Steep,cheap and deep: an ideotype to optimize water and N acquisition by maize root systems

Background

A hypothetical ideotype is presented to optimize water and N acquisition by maize root systems. The overall premise is that soil resource acquisition is optimized by the coincidence of root foraging and resource availability in time and space. Since water and nitrate enter deeper soil strata over time and are initially depleted in surface soil strata, root systems with rapid exploitation of deep soil would optimize water and N capture in most maize production environments.• The ideotype Specific phenes that may contribute to rooting depth in maize include (a) a large diameter primary root with few but long laterals and tolerance of cold soil temperatures, (b) many seminal roots with shallow growth angles, small diameter, many laterals, and long root hairs, or as an alternative, an intermediate number of seminal roots with steep growth angles, large diameter, and few laterals coupled with abundant lateral branching of the initial crown roots, (c) an intermediate number of crown roots with steep growth angles, and few but long laterals, (d) one whorl of brace roots of high occupancy, having a growth angle that is slightly shallower than the growth angle for crown roots, with few but long laterals, (e) low cortical respiratory burden created by abundant cortical aerenchyma, large cortical cell size, an optimal number of cells per cortical file, and accelerated cortical senescence, (f) unresponsiveness of lateral branching to localized resource availability, and (g) low K_m and high V_max for nitrate uptake. Some elements of this ideotype have experimental support, others are hypothetical. Despite differences in N distribution between low-input and commercial maize production, this ideotype is applicable to low-input systems because of the importance of deep rooting for water acquisition. Many features of this ideotype are relevant to other cereal root systems and more generally to root systems of dicotyledonous crops.     

QTL mapping and phenotypic variation for root architectural traits in maize (Zea mays L.)

Key message

QTL were identified for root architectural traits in maize.

Abstract

Root architectural traits, including the number, length, orientation, and branching of the principal root classes, influence plant function by determining the spatial and temporal domains of soil exploration. To characterize phenotypic patterns and their genetic control, three recombinant inbred populations of maize were grown for 28 days in solid media in a greenhouse and evaluated for 21 root architectural traits, including length, number, diameter, and branching of seminal, primary and nodal roots, dry weight of embryonic and nodal systems, and diameter of the nodal root system. Significant phenotypic variation was observed for all traits. Strong correlations were observed among traits in the same root class, particularly for the length of the main root axis and the length of lateral roots. In a principal component analysis, relationships among traits differed slightly for the three families, though vectors grouped together for traits within a given root class, indicating opportunities for more efficient phenotyping. Allometric analysis showed that trajectories of growth for specific traits differ in the three populations. In total, 15 quantitative trait loci (QTL) were identified. QTL are reported for length in multiple root classes, diameter and number of seminal roots, and dry weight of the embryonic and nodal root systems. Phenotypic variation explained by individual QTL ranged from 0.44 % (number of seminal roots, NyH population) to 13.5 % (shoot dry weight, OhW population). Identification of QTL for root architectural traits may be useful for developing genotypes that are better suited to specific soil environments.     

Genetic variability for root hair traits as related to phosphorus status in soybean

Plant root hairs are believed to be very important for phosphorus (P) uptake from the soil by expanding the absorptive surface area of the root and increasing the soil volume explored by the roots, but genetic information about root hair traits in soybean is relatively scarce. In the present study, two contrasting genotypes of soybean (Glycine max and Glycine soja), CN4 and XM6, and their 88 F₉-derived recombinant inbred lines (RILs) were grown in a field with moderately low P availability. Some important root hair traits, including root hair density (RHD), average root hair length (ARHL), and root hair length per unit root (RHLUR) were investigated and quantified with an automatic image analysis system and the genetic variability for these root hair traits was estimated with the RIL population. The results indicated that the two parental genotypes differed significantly in the three root hair traits measured, with XM6 generally having larger RHD and RHLUR (but smaller ARHL) than CN4, which may in part explain the difference in biomass and P status between the two parents. All the three root hair traits were continually segregated in the progenial RIL population with a normal distribution of the phenotypic values, indicating that these traits are possibly controlled by quantitative trait loci (QTLs). Analysis of variance for the RIL population showed that RHD had a low heritability (h² _b = 27.32, 31.04, 33.97% for basal roots, tap roots, total roots, respectively), while ARHL had a relatively higher genetic variance and hence a higher heritability (h² _b = 53.85, 59.18, 60.98% for basal roots, tap roots, total roots, respectively), suggesting that RHD is influenced more by environmental factors than ARHL. Both RHD and ARHL were positively correlated with RHLUR, indicating that the former two traits may be the attributes to the latter. On the other hand, RHD and ARHL were negatively correlated with each other, implying a possible complementary relationship between the two traits. Both RHD and RHLUR (but not ARHL) were positively correlated with P concentration in the plant, suggesting an important role of root hairs in P status. The basal roots had denser and higher total root hair length than the tap roots, and this is in accordance with previous observations with other plants that basal roots are more effective for P uptake than tap roots in cultivated soils.     

