Determination of wheat spike and spikelet architecture and grain traits using X-ray Computed Tomography imaging.

BACKGROUND: Wheat spike architecture is a key determinant of multiple grain yield components and detailed examination of spike morphometric traits is beneficial to explain wheat grain yield and the effects of differing agronomy and genetics. However, quantification of spike morphometric traits has been very limited because it relies on time-consuming manual measurements. RESULTS: In this study, using X-ray Computed Tomography imaging, we proposed a method to efficiently detect the 3D architecture of wheat spikes and component spikelets by clustering grains based on their Euclidean distance and relative positions. Morphometric characteristics of wheat spikelets and grains, e.g., number, size and spatial distribution along the spike can be determined. Two commercial wheat cultivars, one old, Maris Widgeon, and one modern, Siskin, were studied as examples. The average grain volume of Maris Widgeon and Siskin did not differ, but Siskin had more grains per spike and therefore greater total grain volume per spike. The spike length and spikelet number were not statistically different between the two cultivars. However, Siskin had a higher spikelet density (number of spikelets per unit spike length), with more grains and greater grain volume per spikelet than Maris Widgeon. Spatial distribution analysis revealed the number of grains, the average grain volume and the total grain volume of individual spikelets varied along the spike. Siskin had more grains and greater grain volumes per spikelet from spikelet 6, but not spikelet 1-5, compared with Maris Widgeon. The distribution of average grain volume along the spike was similar for the two wheat cultivars. CONCLUSION: The proposed method can efficiently extract spike, spikelet and grain morphometric traits of different wheat cultivars, which can contribute to a more detailed understanding of the sink of wheat grain yield.

Plant roots sense soil compaction through restricted ethylene diffusion.

Soil compaction represents a major challenge for modern agriculture. Compaction is intuitively thought to reduce root growth by limiting the ability of roots to penetrate harder soils. We report that root growth in compacted soil is instead actively suppressed by the volatile hormone ethylene. We found that mutant Arabidopsis and rice roots that were insensitive to ethylene penetrated compacted soil more effectively than did wild-type roots. Our results indicate that soil compaction lowers gas diffusion through a reduction in air-filled pores, thereby causing ethylene to accumulate in root tissues and trigger hormone responses that restrict growth. We propose that ethylene acts as an early warning signal for roots to avoid compacted soils, which would be relevant to research into the breeding of crops resilient to soil compaction.

Wheat growth responses to soil mechanical impedance are dependent on phosphorus supply.

Increased mechanical impedance induced by soil drying or compaction causes reduction in plant growth and crop yield. However, how mechanical impedance interacts with nutrient stress has been largely unknown. Here, we investigated the effect of mechanical impedance on the growth of wheat seedlings under contrasting phosphorus (P) supply in a sand culture system which allows the mechanical impedance to be independent of water and nutrient availability. Two wheat genotypes containing the Rht-B1a (tall) or Rht-B1c (gibberellin-insensitive dwarf) alleles in the Cadenza background were used and their shoot and root traits were determined. Mechanical impedance caused a significant reduction in plant growth under sufficient P supply, including reduced shoot and root biomass, leaf area and total root length. By contrast, under low P supply, mechanical impedance did not affect biomass, tiller number, leaf length, and nodal root number in both wheat genotypes, indicating that the magnitude of the growth restriction imposed by mechanical impedance was dependent on P supply. The interaction effect between mechanical impedance and P level was significant on most plant traits except for axial and lateral root length, suggesting an evident physical and nutritional interaction. Our findings provide valuable insights into the integrated effects of plants in response to both soil physical and nutritional stresses. Understanding the response patterns is critical for optimizing soil tillage and nutrient management in the field.

The interaction between wheat roots and soil pores in structured field soil.

Wheat (Triticum aestivum L.) root growth in the subsoil is usually constrained by soil strength, although roots can use macropores to elongate to deeper layers. The quantitative relationship between the elongation of wheat roots and the soil pore system, however, is still to be determined. We studied the depth distribution of roots of six wheat varieties and explored their relationship with soil macroporosity from samples with the field structure preserved. Undisturbed soil cores (to a depth of 100 cm) were collected from the field and then non-destructively imaged using X-ray computed tomography (at a spatial resolution of 90 µm) to quantify soil macropore structure and root number density (the number of roots cm-2 within a horizontal cross-section of a soil core). Soil macroporosity changed significantly with depth but not between the different wheat lines. There was no significant difference in root number density between wheat varieties. In the subsoil, wheat roots used macropores, especially biopores (i.e. former root or earthworm channels) to grow into deeper layers. Soil macroporosity explained 59% of the variance in root number density. Our data suggested that the development of the wheat root system in the field was more affected by the soil macropore system than by genotype. On this basis, management practices which enhance the porosity of the subsoil may therefore be an effective strategy to improve deep rooting of wheat.

Sustainable Cropping Requires Adaptation to a Heterogeneous Rhizosphere.

