Modelling applicability of fractal analysis to efficiency of soil exploration by roots

BACKGROUND AND AIMS: Fractal analysis allows calculation of fractal dimension, fractal abundance and lacunarity. Fractal analysis of plant roots has revealed correlations of fractal dimension with age, topology or genotypic variation, while fractal abundance has been associated with root length. Lacunarity is associated with heterogeneity of distribution, and has yet to be utilized in analysis of roots. In this study, fractal analysis was applied to the study of root architecture and acquisition of diffusion-limited nutrients. The hypothesis that soil depletion and root competition are more closely correlated with a combination of fractal parameters than by any one alone was tested. MODEL: The geometric simulation model SimRoot was used to dynamically model roots of various architectures growing for up to 16 d in three soil types with contrasting nutrient mobility. Fractal parameters were calculated for whole roots, projections of roots and vertical slices of roots taken at 0, 2.5 and 5 cm from the root origin. Nutrient depletion volumes, competition volumes, and relative competition were regressed against fractal parameters and root length. KEY RESULTS: Root length was correlated with depletion volume, competition volume and relative competition at all times. In analysis of three-dimensional, projected roots and 0 cm slices, log(fractal abundance) was highly correlated with log(depletion volume) when times were pooled. Other than this, multiple regression yielded better correlations than regression with single fractal parameters. Correlations decreased with age of roots and distance of vertical slices from the root origin. Field data were also examined to see if fractal dimension, fractal abundance and lacunarity can be used to distinguish common bean genotypes in field situations. There were significant differences in fractal dimension and fractal abundance, but not in lacunarity. CONCLUSIONS: These results suggest that applying fractal analysis to research of soil exploration by root systems should include fractal abundance, and possibly lacunarity, along with fractal dimension.     

Adaptive Changes and Genotypic Variation for Root Architecture of Common Bean in Response to Phosphorus Deficiency

Root architectural responses to phosphorus (p) availability may be an important trait for P acquisition efficiency. In the present study, The authors examined the effects of P availability on root architectural responses of different common bean genotypes. Five common bean (Phaseolus vulgaris L.) genotypes representing different origins and ecotypic races were compared both in a specially designed paper pouch system and a stratified P buffer sand culture system with computer image analysis. The results showed that root architecture was regulated by P availability. P deficiency led to form a shallower root system, as indicated by increased relative distribution of basal root length in the upper layers and decreased the growth angle of basal roots. There was significant genetic variation in root architecture in response to P deficiency both in the paper pouch system and the stratified sand culture system. Under low P conditions some genotypes were more gravitropically sensitive to low P availability, resulting in producing a shallower root system and enhanced root exploration into the surface soil, where soil available P is more concentrated. G19833 and DOR364, which were most contrasting in P efficiency, were also very different in root architectural response to P availability. The results from this study suggest that P availability regulates root architecture and P deficiency leads to shallower root architecture in beans. The genetic potential of root architecture provides the possibility of selecting this trait for improving P acquisition efficiency in common bean.     

Genotypic Variation in Root Morphological Characteristics of Common Bean in Relation to Phosphorus Efficiency (in English)

Root morphology in plants may be related to phosphorus (P) efficiency by affecting the absorption characteristics of the root system. However, genotypic variation in root morphological characteristics of common bean (Phaseolus vulgaris L.) as affected by P availability has not been well clarified. In the present study, systematic studies were conducted in a P-buffered sand culture system using three pairs of common bean parental materials with contrasting root traits in response to P deficiency. The results indicate that P availability significantly affects bean root morphology. Common bean tends to have smaller root system, shorter and coarser roots at low P availability. Genotypic variation in root morphology was observed among different genotypes in response to P availability. The P efficient genotypes appear to have larger, finer and longer root systems than the P inefficient genotypes, and such a variation was particularly obvious in the basal roots. From allomeric analysis, we found that morphological characteristics of the basal roots contribute more to P efficiency than those of the tap roots. Further studies with the F9 recombinant inbred lines derived from one of the most contrasting parental pairs, DOR364 and G19833, confirmed the above findings, indicating that those morphological characteristics are inheritable hence provide potential for genetic improvement. Root morphology in plants may be related to phosphorus (P) efficiency by affecting the absorption characteristics of the root system. However, genotypic variation in root morphological characteristics of common bean (Phaseolus vulgaris L.) as affected by P availability has not been well clarified. In the present study, systematic studies were conducted in a P-buffered sand culture system using three pairs of common bean parental materials with contrasting root traits in response to P deficiency. The results indicate that P availability significantly affects bean root morphology. Common bean tends to have smaller root system, shorter and coarser roots at low P availability. Genotypic variation in root morphology was observed among different genotypes in response to P availability. The P efficient genotypes appear to have larger, finer and longer root systems than the P inefficient genotypes, and such a variation was particularly obvious in the basal roots. From allomeric analysis, we found that morphological characteristics of the basal roots contribute more to P efficiency than those of the tap roots. Further studies with the F9 recombinant inbred lines derived from one of the most contrasting parental pairs, DOR364 and G19833, confirmed the above findings, indicating that those morphological characteristics are inheritable hence provide potential for genetic improvement.     