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.     

Estimation of tree root lengths using fractal branching rules: a comparison with soil coring for Grevillea robusta

Previous theoretical research has suggested that lengths of tree roots can be estimated on the basis of their branching characteristics, if branching has a fractal pattern that is independent of root diameter. This theory and its underlying assumptions was tested for Grevillea robusta trees at a site in Kenya by comparing estimates of root length from conventional soil coring and the output of a fractal branching algorithm. The trees were in a 4-year-old stand established on a 3 × 4 m planting grid. Root lengths (L _r) in four units of the planting grid were estimated by soil coring. Branching characteristics determined by examination of 32 excavated roots from 16 trees were: The number of branches at each branching point; the length of links between branching points (L _l); the diameter of root tips; and parameters which describe the change in diameter at each branching point. Each was found to be independent of root size. These data were used to parameterise a branching algorithm, which was then used to estimate numbers of root links in the four grid units (n _l) from root diameters at the bases of the four trees at the corners of each unit. Root lengths, from L _r = n₁ L₁, severely underestimated L _r. This discrepancy probably resulted from inaccuracy in the parameterisation of the branching algorithm, as output from the algorithm was very sensitive to small changes in parameter values. Use of fractal branching rules alone to estimate roots length does not appear possible unless the algorithm is calibrated to adjust for errors in parameter estimation. Calibration can be achieved by calculation of an 'effective link length', L ^eff ₁, from L _r/n _l, where L _r is measured by a reference method such as soil coring.     

Fine Root Morphology,Biochemistry and Litter Quality Indices of Fast- and Slow-growing Woody Species in Ethiopian Highland Forest

Fine root turnover of trees is a major C input to soil. However, the quality of litter input is influenced by root morphological traits and tissue chemical composition. In this study, fine roots of ten tropical woody species were collected from an Afromontane forest in the northern highlands of Ethiopia. The fine roots were analysed for root morphological traits and tissue chemistry measured as proxy carbon fractionations. Based on stem increment, the 10 species were divided into faster- and slower-growing species. Faster-growing species exhibited higher specific root length (1362 cm g^?1) than slower-growing species (923 cm g^?1). Similarly specific root area was higher in faster-growing species (223 cm² g^?1) than in slower-growing species (167 cm² g^?1). Among the carbon fractions, the acid-insoluble fraction (AIF) was the highest (44–51%). The carbon content, AIF, and the lignocellulose index were higher for slower-growing species. Root tissue density was lower in faster-growing species (0.33 g cm^?3) than slower-growing species (0.40 g cm^?3) and showed a strong positive correlation with carbon content (r ² = 0.84) and the AIF (r _pearson = 0.93). The morphological traits of fine roots between faster- and slower-growing species reflect the ecological strategy they employ. Slower-growing species have a higher tissue density which may reflect a greater longevity.     

