Heterogeneous nutrient supply promotes maize growth and phosphorus acquisition: additive and compensatory effects of lateral roots and root hairs

Background and AimsRoot proliferation is a response to a heterogeneous nutrient distribution. However, the growth of root hairs in response to heterogeneous nutrients and the relationship between root hairs and lateral roots remain unclear. This study aims to understand the effects of heterogeneous nutrients on root hair growth and the trade-off between root hairs and lateral roots in phosphorus (P) acquisition.MethodsNear-isogenic maize lines, the B73 wild type (WT) and the rth3 root hairless mutant, were grown in rhizoboxes with uniform or localized supply of 40 (low) or 140 (high) mg P kg⁻¹ soil.ResultsBoth WT and rth3 had nearly two-fold greater shoot biomass and P content under local than uniform treatment at low P. Significant root proliferation was observed in both WT and rth3 in the nutrient patch, with the WT accompanied by an obvious increase (from 0.7 to 1.2 mm) in root hair length. The root response ratio of rth3 was greater than that of WT at low P, but could not completely compensate for the loss of root hairs. This suggests that plants enhanced P acquisition through complementarity between lateral roots and root hairs, and thus regulated nutrient foraging and shoot growth. The disappearance of WT and rth3 root response differences at high P indicated that the P application reduced the dependence of the plants on specific root traits to obtain nutrients.ConclusionsIn addition to root proliferation, the root response to a nutrient-rich patch was also accompanied by root hair elongation. The genotypes without root hairs increased their investment in lateral roots in a nutrient-rich patch to compensate for the absence of root hairs, suggesting that plants enhanced nutrient acquisition by regulating the trade-off of complementary root traits.     

Significance of root hairs for plant performance under contrasting field conditions and water deficit

Background and AimsPrevious laboratory studies have suggested selection for root hair traits in future crop breeding to improve resource use efficiency and stress tolerance. However, data on the interplay between root hairs and open-field systems, under contrasting soils and climate conditions, are limited. As such, this study aims to experimentally elucidate some of the impacts that root hairs have on plant performance on a field scale.MethodsA field experiment was set up in Scotland for two consecutive years, under contrasting climate conditions and different soil textures (i.e. clay loam vs. sandy loam). Five barley (Hordeum vulgare) genotypes exhibiting variation in root hair length and density were used in the study. Root hair length, density and rhizosheath weight were measured at several growth stages, as well as shoot biomass, plant water status, shoot phosphorus (P) accumulation and grain yield.Key ResultsMeasurements of root hair density, length and its correlation with rhizosheath weight highlighted trait robustness in the field under variable environmental conditions, although significant variations were found between soil textures as the growing season progressed. Root hairs did not confer a notable advantage to barley under optimal conditions, but under soil water deficit root hairs enhanced plant water status and stress tolerance resulting in a less negative leaf water potential and lower leaf abscisic acid concentration, while promoting shoot P accumulation. Furthermore, the presence of root hairs did not decrease yield under optimal conditions, while root hairs enhanced yield stability under drought.ConclusionsSelecting for beneficial root hair traits can enhance yield stability without diminishing yield potential, overcoming the breeder’s dilemma of trying to simultaneously enhance both productivity and resilience. Therefore, the maintenance or enhancement of root hairs can represent a key trait for breeding the next generation of crops for improved drought tolerance in relation to climate change.     

Phosphorus (P) uptake and growth of a root hairless barley mutant (bald root barley, brb) and wild type in low- and high-P soils

The recently isolated root‐hairless mutant of barley (Hordeum vulgare L), bald root barley, brb offers a unique possibility to quantify the importance of root hairs in phosphorus (P) uptake from soil. In the present study the ability of brb and the wild‐type, barley genotype Pallas producing normal root hairs to deplete P in the rhizosphere soil was investigated and the theory of diffusion and mass flow applied to compare the predicted and measured depletion profiles of diffusible P. Pallas depleted twice as much P from the rhizosphere soil as brb. The P depletion profile of Pallas uniformly extended to 0.8 mm from the root surface, which was equal to the root hair length (RHL). The model based on the theory of diffusion and mass flow explained the observed P‐depletion profile of brb, and the P depletion outside the root‐hair zone of Pallas, suggesting that the model is valid only for P movement in rhizosphere soil outside the root‐hair zone. In low‐P soil (P in soil solution 3 µm ) brb did not survive after 30 d, whereas Pallas continued to grow, confirming the importance of root hairs in plant growth in a P‐limiting environment. In high‐P soil (P in soil solution 10 µm ) both brb and Pallas maintained their growth, and they were able to produce seeds. At the high‐P concentration, RHL of the Pallas was reduced from 0.80 ± 0.2 to 0.68 ± 0.14 mm. In low‐P soil, P‐uptake rate into the roots of Pallas was 4.0 × 10^?7 g mm^?1 d^?1 and that of brb was 1.9 × 10^?7 g mm^?1 d^?1, which agreed well with the double amount of P depleted from the rhizosphere soil of Pallas in comparison with that of brb. In high‐P soil, the P uptake rates into the roots of brb and Pallas were 3.3 and 5.5 × 10^?7 g mm^?1 d^?1, respectively. The results unequivocally confirmed that in a low‐P environment, root hairs are of immense importance in P acquisition and plants survival, but under high‐P conditions they may be dispensable. The characterization of phenotypes brb and Pallas and the ability to reproduce seeds offers a unique possibility of molecular mapping of QTLs and candidate genes conferring root‐hair formation and growth of barley.     