菜豆根形态特性的基因型差异与磷效率

应用磷控释砂培以及计算机图象分析技术,研究了磷效率差异显的菜豆（Phaseolus vulgaris L.)亲本及其重组自交系后代的根形态特性及其与磷效率的关系。试验结果表明,供磷状况显影响菜豆根系形态学特性。在低磷胁迫下,菜豆根系总根长变短、根部生物量减少,根直径增大。菜豆根形态特性对低磷有效性的适应性反应具有显的基因型差异。在低磷条件下磷高效率基因型的根系比磷低效率基因型相对根部生物量较大、总根长较长,根表面积较大。异计分析表明,菜豆基根根形态特性在低磷条件下的适应性变化对磷效率的贡献远远大于主根,并且这些适应性变化是可以遗传的,表明通过对菜豆根形态特性进行遗传改良来提高磷效率有一定的可行性。     

Effect of phosphorus deficiency on growth angle of basal roots in Phaseolus vulgaris

Root architectural plasticity might be an important factor in the acquisition by plants of immobile nutrients such as phosphorus (P). In this study, we examined the effect of P availability on the orientation of basal roots with respect to gravity, and thereby on the growth angle of these roots of common bean (Phaseolus vulgaris L.). In one set of studies the growth angle of basal roots of bean seedlings was measured over time. Sixteen bean genotypes were examined; six showed a decrease in root orientation with respect to gravity in low P media, one increased orientation, and nine showed no difference within 5 d of basal root emergence. Bean taproots also showed decreased root orientation with respect to gravity in low P. Growth angle after 5 d was correlated with field performance of contrasting genotypes in low P tropical soils. Mineral deficiencies other than P did not cause changes in root angle. In a split pouch system that provided high or low P solution to different parts of the root system, the decrease in root angle in low P was found to be a response to global P availability, and not local to the portion of the root system in low P. Effects of P availability on root angle were associated with reduced shoot P concentration, but preceded effects on plant biomass accumulation and leaf area expansion. Results from growth pouches for genotype G 19833 were confirmed using a solid-phase buffered sand-culture system supplying P at three levels. Pea (Pisum sativum), soybean (Glycine max Williams), chickpea (Cicer arietinum), lima bean (Phaseolus lunatus), and lentil (Lens culinaris) were grown with and without P; soybean and pea also showed decreased basal root angles in low P.     

Induction of maize acid phosphatase activities under phosphorus starvation

Large variation in phosphorus-(P) acquisition efficiency exists among maize inbred and hybrid genotypes. Acid phosphatases are a type of enzyme that affects P acquisition and P-use efficiency in plants. The objectives of this research were (1) to characterize acid phosphatase activity in maize grown hydroponically under P starvation, and (2) to determine if there is differential induction of acid phosphatases in two maize genotypes previously characterized as P efficient (Mo17) and P inefficient (B73). B73 and Mo17 seedlings were grown hydroponically and both intracellular and secreted acid phosphatase activities were characterized. Fresh seedling weight of both genotypes decreased under P starvation, but percent fresh weight allocated to roots increased 14 days after P starvation in B73. Soluble protein concentration in shoots and roots was affected little, but secreted protein decreased by 40 and 20% in B73 and Mo17 seedlings grown without P for 14 days. Intracellular and secreted acid phosphate activity increased substantially in leaves and roots in B73 and Mo17 in response to P starvation. Secreted APase activity per unit protein increased 310 and 300% in B73 and Mo17, respectively, 7 days after P withdrawal. One of the minor isozymes identified on non-denaturing PAGE, was increased specifically in response to P starvation in both maize genotypes. The patterns and levels of change in APase activities in B73 and Mo17 were not sufficiently different to account for the diverse growth response of these genotypes in low-P conditions. The results suggest that APases may not be a major mechanism for scavenging or acquiring P and changes in APases may reflect a state of P stress in both varieties. Other factors such as root architecture, secretion of low-molecular weight carboxylates and microbial interactions might explain the difference between these two genotypes.     