QTL mapping and epistasis analysis of brace root traits in maize

Root architecture is a major factor influencing root lodging, which limits greater yield stability at high planting density. Total brace root tier number (TBRTN) and effective brace root tier number (EBRTN) are the two most important root architecture traits influencing root lodging. However, the genetic mechanisms that underlie these traits remain poorly understood. In this study, quantitative trait loci (QTL) for TBRTN and EBRTN were mapped using a set of 201 recombinant inbred lines (RILs) and 278 immortalized F2 (IF2) populations derived from these RILs, which were evaluated in three environments. Ten QTL in the RILs and 15 QTL in the IF2 population were detected. In the two populations, we identified two coincident major QTL for TBRTN and a single identical major QTL for EBRTN. The QTL for TBRTN showed the largest additive effect, accounting for 16.36 and 17.88% of the phenotypic variance in the RILs and IF2 population, respectively. Additional epistatic effects were identified for all the maize chromosomes, except for chromosome 4. Most epistatic effects involved pairs of loci that were on different chromosomes. At the same time, we found loci that interacted simultaneously with several other loci to affect expression of the traits, which was particularly evident in the IF2 population. For example, qTAR1-2 interacted simultaneously with qTAR2-1, qTAR3-1, qTAR5-1, and qITAR8-2 to affect the expression of TBRTN. Therefore, a complex network controlling the traits was found in maize. These results provide useful information for understanding the molecular mechanisms controlling root architecture.     

黄河三角洲滨海盐碱地11个造林树种细根解剖性状对土壤条件的响应

细根作为植物与土壤连接的重要部位,能够反映植物对生存环境的适应性。以黄河三角洲滨海盐碱地不同立地条件下11个造林树种为对象,基于细根分支等级划分1-4级根序并进行解剖特征测定,分析细根解剖性状对滨海盐碱地不同土壤条件的响应规律。结果表明：（1）不同根序的细根直径存在显著差异,细根直径随根序升高呈增大趋势,而同根序的细根直径在不同树种间表现出显著的种间差异（P < 0.05）。1-2级细根皮层厚度、3-4级细根导管密度在树种间的差异均达显著水平（P < 0.05）。（2）在较为严重盐渍化土壤条件下（立地1）,细根皮层厚度较其他立地显著增大,但细根导管密度较小;在轻度盐碱立地条件下（立地3）,细根导管密度较大;较为严重的盐碱立地具有更为发达的细根直径及维管柱直径。（3）树种1-2级细根解剖结构与土壤环境关系最为密切,其中1级根直径与土壤pH值显著正相关（P < 0.05）,与土壤硝态氮含量呈显著负相关（P < 0.05）。对土壤理化性质与细根解剖性状的冗余分析表明,前两个轴的特征值达0.640和0.196,土壤速效养分含量与轴一（RDA1）呈正相关,低级根解剖性状则与轴二（RDA2）呈显著负相关。低级根解剖结构以及土壤的pH值能解释较多树种的差异性,其中低级根直径与皮层厚度对盐碱环境表现出较强的响应。     

Focus Issue on Roots: Branching Out in Roots: Uncovering Form,Function, and Regulation

Root branching is critical for plants to secure anchorage and ensure the supply of water, minerals, and nutrients. To date, research on root branching has focused on lateral root development in young seedlings. However, many other programs of postembryonic root organogenesis exist in angiosperms. In cereal crops, the majority of the mature root system is composed of several classes of adventitious roots that include crown roots and brace roots. In this Update, we initially describe the diversity of postembryonic root forms. Next, we review recent advances in our understanding of the genes, signals, and mechanisms regulating lateral root and adventitious root branching in the plant models Arabidopsis (Arabidopsis thaliana), maize (Zea mays), and rice (Oryza sativa). While many common signals, regulatory components, and mechanisms have been identified that control the initiation, morphogenesis, and emergence of new lateral and adventitious root organs, much more remains to be done. We conclude by discussing the challenges and opportunities facing root branching research.Branching through lateral and adventitious root formation represents an important element of the adaptability of the root system to its environment. Both are regulated by nutrient and hormonal signals (Bellini et al., 2014; Giehl and von Wirén, 2014), which act locally to induce or inhibit root branching. The net effect of these adaptive responses is to increase the surface area of the plant root system in the most important region of the soil matrix for resource capture (e.g. surface layers for phosphorus uptake and deeper layers for nitrate uptake) or to secure anchorage. Different species use different combinations of lateral or adventitious roots to achieve this, with lateral roots dominating the root system of eudicots while adventitious (crown and brace) roots dominate the root system of monocots, including in cereal crops.Our understanding of the mechanisms controlling lateral and adventitious root development has accelerated in recent years, primarily through research on model plants. The simple anatomy and the wealth of genetic resources in Arabidopsis (Arabidopsis thaliana) have greatly aided embryonic and postembryonic root developmental studies (De Smet et al., 2007; Péret et al., 2009a; Fig. 1, A and E). Nevertheless, impressive recent progress has been made studying root branching in other crop species, notably cereals such as maize (Zea mays) and rice (Oryza sativa).Open in a separate windowFigure 1.A to D, Schematics showing diversity in root system architecture at both seedling (left) and mature (right) stages in eudicots (A and C) and monocots (B and D). A, Arabidopsis root system. B, Maize root system. C, Tomato root system (for clarity, stem-derived adventitious roots are only shown in the labeled region). D, Wheat root system. E and F, Cross sections of emerging lateral root primordia in Arabidopsis (E) and rice (F). E and F are adapted from Péret et al. (2009b).In this Update, we initially describe the diversity of postembryonic root forms in eudicots and monocots (Fig. 1). Next, we highlight recent advances in our understanding of the genes, signals, and mechanisms regulating lateral root and adventitious root branching in Arabidopsis, rice, and maize. Due to space limits, we cannot provide an exhaustive review of this subject area, focusing instead on recent research advances. However, we direct readers to several recent in-depth reviews on lateral root (Lavenus et al., 2013; Orman-Ligeza et al., 2013) and adventitious root development (Bellini et al., 2014).     