Root-soil interactions in the rhizosphere are central to resource acquisition and crop production in agricultural systems. However, apart from studies in idealized experimental systems, rhizosphere processes in real agricultural soils in situ are largely uncharacterized. This limits the contribution of rhizosphere science to agriculture and the ongoing Green Revolution. Here, we argue that understanding plant responses to soil heterogeneity is key to understanding rhizosphere processes. We highlight rhizosphere sensing and root-induced soil modification in the context of heterogeneous soil structure, resource distribution, and root-soil interactions. A deeper understanding of the integrated and dynamic root-soil interactions in the heterogeneously structured rhizosphere could increase crop production and resource use efficiency towards sustainable agriculture.

Neighbouring plants modify maize root foraging for phosphorus: coupling nutrients and neighbours for improved nutrient-use efficiency.

Nutrient distribution and neighbours can impact plant growth, but how neighbours shape root-foraging strategy for nutrients is unclear. Here, we explore new patterns of plant foraging for nutrients as affected by neighbours to improve nutrient acquisition. Maize (Zea mays) was grown alone (maize), or with maize (maize/maize) or faba bean (Vicia faba) (maize/faba bean) as a neighbour on one side and with or without a phosphorus (P)-rich zone on the other in a rhizo-box experiment. Maize demonstrated root avoidance in maize/maize, with reduced root growth in 'shared' soil, and increased growth away from its neighbours. Conversely, maize proliferated roots in the proximity of neighbouring faba bean roots that had greater P availability in the rhizosphere (as a result of citrate and acid phosphatase exudation) compared with maize roots. Maize proliferated more roots, but spent less time to reach, and grow out of, the P patches away from neighbours in the maize/maize than in the maize/faba bean experiment. Maize shoot biomass and P uptake were greater in the heterogeneous P treatment with maize/faba bean than with maize/maize system. The foraging strategy of maize roots is an integrated function of heterogeneous distribution of nutrients and neighbouring plants, thus improving nutrient acquisition and maize growth. Understanding the foraging patterns is critical for optimizing nutrient management in crops.

Soil strength influences wheat root interactions with soil macropores.

Deep rooting is critical for access to water and nutrients found in subsoil. However, damage to soil structure and the natural increase in soil strength with depth, often impedes root penetration. Evidence suggests that roots use macropores (soil cavities greater than 75 µm) to bypass strong soil layers. If roots have to exploit structures, a key trait conferring deep rooting will be the ability to locate existing pore networks; a trait called trematotropism. In this study, artificial macropores were created in repacked soil columns at bulk densities of 1.6 g cm-3 and 1.2 g cm-3 , representing compact and loose soil. Near isogenic lines of wheat, Rht-B1a and Rht-B1c, were planted and root-macropore interactions were visualized and quantified using X-ray computed tomography. In compact soil, 68.8% of root-macropore interactions resulted in pore colonization, compared with 12.5% in loose soil. Changes in root growth trajectory following pore interaction were also quantified, with 21.0% of roots changing direction (±3°) in loose soil compared with 76.0% in compact soil. These results indicate that colonization of macropores is an important strategy of wheat roots in compacted subsoil. Management practices to reduce subsoil compaction and encourage macropore formation could offer significant advantage in helping wheat roots penetrate deeper into subsoil.

Shaping an Optimal Soil by Root-Soil Interaction.

Crop production depends on the availability of water and mineral nutrients, and increased yields might be facilitated by a greater focus on roots-soil interactions. Soil properties affecting plant growth include drought, compaction, nutrient deficiency, mineral toxicity, salinity, and submergence. Plant roots respond to the soil environment both spatially and temporally by avoiding stressful soil environments and proliferating in more favorable environments. We observe that crops can be bred for specific root architectural and biochemical traits that facilitate soil exploration and resource acquisition, enabling greater crop yields. These root traits affect soil physical and chemical properties and might be utilized to improve the soil for subsequent crops. We argue that optimizing root-soil interactions is a prerequisite for future food security.

Major Crop Species Show Differential Balance between Root Morphological and Physiological Responses to Variable Phosphorus Supply.

The relationship between root morphological and physiological responses to variable P supply in different plant species is poorly understood. We compared root morphological and physiological responses to P supply in seven crop species (Zea mays, Triticum aestivum, Brassica napus, Lupinus albus, Glycine max, Vicia faba, Cicer arietinum) treated with or without 100 mg P kg-1 in two soils (acidic and calcareous). Phosphorus deficiency decreased root length more in fibrous root species (Zea mays, Triticum aestivum, Brassica napus) than legumes. Zea mays and Triticum aestivum had higher root/shoot biomass ratio and Brassica napus had higher specific root length compared to legumes, whereas legumes (except soybean) had higher carboxylate exudation than fibrous root species. Lupinus albus exhibited the highest P-acquisition efficiency due to high exudation of carboxylates and acid phosphatases. Lupinus albus and Cicer arietinum depended mostly on root exudation (i.e., physiological response) to enhance P acquisition, whereas Zea mays, Triticum aestivum and Brassica napus had higher root morphology dependence, with Glycine max and Vicia faba in between. Principal component analysis using six morphological and six physiological responses identified root size and diameter as the most important morphological traits, whereas important physiological responses included carboxylate exudation, and P-acquisition and P-utilization efficiency followed by rhizosphere soil pH and acid phosphatase activity. In conclusion, plant species can be grouped on the basis of their response to soil P being primarily via root architectural or exudation plasticity, suggesting a potential benefit of crop-specific root-trait-based management to cope with variable soil P supply in sustainable grain production.