Genotypic variation in seedling root architectural traits and implications for drought adaptation in wheat (Triticum aestivum L.)

Root system characteristics are of fundamental importance to soil exploration and below-ground resource acquisition. Root architectural traits determine the in situ space-filling properties of a root system or root architecture. The growth angle of root axes is a principal component of root system architecture that has been strongly associated with acquisition efficiency in many crop species. The aims of this study were to examine the extent of genotypic variability for the growth angle and number of seminal roots in 27 current Australian and 3 CIMMYT wheat (Triticum aestivum L.) genotypes, and to quantify using fractal analysis the root system architecture of a subset of wheat genotypes contrasting in drought tolerance and seminal root characteristics. The growth angle and number of seminal roots showed significant genotypic variation among the wheat genotypes with values ranging from 36 to 56 (degrees) and 3 to 5 (plant⁻¹), respectively. Cluster analysis of wheat genotypes based on similarity in their seminal root characteristics resulted in four groups. The group composition reflected to some extent the genetic background and environmental adaptation of genotypes. Wheat cultivars grown widely in the Mediterranean environments of southern and western Australia generally had wider growth angle and lower number of seminal axes. In contrast, cultivars with superior performance on deep clay soils in the northern cropping region, such as SeriM82, Baxter, Babax, and Dharwar Dry exhibited a narrower angle of seminal axes. The wheat genotypes also showed significant variation in fractal dimension (D). The D values calculated for the individual segments of each root system suggested that, compared to the standard cultivar Hartog, the drought-tolerant genotypes adapted to the northern region tended to distribute relatively more roots in the soil volume directly underneath the plant. These findings suggest that wheat root system architecture is closely linked to the angle of seminal root axes at the seedling stage. The implications of genotypic variation in the seminal root characteristics and fractal dimension for specific adaptation to drought environment types are discussed with emphasis on the possible exploitation of root architectural traits in breeding for improved wheat cultivars for water-limited environments.     

The effect of phosphorus availability on the carbon economy of contrasting common bean (Phaseolus vulgaris L.) genotypes

A common response to low phosphorus availability is increased relative biomass allocation to roots. The resulting increase in root:shoot ratio presumably enhances phosphorus acquisition, but may also reduce growth rates by diverting carbon to the production of heterotrophic rather than photosynthetic tissues. To assess the importance of increased carbon allocation to roots for the adaptation of plants to low P availability, carbon budgets were constructed for four common bean genotypes with contrasting adaptation to low phosphorus availability in the field ("phosphorus efficiency"). Solid-phase-buffered silica sand provided low (1 microM), medium (10 microM), and high (30 microM) phosphorus availability. Compared to the high phosphorus treatment, plant growth was reduced by 20% by medium phosphorus availability and by more than 90% by low phosphorus availability. Low phosphorus plants utilized a significantly larger fraction of their daytime net carbon assimilation on root respiration (c. 40%) compared to medium and high phosphorus plants (c. 20%). No significant difference was found among genotypes in this respect. Genotypes also had similar rates of P absorption per unit root weight and plant growth per unit of P absorbed. However, P-efficient genotypes allocated a larger fraction of their biomass to root growth, especially under low P conditions. Efficient genotypes had lower rates of root respiration than inefficient genotypes, which enabled them to maintain greater root biomass allocation than inefficient genotypes without increasing overall root carbon costs.     

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.     