Genetic dissection of nitrogen nutrition in pea through a QTL approach of root, nodule, and shoot variability

Pea (Pisum sativum L.) is the third most important grain legume worldwide, and the increasing demand for protein-rich raw material has led to a great interest in this crop as a protein source. Seed yield and protein content in crops are strongly determined by nitrogen (N) nutrition, which in legumes relies on two complementary pathways: absorption by roots of soil mineral nitrogen, and fixation in nodules of atmospheric dinitrogen through the plant–Rhizobium symbiosis. This study assessed the potential of naturally occurring genetic variability of nodulated root structure and functioning traits to improve N nutrition in pea. Glasshouse and field experiments were performed on seven pea genotypes and on the ‘Cameor’ × ‘Ballet’ population of recombinant inbred lines selected on the basis of parental contrast for root and nodule traits. Significant variation was observed for most traits, which were obtained from non-destructive kinetic measurements of nodulated root and shoot in pouches, root and shoot image analysis, ¹⁵N quantification, or seed yield and protein content determination. A significant positive relationship was found between nodule establishment and root system growth, both among the seven genotypes and the RIL population. Moreover, several quantitative trait loci for root or nodule traits and seed N accumulation were mapped in similar locations, highlighting the possibility of breeding new pea cultivars with increased root system size, sustained nodule number, and improved N nutrition. The impact on both root or nodule traits and N nutrition of the genomic regions of the major developmental genes Le and Af was also underlined.     

Fine-root morphological and growth traits in a Turkey-oak stand in relation to seasonal changes in soil moisture in the Southern Apennines, Italy

We investigated the effects of seasonal changes in soil moisture on the morphological and growth traits of fine roots (<2?mm in diameter) in a mature Turkey-oak stand (Quercus cerris L.) in the Southern Apennines of Italy. Root samples (diameter:?<0.5, 0.5?C1.0, 1.0?C1.5, and 1.5?C2.0?mm) were collected with the Auger method. Mean annual fine-root mass and length on site was 443?g?m^?2 (oak fine roots 321?g?m^?2; other species 122?g?m^?2) and 3.18?km?m^?2 (oak fine roots 1.14?km?m^?2; other species 2.04?km?m^?2), respectively. Mean specific root length was 8.3?m?g^?1. All fine-root traits displayed a complex pattern that was significantly related to season. In the four diameter classes, both fine-root biomass and length peaked in summer when soil water content was the lowest and air temperature the highest of the season. Moreover, both fine-root biomass and length were inversely related with soil moisture (p?<?0.001). The finest roots (<0.5?mm in diameter) constituted an important fraction of total fine-root length (79?%), but only 21?% of biomass. Only in this root class, consequent to change in mean diameter, specific root length peaked when soil water content was lowest showing an inverse relationship (p?<?0.001). Furthermore, fine-root production and turnover decreased with increasing root diameter. These results suggest that changes in root length per unit mass, and pulses in root growth to exploit transient periods of low soil water content may enable trees to increase nutrient and water uptake under seasonal drought conditions.     