Increased soil phosphorus availability induced by faba bean root exudation stimulates root growth and phosphorus uptake in neighbouring maize.

Root growth is influenced by soil nutrients and neighbouring plants, but how these two drivers affect root interactions and regulate plant growth dynamics is poorly understood. Here, interactions between the roots of maize (Zea mays) and faba bean (Vicia faba) are characterized. Maize was grown alone (maize) or with maize (maize/maize) or faba bean (maize/faba bean) as competitors under five levels of phosphorus (P) supply, and with homogeneous or heterogeneous P distribution. Maize had longer root length and greater shoot biomass and P content when grown with faba bean than with maize. At each P supply rate, faba bean had a smaller root system than maize but greater exudation of citrate and acid phosphatase, suggesting a greater capacity to mobilize P in the rhizosphere. Heterogeneous P availability enhanced the root-length density of maize but not faba bean. Maize root proliferation in the P-rich patches was associated with increased shoot P uptake. Increased P availability by localized P application or by the presence of faba bean exudation stimulated root morphological plasticity and increased shoot growth in maize in the maize/faba bean mixture, suggesting that root interactions of neighbouring plants can be modified by increased P availability.

The effect of impedance to root growth on plant architecture in wheat.

BACKGROUND AND AIMS: We were interested in the effect of impedance to root growth on root and shoot architecture of wheat. It is known that Rht-1 semi-dwarfing alleles decrease the degree of leaf stunting due to root impedance. We compared commercial wheat cultivars containing different Rht-1 alleles to determine whether leaf stunting caused by root impedance differed between cultivars. We investigated effects of impedance to root growth on the angular spread of roots. METHODS: The wheat cultivars Avalon, Robigus and Battalion, carrying semi-dwarfing alleles of Rht-1, and cv. Cadenza, carrying the tall, wild-type allele, were grown under two levels of soil strength in a sand culture system designed to allow the mechanical impedance of the root growth environment to be adjusted independently of water and nutrient availability. RESULTS: Impeded roots grew more steeply than non-impeded roots: the angular spread of roots decreased from 55° to 43° from the vertical, but the genotypic effects were weak. Root impedance reduced leaf elongation and the number of tillers. Leaf area and total root length provided a common relationship across all genotype x treatment combinations. Leaf stunting in Cadenza was more severe. CONCLUSION: Our data support the hypothesis that the severity of leaf stunting due to root impedance is related to the Rht allele. Impeded roots had a smaller angular spread.

How do roots elongate in a structured soil?

In this review, we examine how roots penetrate a structured soil. We first examine the relationship between soil water status and its mechanical strength, as well as the ability of the soil to supply water to the root. We identify these as critical soil factors, because it is primarily in drying soil that mechanical constraints limit root elongation. Water supply to the root is important because root water status affects growth pressures and root stiffness. To simplify the bewildering complexity of soil-root interactions, the discussion is focused around the special cases of root elongation in soil with pores much smaller than the root diameter and the penetration of roots at interfaces within the soil. While it is often assumed that the former case is well understood, many unanswered questions remain. While low soil-root friction is often viewed as a trait conferring better penetration of strong soils, it may also increase the axial pressure on the root tip and in so doing reduce the rate of cell division and/or expansion. The precise trade-off between various root traits involved in root elongation in homogeneous soil remains to be determined. There is consensus that the most important factors determining root penetration at an interface are the angle at which the root attempts to penetrate the soil, root stiffness, and the strength of the soil to be penetrated. The effect of growth angle on root penetration implicates gravitropic responses in improved root penetration ability. Although there is no work that has explored the effect of the strength of the gravitropic responses on penetration of hard layers, we attempt to outline possible interactions. Impacts of soil drying and strength on phytohormone concentrations in roots, and consequent root-to-shoot signalling, are also considered.

A bioinformatic and transcriptomic approach to identifying positional candidate genes without fine mapping: an example using rice root-growth QTLs.

Fine mapping can accurately identify positional candidate genes for quantitative trait loci (QTLs) but can be time consuming, costly, and, for small-effect QTLs with low heritability, difficult in practice. We propose an alternative approach, which uses meta-analysis of original mapping data to produce a relatively small confidence interval for target QTLs, lists the underlying positional candidates, and then eliminates them using whole-genome transcriptomics. Finally, sequencing is conducted on the remaining candidate genes allowing identification of allelic variation in either expression or protein sequence. We demonstrate the approach using root-growth QTLs on chromosomes 2, 5, and 9 of the Bala x Azucena rice mapping population. Confidence intervals of 10.5, 9.6, and 5.4 cM containing 189, 322, and 81 genes, respectively, were produced. Transcriptomics eliminated 40% of candidate genes and identified nine expression polymorphisms. Sequencing of 30 genes revealed that 57% of the predicted proteins were polymorphic. The limitations of this approach are discussed.