低磷胁迫下磷高效基因型大麦的根系形态特征

在根袋土培盆栽条件下,以磷高效基因型DH110+、DH147和低效基因型DH49大麦为试验材料,利用根系分析系统分析不同施磷(P₂O₅)水平(极低磷25 mg·kg^-1、低磷50 mg·kg^-1和正常磷75 mg·kg^-1)下,磷高效基因型大麦的根系形态特征及其与植株磷素吸收的关系.结果表明： 低磷胁迫显著降低大麦生物量和磷吸收量,其中磷高效基因型的生物量和磷吸收量在各施磷水平下分别为低效基因型的1.24～1.70和1.18～1.83倍;大麦的总根长、总根表面积、平均根系直径、不定根长及其根表面积、侧根长及其根表面积均随施磷水平的降低而显著降低,其中磷高效基因型大麦在各施磷水平下的总根长、总根表面积、比根长、侧根长及根表面积分别为低效基因型的1.46～2.06、1.12～1.51、1.35～1.72、1.69～2.42和1.40～1.78倍,而平均根系直径为低效基因型的70.6%～90.2%;主成分分析表明,平均根系直径、比根表面积和比根长受基因型差异的影响较为明显,是区分两类磷效率基因型大麦根系形态差异的主要指标;偏最小二乘回归分析表明,各施磷水平下,总根长、总根表面积对大麦植株磷素吸收贡献均较大,随施磷水平降低,不定根长、不定根表面积对大麦植株磷素吸收的贡献明显降低,而平均根系直径、比根长、侧根长及其根表面积的贡献明显增加.磷高效基因型大麦可通过维持侧根的生长、根细度和比根长的增加来适应低磷胁迫.     

The importance of root gravitropism for inter-root competition and phosphorus acquisition efficiency: results from a geometric simulation model

We have observed that low soil phosphorus availability alters the gravitropic response of basal roots in common bean (Phaseolus vulgaris L.), resulting in a shallower root system. In this study we use a geometric model to test the hypotheses that a shallower root system is a positive adaptive response to low soil P availability by (1) concentrating root foraging in surface soil horizons, which generally have the highest P availability, and (2) reducing spatial competition for P among roots of the same plant. The growth of nine root systems contrasting in gravitropic response over 320 h was simulated in SimRoot, a dynamic three-dimensional geometric model of root growth and architecture. Phosphorus acquisition and inter-root competition were estimated with Depzone, a program that dynamically models nutrient diffusion to roots. Shallower root systems had greater P acquisition per unit carbon cost than deeper root systems, especially in older root systems. This was due to greater inter-root competition in deeper root systems, as measured by the volume of overlapping P depletion zones. Inter-root competition for P was a significant fraction of total soil P depletion, and increased with increasing values of the P diffusion coefficient (De), with root age, and with increasing root gravitropism. In heterogenous soil having greater P availability in surface horizons, shallower root systems had greater P acquisition than deeper root systems, because of less inter-root competition as well as increased root foraging in the topsoil. Root P acquisition predicted by SimRoot was validated against values for bean P uptake in the field, with an r ² between observed and predicted values of 0.75. Our results support the hypothesis that altered gravitropic sensitivity in P-stressed roots, resulting in a shallower root system, is a positive adaptive response to low P availability by reducing inter-root competition within the same plant and by concentrating root activity in soil domains with the greatest P availability. This revised version was published online in August 2006 with corrections to the Cover Date.     

不同基因型小麦磷素营养阈值的研究

利用温室盆栽试验,对不同磷效率小麦基因型进行了磷素营养阈值的研究。结果表明,磷低效基因对缺磷反应敏感,在不施或施磷量较低时,籽粒产量及生物量均较低,随着施磷量增加,产量缓慢增加,表现出对磷肥效应较迟钝的钝的现象,而磷高效小麦基因型在不同施磷肥或施磷量较低条件下的产量与生物量相对较高,表现出明显的耐低磷特性及对土壤难溶性磷的高效活化、吸收能力。且随产丰施磷量增加,磷高效基因型的产量急剧增加,达到一定     

小麦与蚕豆间作体系根系形态与磷吸收的定量解析

豆科与禾本科作物间作能够改变作物根系生长,但不同施磷水平下间作-根系形态-磷吸收之间的关系尚未明确。本研究通过田间定位试验和根箱模拟试验,研究不同种植模式(小麦单作、蚕豆单作和小麦-蚕豆间作)和不同磷水平下小麦和蚕豆的产量、生物量、磷吸收及根系形态特征,分析探讨不同施磷条件下小麦-蚕豆间作对根系形态和磷吸收的影响。结果表明: 根箱试验中,与单作相比,间作小麦的根干重、根冠比分别增加21.2%、61.5%,地上部干重降低14.6%,根系磷含量和总吸磷量分别提高23.8%和12.1%;间作蚕豆的地上部干重、根干重、根冠比、总根长和根体积分别增加16.5%、47.3%、24.0%、3.5%和8.4%,间作蚕豆地上部磷含量、根系磷含量和总吸磷量分别提高14.7%、26.2%和21.5%。田间试验中,与单作相比,分蘖期间作小麦地上部磷吸收量降低8.7%,而在拔节期、抽穗期、灌浆期和成熟期分别提高40.6%、19.7%、7.8%和12.4%;但种间互作导致开花期、结荚期和成熟期间作蚕豆的地上部磷吸收量分别降低9.8%、9.0%和5.2%。偏最小二乘法(PLS)回归分析表明,小麦的根表面积和根体积、蚕豆的根表面积对作物磷吸收的贡献最大,在低磷胁迫条件下,间作同时提高了小麦和蚕豆的根体积和根表面积,促进了磷的吸收。总之,在缺磷或低磷条件下,种间互作扩大了小麦和蚕豆根土接触面积,促进了苗期磷的吸收,为后期间作优势的形成奠定了基础。     