Assessing variability in root traits of wild Lupinus angustifolius germplasm: basis for modelling root system structure

Background and aims

Intra-specific variation in root system architecture and consequent efficiency of resource capture by major crops has received recent attention. The aim of this study was to assess variability in a number of root traits among wild genotypes of narrow-leafed lupin (Lupinus angustifolius L.), to provide a basis for modelling of root structure.

Methods

A subset of 111 genotypes of L. angustifolius was selected from a large germplasm pool based on similarity matrices calculated using Diversity Array Technology markers. Plants were grown for 6?weeks in the established semi-hydroponic phenotyping systems to measure the fine-scale features of the root systems.

Results

Root morphology of wild L. angustifolius was primarily dominated by the taproot and first-order branches, with the presence of densely or sparsely distributed second-order branches in the late growth stage. Large variation in most root traits was identified among the tested genotypes. Total root length, branch length and branch number in the entire root system and in the upper roots were the most varied traits (coefficient of variation CV >0.50). Over 94% of the root system architectural variation determined from the principal components analysis was captured by six components (eigenvalue >1). Five relatively homogeneous groups of genotypes with distinguished patterns of root architecture were separated by k-means clustering analysis.

Conclusions

Variability in the fine-scale features of root systems such as branching behaviour and taproot growth rates provides a basis for modelling root system structure, which is a promising path for selecting desirable root traits in breeding and domestication of wild and exotic resources of L. angustifolius for stressful or poor soil environments.     

Changes in water relations for Acacia ancistrocarpa on natural and mine-rehabilitation sites in response to an experimental wetting pulse in the Great Sandy Desert

Vegetation plays an essential role in mine-site rehabilitation as it serves to reduce erosion and extract moisture from the soil cover, thus minimising deep drainage to potentially hazardous materials below. Plant water-use patterns and root development are likely to be impacted where subsoils are physically and/or chemically inhospitable and close to the surface, particularly in arid and semi-arid environments where pulses of rainfall are infrequent and interspersed with long dry periods. The aim of this investigation was to determine how plant water relations of the woody shrub species Acacia ancistrocarpa (Maiden and Blakely) were affected by a summer wetting pulse on a degraded mine-rehabilitation site and a pristine natural site in the Great Sandy Desert of north-western Australia. At both the natural and rehabilitation sites, 20 m³ of water was applied to emulate an 80-mm rainfall event comparable with cyclones that occur during the summer wet season. We found that plants responded with significant increases in stomatal conductance, leaf water potential and sap-flow in lateral roots within three days of irrigation at the natural site and two days at the rehabilitation site. At the rehabilitation site this response occurred despite the fact that sinker roots were stunted by impeding physical properties of the mine-waste material. Sap-flow velocity, using the heat ratio method, in lateral and primary sinker roots was close to zero at night and positive during the day for trees at both sites during the dry period before the irrigation. These data indicate hydraulic redistribution was not occurring between the primary sinker and lateral roots at night, and water was transported along the laterals towards the crown, despite the fact they were in dry soil. Excavations revealed that lateral roots extended up to 6 m and displayed secondary sinker roots accessing water from deeper soil layers, even from mine-waste material. These morphological traits likely improved the water relations and survival of A. ancistrocarpa at the rehabilitation site. Considering these traits and the rapid pulse responsiveness of this species, we recommend A. ancistrocarpa for future rehabilitation projects at this and other mine-sites in the region.     

Experimentally reduced root–microbe interactions reveal limited plasticity in functional root traits in Acer and Quercus

Background and Aims

Interactions between roots and soil microbes are critical components of below-ground ecology. It is essential to quantify the magnitude of root trait variation both among and within species, including variation due to plasticity. In addition to contextualizing the magnitude of plasticity relative to differences between species, studies of plasticity can ascertain if plasticity is predictable and whether an environmental factor elicits changes in traits that are functionally advantageous.

