.). We computed the three-dimensional FD, FD of horizontal plane intercepts (planar FD), FD of vertical line intercepts (linear FD), and FD of orthogonal projections onto planes (projected FD). Three-dimensional FD was found to differ from corresponding projected FD, suggesting that the analysis of roots grown in a narrow space or excavated and flattened prior to analysis is problematic. A log-linear relationship was found between FD of roots and spatial dimension. This log-linear relationship suggests that the three-dimensional FD of root systems may be accurately estimated from excavations and tracing of root intersections on exposed planes.     

Study of differences between vertical root maps observed in a maize crop and simulated maps obtained using a model for the three-dimensional architecture of the root system

Differences between observed and simulated vertical root maps were studied in an attempt to evaluate the predictive ability of a simulation model of root system architecture under field conditions on mature plants, and to identify avenues for improvement. Some methodological problems associated with root mapping in the field are considered with a sensitivity analysis.Comparisons were made on a maize crop (early maturing hybrid F1 cultivar Dea) 15 days after silking. Four vertical root maps, perpendicular to the row and midway between two successive plants, were observed. Simulated root maps for different locations along the row showed essentially the same pattern, attesting of an approximately two-dimensional distribution of the roots in such a crop. Simulation of the intesection of roots with thin layers (thickness from 0 to 20 mm) instead of a perfect plane allowed us to assess effects due to the roughness of actual trench walls, and possible artefacts in the observation of root intersections. The simulated root profiles were very sensitive to this thickness, especially in the 0–5 mm range, in both average values, and overall shape. Actual data were close to the 3 mm thick simulations. This value seems plausible under our field conditions.Differences between simulated and actual root maps were shown to be mostly accounted for by the variations in soil bulk density. Thus, this environmental parameter appears as the most important one to include into the model for improving its predictions.     

Role of cytokinin and auxin in shaping root architecture: regulating vascular differentiation, lateral root initiation, root apical dominance and root gravitropism

BACKGROUND AND AIMS: Development and architecture of plant roots are regulated by phytohormones. Cytokinin (CK), synthesized in the root cap, promotes cytokinesis, vascular cambium sensitivity, vascular differentiation and root apical dominance. Auxin (indole-3-acetic acid, IAA), produced in young shoot organs, promotes root development and induces vascular differentiation. Both IAA and CK regulate root gravitropism. The aims of this study were to analyse the hormonal mechanisms that induce the root's primary vascular system, explain how differentiating-protoxylem vessels promote lateral root initiation, propose the concept of CK-dependent root apical dominance, and visualize the CK and IAA regulation of root gravitropiosm. KEY ISSUES: The hormonal analysis and proposed mechanisms yield new insights and extend previous concepts: how the radial pattern of the root protoxylem vs. protophloem strands is induced by alternating polar streams of high IAA vs. low IAA concentrations, respectively; how differentiating-protoxylem vessel elements stimulate lateral root initiation by auxin-ethylene-auxin signalling; and how root apical dominance is regulated by the root-cap-synthesized CK, which gives priority to the primary root in competition with its own lateral roots. CONCLUSIONS: CK and IAA are key hormones that regulate root development, its vascular differentiation and root gravitropism; these two hormones, together with ethylene, regulate lateral root initiation.     

The evolution of physiology and development in the cluster root: teaching an old dog new tricks?

In this paper we examine the key elements of cluster or proteoid roots, and trace their origins back to regular root properties. By viewing the root system as being composed of two categories of surface, the high transport capacity (HTC) area, just behind the meristem, and the low transport capacity (LTC) area (the rest of the root system), based on export and import capacities, we examine root system architecture in terms of structure–function relationships, and conclude that measuring total root exudation per unit area, volume or mass will not give useful comparative data for root transport properties. Furthermore, the cluster root represents a manipulation of the HTC to LTC root surface area ratio. Increased exudation and P uptake may be no higher in individual rootlets than in other HTC regions of the root system. We also examine the transformation theory (the theory of form resulting from a series of forces, which, when altered, lead to a change, or transformation in form) as an explanation of cluster root evolution, and conclude that the cluster root requires only a change in pericycle response to depleted internal nutrient levels, with the other characteristics representing consequences stemming from the form and constraints of the root system.     