Carbon cost of root systems: an architectural approach

Root architecture is an important component of nutrient uptake and may be sensitive to carbon allocational changes brought about by rising CO₂. We describe a deformable geometric model of root growth, SimRoot, for the dynamic morphological and physiological simulation of root architectures. Using SimRoot, and measurements of root biomass deposition, respiration and exudation, carbon/phosphorus budgets were developed for three contrasting root architectures. Carbon allocation patterns and phosphorus acquisition efficiencies were estimated for Phaseolus vulgaris seedlings with either a dichotomous, herringbone, or empirically determined bean root architecture. Carbon allocation to biomass, respiration, and exudation varied significantly among architectures. Root systems also varied in the relationship between C expenditure and P acquisition, providing evidence for the importance of architecture in nutrient acquisition efficiency.     

异质低磷胁迫下马尾松家系根构型和磷效率的遗传变异

以7个马尾松(Pinus massoniana)一代种子园自由授粉家系为材料, 设置同质低磷(P)胁迫和异质低P胁迫模拟的盆栽试验, 系统研究马尾松家系对不同类型低P胁迫的适应机制和P效率变异规律。结果表明, 参试马尾松家系的苗高、地径和生物量等P效率指标均表现出显著的家系变异, 主要P效率指标的家系遗传力均较高, 干物质积累量的广义遗传力大于0.80, 揭示了马尾松P营养效率的较大遗传改良潜力。马尾松对不同类型低P胁迫的适应机制有所差异。在同质低P胁迫下, ‘3201’、‘1217’等高P效率家系的根系主要参数均高于低P效率家系, 表明整体根系参数的适应性变化是P效率和生物量形成的决定因素。在异质低P胁迫下, 高P效率马尾松家系在表层富P介质的根系分布量、分布比例均显著增加, 表层根系参数与马尾松家系P效率呈显著正相关, 揭示根系空间构型的适应性变化是决定马尾松高P效率的重要生物学基础。表层根系生物量、表层根相对比例的家系遗传力达0.88和0.72, 证实了以马尾松根构型的适应变化为突破口, 选育具有理想根构型和较高P效率的马尾松家系。     

Root metaxylem and architecture phenotypes integrate to regulate water use under drought stress

At the genus and species level, variation in root anatomy and architecture may interact to affect strategies of drought avoidance. To investigate this idea, root anatomy and architecture of the drought‐sensitive common bean (Phaseolus vulgaris) and drought‐adapted tepary bean (Phaseolus acutifolius) were analyzed in relation to water use under terminal drought. Intraspecific variation for metaxylem anatomy and axial conductance was found in the roots of both species. Genotypes with high‐conductance root metaxylem phenotypes acquired and transpired more water per unit leaf area, shoot mass, and root mass than genotypes with low‐conductance metaxylem phenotypes. Interspecific variation in root architecture and root depth was observed where P. acutifolius has a deeper distribution of root length than P. vulgaris. In the deeper‐rooted P. acutifolius, genotypes with high root conductance were better able to exploit deep soil water than genotypes with low root axial conductance. Contrastingly, in the shallower‐rooted P. vulgaris, genotypes with low root axial conductance had improved water status through conservation of soil moisture for sustained water capture later in the season. These results indicate that metaxylem morphology interacts with root system depth to determine a strategy of drought avoidance and illustrate synergism among architectural and anatomical phenotypes for root function.     

Basal root whorl number: a modulator of phosphorus acquisition in common bean (Phaseolus vulgaris)

Background and Aims

Root architectural phenes enhancing topsoil foraging are important for phosphorus acquisition. In this study, the utility of a novel phene is described, basal root whorl number (BRWN), that has significant effects on topsoil foraging in common bean (Phaseolus vulgaris).