Methods

To compare functional traits and trait plasticities in fine root tissues with natural and reduced levels of colonization by microbial symbionts, trimmed and surface-sterilized root segments of 2-year-old Acer rubrum and Quercus rubra seedlings were manipulated. Segments were then replanted into satellite pots filled with control or heat-treated soil, both originally derived from a natural forest. Mycorrhizal colonization was near zero in roots grown in heat-treated soil; roots grown in control soil matched the higher colonization levels observed in unmanipulated root samples collected from field locations.

Key Results

Between-treatment comparisons revealed negligible plasticity for root diameter, branching intensity and nitrogen concentration across both species. Roots from treated soils had decreased tissue density (approx. 10–20 %) and increased specific root length (approx. 10–30 %). In contrast, species differences were significant and greater than treatment effects in traits other than tissue density. Interspecific trait differences were also significant in field samples, which generally resembled greenhouse samples.

Conclusions

The combination of experimental and field approaches was useful for contextualizing trait plasticity in comparison with inter- and intra-specific trait variation. Findings that root traits are largely species dependent, with the exception of root tissue density, are discussed in the context of current literature on root trait variation, interactions with symbionts and recent progress in standardization of methods for quantifying root traits.     

华北山地6个外生菌根树种的根属性及其与菌根侵染率的关系

该研究以共存于同一暖温带森林的6个外生菌根(ECM)树种为研究对象,测定分析不同根序(1～5级)和功能根系(吸收细根和运输细根)的主要形态和构型属性及ECM侵染率,探究不同外生菌根树种的根属性变异模式及其与菌根真菌侵染程度的关系。结果表明:(1)随着根序的增加,不同树种根直径和单根长度均增加,而比根长和根分支强度均降低;根属性在同一根序下均存在显著的种间差异,尤其是2个裸子植物(落叶松和油松)的根直径较其他4个被子植物大。(2)同一树种的所有根属性在吸收细根和运输细根之间均有显著差异;吸收细根和运输细根的根直径、比根长和根组织密度在树种间均存在显著差异,而其单根长度和根分支强度在树种间无显著差异。(3)ECM侵染率以落叶松最高,千金榆和白桦最低,且与根尖直径呈显著正相关关系,与根尖比根长呈显著负相关关系。研究发现,基于根序或者功能根系,根属性在种间的变异模式不完全一致,单根长度和根分支强度在两个功能根系中均没有表现出显著的种间差异;吸收细根的比根长和根分支强度的变异系数较大,对环境变化有较敏感的响应;古老树种的根直径相对较粗,对菌根真菌的依赖性更高。     

Early vertical distribution of roots and its association with drought tolerance in tropical maize

Background and aims

Selection for deep roots to improve drought tolerance of maize (Zea mays L.) requires presence of genetic variation and suitable screening methods.

Methods

We examined a diverse set of 33 tropical maize inbred lines that were grown in growth columns in the greenhouse up to the 2-, 4-, and 6-leaf stage and in the field in Mexico. To determine length of roots from different depths at high throughput, we tested an approach based on staining roots with methylene blue and measuring the amount of absorbed dye as proxy measure for root length.

Results

Staining provided no advantage over root weights that are much easier to measure and therefore preferable. We found significant genotypic variation for all traits at the 6-leaf stage. For development rates between the 2-leaf and the 6-leaf stage, genotypes only differed for rooting depth and the number of crown roots. Positive correlations of leaf area with root length and rooting depth indicated a common effect of plant vigor. However, leaf area in growth columns was negatively related to grain yield under drought (r?=??0.50).

Conclusion

The selection for deeper roots by an increase in plant vigor likely results in a poorer performance under drought conditions. The proportion of deep roots was independent of other traits but showed a low heritability and was not correlated to field performance. An improved screening protocol is proposed to increase throughput and heritability for this trait.

     

Spatial distribution and phenotypic variation in root cortical aerenchyma of maize (Zea mays L.)

Background and aims

Previous research has suggested that root cortical aerenchyma (RCA) can enhance soil exploration and crop performance by decreasing root respiration. This trait is a potential breeding target for adaptation to abiotic stresses such as drought and low nutrient availability. However, little is known of phenotypic variation in aerenchyma or its distribution among root classes.