分形理论及其在土壤空间变异研究中的应用

土壤具有不同程度的空间变异性,土壤空间变异研究对于土壤管理有重要意义.本文简要综述了分形理论及其在土壤空间变异研究中的应用,重点讨论了利用矩方法计算土壤属性分形维,多重分形分析土壤空间变异性及基于多重分形谱参数的土壤属性尺度转换.早期研究验证了分形理论在分析土壤空间变异中的有效性和应用潜力,国内外近期研究则报道了利用分形及多重分形理论分析土壤空间变异的最新进展.分形理论可以成为量化土壤属性空间变异性及尺度转换的重要工具.     

Evaluation in field conditions of a three-dimensional architectural model of the maize root system: Comparison of simulated and observed horizontal root maps

Most existing water and nutrient uptake models are based on the assumption that roots are evenly distributed in the soil volume. This assumption is not realistic for field conditions, and significantly alters water or nutrient uptake calculations. Therefore, development of models of root system growth that account for the spatial distribution of roots is necessary.The objective of this work was to test a three dimensional architectural model of the maize root system by comparing simulated horizontal root maps with observed root maps obtained from the field. The model was built using the current knowledge on maize root system morphogenesis and parameters obtained under field conditions. Simulated root maps (0.45 × 0.75 m) of horizontal cross sections at 3 depths and 3 dates were obtained by using the model for a plant population. Actual root maps were obtained in a deep, barrier-free clay-loamy soil by digging pits, preparing selected horizontal planes and recording root contacts on plastic sheets.Results showed that both the number of cross-sections of axile roots, and their spatial distribution characterized with the R-index value of Clark and Evans (1954), were correctly accounted for by the model at all dates and depths. The number of cross-sections of laterals was also correctly predicted. However, laterals were more clustered around axile roots on simulated root maps than on observed root maps. Although slight discrepancies appeared between simulated and observed root maps in this respect, it was concluded that the model correctly accounted for the general colonization pattern of the soil volume by roots under a maize crop.     

Patterns in root traits of woody species hosting arbuscular and ectomycorrhizas: implications for the evolution of belowground strategies

Root traits vary enormously among plant species but we have little understanding of how this variation affects their functioning. Of central interest is how root traits are related to plant resource acquisition strategies from soil. We examined root traits of 33 woody species from northeastern US forests that form two of the most common types of mutualisms with fungi, arbuscular mycorrhizas (AM) and ectomycorrhizas (EM). We examined root trait distribution with respect to plant phylogeny, quantifying the phylogenetic signal (K statistic) in fine root morphology and architecture, and used phylogenetically independent contrasts (PICs) to test whether taxa forming different mycorrhizal associations had different root traits. We found a pattern of species forming roots with thinner diameters as species diversified across time. Given moderate phylogenetic signals (K = 0.44–0.68), we used PICs to examine traits variation among taxa forming AM or EM, revealing that hosts of AM were associated with lower branching intensity (r_PIC = −0.77) and thicker root diameter (r_PIC = −0.41). Because EM evolved relatively more recently and intermittently across plant phylogenies, significant differences in root traits and colonization between plants forming AM and EM imply linkages between the evolution of these biotic interactions and root traits and suggest a history of selection pressures, with trade-offs for supporting different types of associations. Finally, across plant hosts of both EM and AM, species with thinner root diameters and longer specific root length (SRL) had less colonization (r_PIC = 0.85, −0.87), suggesting constraints on colonization linked to the evolution of root morphology.     

Morphometric analysis of root systems: application of the technique and influence of soil fertility on root system development in two herbaceous species

Abstract. A method for describing root systems based on geomorphological techniques developed for river systems is described. Root systems, in common with other natural branching structures (rivers, bronchioles, trees), appear to obey Morton's Law of Branching: there is a constant ratio, the bifurcation or branching ratio, R_b, between the number of branches of a given order, N_u , and that of the next order. N_u+1 , In experiments where Poa annua , and Rumex cripus , were grown at two levels of fertility, the first-order roots (the youngest members in this system) were generally unresponsive to fertility, and differences in the root systems were largely the result of changes in the second-order roots, those formed at the junction of two first-order roots. These differences were reflected in the branching ratio, R_b Although it is possible to explain these results by a stochastic model of branch development, the R_b values for roots are higher than for other natural branching structures, and higher than the random model predicts. It is possible that a model based on optimum exploration of space may be more appropriate and provide a key to the factors governing root branching patterns.     