Methods

Whorls are defined as distinct tiers of basal roots that emerge in a tetrarch fashion along the base of the hypocotyl. Wild and domesticated bean taxa as well as two recombinant inbred line (RIL) populations were screened for BRWN and basal root number (BRN). A set of six RILs contrasting for BRWN was evaluated for performance under low phosphorus availability in the greenhouse and in the field. In the greenhouse, plants were grown in a sand–soil media with low or high phosphorus availability. In the field, plants were grown in an Oxisol in Mozambique under low and moderate phosphorus availability.

Key Results

Wild bean accessions tended to have a BRWN of one or two, whereas cultivated accessions had BRWN reaching four and sometimes five. BRWN and BRN did not vary with phosphorus availability, i.e. BRWN was not a plastic trait in these genotypes. Greater BRWN was beneficial for phosphorus acquisition in low phosphorus soil. Genotypes with three whorls had almost twice the shoot biomass, greater root length and greater leaf area than related genotypes with two whorls. In low phosphorus soil, shoot phosphorus content was strongly correlated with BRWN (R² = 0·64 in the greenhouse and R² = 0·88 in the field). Genotypes with three whorls had shallower root systems with a greater range of basal root growth angles (from 10 to 45 ° from horizontal) than genotypes with two whorls (angles ranged from 60 to 85 ° from horizontal).

Conclusions

The results indicate that BRWN is associated with increased phosphorus acquisition and that this trait may have value for selection of genotypes with better performance in low phosphorus soils.     

Modelling root plasticity and response of narrow-leafed lupin to heterogeneous phosphorus supply

Background & Aims

Searching for root traits underpinning efficient nutrient acquisition has received increased attention in modern breeding programs aimed at improved crop productivity. Root models provide an opportunity to investigate root-soil interactions through representing the relationships between rooting traits and the non-uniform supply of soil resources. This study used simulation modelling to predict and identify phenotypic plasticity, root growth responses and phosphorus (P) use efficiency of contrasting Lupinus angustifolius genotypes to localised soil P in a glasshouse.

Methods

Two L. angustifolius genotypes with contrasting root systems were grown in cylindrical columns containing uniform soil with three P treatments (nil and 20 mg P kg^?1 either top-dressed or banded) in the glasshouse. Computer simulations were carried out with root architecture model ROOTMAP which was parameterized with root architectural data from an earlier published hydroponic phenotyping study.

Results

The experimental and simulated results showed that plants supplied with banded P had the largest root system and the greatest P-uptake efficiency. The P addition significantly stimulated root branching in the topsoil, whereas plants with nil P had relatively deeper roots. Genotype-dependent root growth plasticity in response to P supply was shown, with the greatest response to banded P.

Conclusions

Both experimental and simulation outcomes demonstrated that 1) root hairs and root proliferation increased plant P acquisition and were more beneficial in the localised P fertilisation scenario, 2) placing P deeper in the soil might be a more effective fertilisation method with greater P uptake than top dressing, and 3) the combination of P foraging strategies (including root architecture, root hairs and root growth plasticity) is important for efficient P acquisition from a localised source of fertiliser P.     

Phosphorus allocation in P-efficient and inefficient barley cultivars as affected by mycorrhizal infection

A glasshouse experiment was undertaken to investigate the effect of mycorrhizal infection on the allocation of phosphorus (P) in agronomically P-efficient (i.e. high yields at low P supply) and inefficient barley (Hordeum vulgare L.) cultivars. Four barley cultivars differing in agronomic P-efficiency were inoculated or not inoculated with Glomus etunicatum. Cultivars did not differ in percentage of root length infected. The concentration of P in roots of the inefficient cultivars was higher than that of the efficient cultivars. However, because of changes in root to shoot dry weight ratio and below-ground productivity, mycorrhizal infection significantly reduced the percentage of total plant P in roots of the inefficient cultivars. The distribution of P between root and shoot of P-efficient cultivars was not affected by mycorrhizal infection. Root to shoot dry weight ratio of the P-efficient cultivars was lower than that of the inefficient cultivars, and the decrease in the ratio following infection was significant in inefficient but not in P-efficient cultivars. This study indicates that mycorrhizal infection alters the allocation of P in inefficient cultivars and effectively improves the efficiency of P utilization with respect to shoot growth.