Methods

The spatial distribution of RCA was evaluated in freehand sections from 13 sites in the root systems of 11 recombinant inbred and commercial lines of maize (Zea mays). RCA variation was evaluated in 583 recombinant inbred lines of maize at one sampling position.

Results

RCA varied significantly among root classes and axial positions. Genotypic differences were observed for the amount of RCA at corresponding sampling locations and for the mean amount of RCA across all sampling locations, but genotypes did not differ in the proportional distribution of RCA within the whole root system. The amount of RCA in a cross-section was independent of several other anatomical traits.

Conclusions

There is substantial genetic variation for RCA, and this variation is independent of other anatomical traits. RCA can be phenotyped in greenhouse-grown plants by sampling the middle parts of second- or third-whorl crown roots.     

Distribution of Hydrogen-Metabolizing Bacteria in Alfalfa Field Soil

H₂ evolved by alfalfa root nodules during the process of N₂ fixation may be an important factor influencing the distribution of soil bacteria. To test this hypothesis under field conditions, over 700 bacterial isolates were obtained from fallow soil or from the 3-mm layer of soil surrounding alfalfa (Medicago sativa L.) root nodules, alfalfa roots, or bindweed (Convolvulus arvensis L.) roots. Bacteria were isolated under either aerobic or microaerophilic conditions and were tested for their capacity to metabolize H₂. Isolates showing net H₂ uptake and ³H₂ incorporation activity under laboratory conditions were assigned a Hup⁺ phenotype, whereas organisms with significant H₂ output capacity were designated as a Hout⁺ phenotype. Under aerobic isolation conditions two Hup⁺ isolates were obtained, whereas under microaerophilic conditions five Hup⁺ and two Hout⁺ isolates were found. The nine isolates differed on the basis of 24 standard bacteriological characteristics or fatty acid composition. Five of the nine organisms were isolated from soil around root nodules, whereas the other four were found distributed among the other three soil environments. On the basis of the microaerophilic isolations, 4.8% of the total procaryotic isolates from soil around root nodules were capable of oxidizing H₂, and 1.2% could produce H₂. Two of the Hup⁺ isolates were identified as Rhizobium meliloti by root nodulation tests, but the fact that none of the isolates reduced C₂H₂ under the assay conditions suggested that the H₂ metabolism traits were associated with various hydrogenase systems rather than with nitrogenase activity. Results from this study support the concept that H₂ evolution by alfalfa root nodules has a significant effect on the surrounding microenvironment and influences the number and diversity of bacteria occupying that region.     

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.     

Insights into root growth, function, and mycorrhizal abundance from chemical and isotopic data across root orders

Background and aims

Detailed analyses of root chemistry by branching order may provide insights into root function, root lifespan and the abundance of root-associated mycorrhizal fungi in forest ecosystems.

Methods

We examined the nitrogen and carbon stable isotopes (δ¹⁵N and δ¹³C) and concentration (%N and %C) in the fine roots of an arbuscular mycorrhizal tree, Fraxinus mandshurica, and an ectomycorrhizal tree, Larix gmelinii, over depth, time, and across five root branching orders.

Results and conclusions

Larix δ¹⁵N increased by 2.3?‰ from 4th order to 1st order roots, reflecting the increased presence of ¹⁵N-enriched ECM fungi on the lower root orders. In contrast, arbuscular mycorrhizal Fraxinus only increased by 0.7?‰ from 4th order to 1st order roots, reflecting the smaller ¹⁵N enrichment and lower fungal mass on arbuscular mycorrhizal fine roots. Isotopic and anatomical mass balance calculations indicate that first, second, and third order roots in ectomycorrhizal Larix averaged 36 %, 23 %, and 8 % fungal tissue by mass, respectively. Using literature values of root production by root branching order, we estimate that about 25 % of fine root production in ECM species like Larix is actually of fungal sheaths. In contrast to %N, %C, and δ¹⁵N, δ¹³C changed minimally across depth, time, and branching order. The homogeneity of δ¹³C suggests root tissues are constructed from a large well-mixed reservoir of carbon, although compound specific δ¹³C data is needed to fully interpret these patterns. The measurements developed here are an important step towards explicitly including mycorrhizal production in forest ecosystem carbon budgets.