Plant physiology in theory and practice: An analysis of the WBE model for vascular plants

The theoretical model of West, Brown and Enquist (hereafter WBE) proposed the fractal geometry of the transport system as the origin of the allometric scaling laws observed in nature. The WBE model has either been criticized for some restrictive and biologically unrealistic constraints or its reliability debated on the evidence of empirical tests. In this work, we revised the structure of the WBE model for vascular plants, highlighting some critical assumptions and simplifications and discuss them with regard to empirical evidence from plant anatomy and physiology. We conclude that the WBE model had the distinct merit of shedding light on some important features such as conduit tapering. Nonetheless, it is over-simplistic and a revised model would be desirable with an ontogenetic perspective that takes some important phenomena into account, such as the transformation of the inner sapwood into heartwood and the effect of hydraulic constraints in limiting the growth in height.     

Short- and long-term effects of a change in the spatial distribution of nitrate in the root zone on N uptake, growth and root development of young lettuce plants

Abstract. The effects of a change in the distribution of nitrate within the root zone on N uptake and growth were studied using young lettuce plants after reducing the proportion of their root systems supplied with nitrate from 100 to ca 10% in split-root experiments in the glasshouse. The main effects of the localized nitrate supply were concentrated in a 2-week period immediately after the treatment was imposed, when a temporary reduction in nitrate uptake caused the gradual development of N deficiency and a decline in plant growth rate. The plants adapted to the change in nitrate distribution, initially by increasing unit absorption rates (uptake rates per unit weight of root) and more gradually by increasing production of new roots in the high-nitrate zone. As a result, relative N uptake rates and relative growth rates were restored to the same levels as for control plants (given a spatially uniform N supply throughout) after ca 12d, even though only ca 12–15% of their roots were exposed to nitrate at this time. Thereafter, the plants continued to adapt by concentrating new root growth in the nitrate-containing zone, ultimately allowing unit absorption rates to return to normal. There was no evidence of any significant N deficiency in the plants after the initial adaptive response was complete, even though the total-N concentrations of the plants given the localized supply were consistently less than those given the uniform N treatment, and nitrate concentrations in the petiole sap were generally lower in leaves on one side of the plant (because of limited lateral movement of nitrate between xylem vessels during its transport to the shoot). The delay in the initiation of an adaptive response caused a significant check in growth, and the resulting relative weight differences were maintained throughout the subsequent life of the plant. Plants in all treatments matured on the same date, so yields for those grown with the localized supply were less than those of the control, and could not be recovered by delaying final harvest without unacceptable loss of quality. The pattern of the changes in N uptake and plant growth, and the effect on final yield, were similar to those exhibited by young lettuce plants subjected to a temporary interruption in nitrate supply, suggesting that the reduction in final yield for plants grown with the localized supply was largely the effect of the check in growth which occurred whilst the Plants were adapting to the change in nitrate distribution during the early part of the experiment. This implies that the rate of dry matter production of young lettuce plants can be altered by N treatment without affecting their rate of physiological development.     

RATP: a model for simulating the spatial distribution of radiation absorption, transpiration and photosynthesis within canopies: application to an isolated tree crown

The model RATP (radiation absorption, transpiration and photosynthesis) is presented. The model was designed to simulate the spatial distribution of radiation and leaf-gas exchanges within vegetation canopies as a function of canopy structure, canopy microclimate within the canopy and physical and physiological leaf properties. The model uses a three-dimensional (3D) representation of the canopy (i.e. an array of 3D cells, each characterized by a leaf area density). Radiation transfer is computed by a turbid medium analogy, transpiration by the leaf energy budget approach, and photosynthesis by the Farquhar model, each applied for sunlit and shaded leaves at the individual 3D cell-scale. The model typically operates at a 20–30 min time step. The RATP model was applied to an isolated, 20-year-old walnut tree grown in the field. The spatial distribution of wind speed, stomatal response to environmental variables, and light acclimation of leaf photosynthetic properties were taken into account. Model outputs were compared with data acquired in the field. The model was shown to simulate satisfactorily the intracrown distribution of radiation regime, transpiration and photosynthetic rates, at shoot or branch scales.     

Use of the root contact concept,an empirical leaf conductance model and pressure-volume curves in simulating crop water relations

A simulation model “DanStress” was developed for studying the integrated effects of soil, crop and climatic conditions on water relations and water use of field grown cereal crops. The root zone was separated into 0.1 m deep layers of topsoil and subsoil. For each layer the water potential at the root surface was calculated by a single root model, and the uptake of water across the root was calculated by a root contact model. Crop transpiration was calculated by Monteith's combination equation for vapour flow. Crop conductance to water vapour transfer for use in Monteith's combination equation was scaled up from an empirical stomatal conductance model used on sunlit and shaded crop surfaces of different crop layers. In the model, transpirational water loss originates from root water uptake and changes in crop water storage. Crop water capacitance, used for describing the water storage, was derived from the slope of pressure-volume (PV) curves of the leaves. PV curves were also used for deriving crop water potential, osmotic potential, and turgor pressure. The model could simulate detailed diurnal soil-crop water relations during a 23-day-drying cycle with time steps of one hour. During the grain filling period in spring barley (Hordeum distichum L.), grown in a sandy soil in the field, measured and predicted values of leaf water and osmotic potential, RWC, and leaf stomatal conductance were compared. Good agreement was obtained between measured and predicted values at different soil water deficits and climatic conditions. In the field, measured and predicted volumetric soil water contents (θ) of topsoil and subsoil layers were also compared during a drying cycle. Predicted and measured θ-values as a function of soil water deficits were similar suggesting that the root contact model approach was valid. From the investigation we concluded: (I) a model, which takes the degree of contact between root surface and soil water into account, can be used in sandy soil for calculation of root water uptake, so that the root conductance during soil water depletion only varies by the degree of contact; (II) crop conductance, used for calculation of crop transpiration, can be scaled up from an empirical single leaf stomatal conductance model controlled by the level of leaf water potential and micrometeorological conditions; (III) PV curves are usable for describing crop water status including crop water storage.     

A model for the prediction of Oncomelania hupensis in the lake and marshland regions, China

A model has been developed for predicting the density of Oncomelania hupensis, the intermediate host snail of Schistosoma japonicum. The model takes into account different environmental factors, including elevation, air and soil temperature, type of vegetation, mean height of preponderant vegetation and soil humidity. Deviance and Akaike information criteria were used to determine the best model fits. Model diagnostics and internal and external validations of model efficiency were also performed. From the final prediction model, two important results emerge. First, air temperature should be used with care to study the distribution of O. hupensis and to predict its potential survival because the impact is indirect, and it is weaker and more unstable than soil temperature. Second, the more important environmental factor for O. hupensis prediction at the microscale is soil humidity, but the more important macroscale environmental factor is soil temperature. This finding might help in selecting different environmental features for studying O. hupensis at different spatial scales. Our model is promising for predicting the density of O. hupensis, and hence can provide more objective information about snail dispersal, which might eventually replace the tedious and imprecise field work for annual surveillance of O. hupensis.     

Diversity analysis of the response to Zn within the Arabidopsis thaliana species revealed a low contribution of Zn translocation to Zn tolerance and a new role for Zn in lateral root development

This work reports the first characterization of the natural variation of Zn tolerance and accumulation in Arabidopsis thaliana. Root and shoot growth as well as Zn content were determined for 27 A. thaliana accessions grown in vitro in presence of Zn concentrations ranging from 1 to 250 μm. All traits varied by at least twofold and their broad sense heritability varied from 0.36 to 0.91. Primary and lateral root developments were differently affected by Zn in the different accessions. Remarkably, Zn was for the first time shown to be essential for the development of lateral roots. As a general rule, the different traits showed uncorrelated variations. In particular, variation in Zn tolerance was not linked to either root or shoot Zn contents. The only detectable relationship between different traits linked Zn sensitivity of roots to root-to-shoot Zn translocation but the correlation between variation of these traits was pretty low. This suggests that Zn translocation from root to shoots explains only a part of Zn tolerance. Our analysis opens the way to the characterization of genetic determinants controlling different Zn-related traits through the identification of particular accessions displaying contrasted phenotypes and representing excellent starting material to develop quantitative trait locus (QTL) studies.     

Understanding plant rooting patterns in semi‐arid systems: an integrated model analysis of climate,soil type and plant biomass

Aim A consistent set of root characteristics for herbaceous plants growing in water‐limited environments has been developed based on compilations of global root databases, but an overall analysis of why these characteristics occur is still missing. The central question in this study is whether an ecohydrological model which assumes that rooting strategies reflect maximization of transpiration can predict the variations in rooting strategies of plants in dry environments. Location Arid ecosystems across the globe. Methods A model was used to explore interactions between plant biomass, root–shoot allocation, root distribution, rainfall, soil type and water use by plants. Results Model analyses showed that the predicted shifts in rooting depth and root–shoot allocation due to changes in rainfall, soil type and plant biomass were quite similar to observed shifts. The model predicted that soil type, annual rainfall and plant biomass each had strong effects on the rooting strategies that optimize transpiration, but also that these factors have strong interactive effects. The process by which plants compete for water availability (soil evaporation or drainage) especially affected the depth distribution of roots in the soil, whereas the availability of rainfall mainly affected the optimal root–shoot allocation strategy. Main conclusions The empirically observed key patterns in rooting characteristics of herbaceous plant species in arid environments could be explained in this theoretical study by using the concept of hydrological optimality, represented here by the maximization of transpiration.     

Genetic and physiological analysis of doubled-haploid,aluminium-resistant lines of wheat provide evidence for the involvement of a 23 kD,root exudate polypeptide in mediating resistance

We have made use of a genetic approach to develop homozygous, near-isogenic germplasm for investigating aluminium (Al) resistance in Triticum aestivum L. A conventional backcross program was used to transfer Al resistance from the Al-resistant cultivar, Maringa, to a locally-adapted, Al-sensitive cultivar, Katepwa. At the third backcross stage, a single, resistant isoline (Alikat = Katepwa*3/Maringa) was chosen on the basis of superior root growth after 14 days of exposure to a broad range of Al concentrations (0 to 600 µM). Genetic analysis of doubled-haploid lines (DH) developed from this isoline suggested that resistance is controlled by a single dominant gene. Crosses between DH Alikat and DH Katepwa yielded an Al-resistant F1 population. Backcrossing this F1 population to DH Katepwa produced a population which segregated 1:1 for Al resistance, while selfing produced a population segregating 3 : 1 for Al resistance. Under conditions of Al stress, Al-resistant F2 plants released a suite of novel low molecular weight polypeptides into the rhizosphere. One of these polypeptides (23 kD) shows substantive Al-binding capacity and segregates with the resistant phenotype. While the precise mechanisms that mediate Al resistance are still unknown, this research has provided support for a possible role of the 23 kD exudate polypeptide in mediating resistance to Al. To more fully understand the role that this polypeptide plays in Al-resistance, we are attempting to clone this gene from microsequence data obtained from purified protein.     

中国亚热带不同菌根树种的根叶形态学性状特征与生长差异: 以江西新岗山为例

探究植物功能性状的种内和种间变异不仅有助于揭示植物对环境的适应, 也能够反映植物的生态策略, 但不同菌根类型树木生长过程中根叶形态学功能性状的适应策略仍有待探究。本研究依托中国亚热带森林生物多样性与生态系统功能实验研究平台(BEF-China)选取7种丛枝菌根(AM)树木和7种外生菌根(EM)树木的纯林, 测定各个树种的比叶面积、叶干物质含量、比根长、根系直径、树高生长速率、地径生长速率及细根生物量等根叶形态学功能性状和生长指标, 探讨了两种菌根类型树种间的根叶形态学特征的差异。结果表明: 与AM树种相比, EM树种具有较小的比叶面积、吸收根平均直径和生长速率, 但具有更大的叶干物质含量; 两种菌根树种之间的比根长和细根生物量无显著差异。比叶面积、叶干物质含量、树高生长速率、地径生长速率和细根生物量等功能性状及生长指标在不同菌根类型、树种及二者的交互作用中均存在显著差异; 且树种、根功能型、菌根类型及三者之间的交互作用均对根功能性状有显著影响。EM树种地上指标的种内变异均大于种间变异, 而AM树种地上指标的种内和种间变异程度类似; 但两种菌根树种细根生物量的种间变异均大于种内变异。尽管两种菌根树种地上部分生长速率较快通常表现为较低的叶干物质含量, 但AM树种通常拥有较高的吸收根比根长, 而EM树种拥有较粗的运输根平均直径。吸收根比根长越低, 两类菌根树种的细根生物量就越多。由此可见, 根叶功能性状对植物地上部分的生长具有一定的协同效应, 其中运输根主要在EM树种地上生长过程中发挥重要作用, 吸收根主要与AM树种的地上部分生长有关; 但两类菌根树种的地下细根生物量均与吸收根有关。