A root hairless barley mutant for elucidating genetic of root hairs and phosphorus uptake

This paper reports a new barley mutant missing root hairs. The mutant was spontaneously discovered among the population of wild type (Pallas, a spring barley cultivar), producing normal, 0.8 mm long root hairs. We have called the mutant bald root barley (brb). Root anatomical studies confirmed the lack of root hairs on mutant roots. Amplified Fragment Length Polymorphism (AFLP) analyses of the genomes of the mutant and Pallas supported that the brb mutant has its genetic background in Pallas. The segregation ratio of selfed F₂ plants, resulting from mutant and Pallas outcross, was 1:3 (–root hairs:+root hairs), suggesting a monogenic recessive mode of inheritance.In rhizosphere studies, Pallas absorbed nearly two times more phosphorus (P) than the mutant. Most of available inorganic P in the root hair zone (0.8 mm) of Pallas was depleted, as indicated by the uniform P depletion profile near its roots. The acid phosphatase (Apase) activity near the roots of Pallas was higher and Pallas mobilised more organic P in the rhizosphere than the mutant. The higher Apase activity near Pallas roots also suggests a link between root hair formation and rhizosphere Apase activity. Hence, root hairs are important for increasing plant P uptake of inorganic as well as mobilisation of organic P in soils.Laboratory, pot and field studies showed that barley cultivars with longer root hairs (1.10 mm), extracted more P from rhizosphere soil, absorbed more P in low-P field (Olsen P=14 mg P kg^–1 soil), and produced more shoot biomass than shorter root hair cultivars (0.63 mm). Especially in low-P soil, the differences in root hair length and P uptake among the cultivars were significantly larger. Based on the results, the perspectives of genetic analysis of root hairs and their importance in P uptake and field performance of cereals are discussed.     

Arbuscular mycorrhizal colonization outcompetes root hairs in maize under low phosphorus availability

Background and AimsAn increase in root hair length and density and the development of arbuscular mycorrhiza symbiosis are two alternative strategies of most plants to increase the root–soil surface area under phosphorus (P) deficiency. Across many plant species, root hair length and mycorrhization density are inversely correlated. Root architecture, rooting density and physiology also differ between species. This study aims to understand the relationship among root hairs, arbuscular mycorrhizal fungi (AMF) colonization, plant growth, P acquisition and mycorrhizal-specific P_i transporter gene expression in maize.MethodsUsing nearly isogenic maize lines, the B73 wild type and the rth3 root hairless mutant, we quantified the effect of root hairs and AMF infection in a calcareous soil under P deficiency through a combined analysis of morphological, physiological and molecular factors.Key ResultsWild-type root hairs extended the rhizosphere for acid phosphatase activity by 0.5 mm compared with the rth3 hairless mutant, as measured by in situ zymography. Total root length of the wild type was longer than that of rth3 under P deficiency. Higher AMF colonization and mycorrhiza-induced phosphate transporter gene expression were identified in the mutant under P deficiency, but plant growth and P acquisition were similar between mutant and the wild type. The mycorrhizal dependency of maize was 33 % higher than the root hair dependency.ConclusionsThe results identified larger mycorrhizal dependency than root hair dependency under P deficiency in maize. Root hairs and AMF inoculation are two alternative ways to increase P_i acquisition under P deficiency, but these two strategies compete with each other.     

Root hair length and rhizosheath mass depend on soil porosity,strength and water content in barley genotypes

Selecting plants with improved root hair growth is a key strategy for improving phosphorus-uptake efficiency in agriculture. While significant inter- and intra-specific variation is reported for root hair length, it is not known whether these phenotypic differences are exhibited under conditions that are known to affect root hair elongation. This work investigates the effect of soil strength, soil water content (SWC) and soil particle size (SPS) on the root hair length of different root hair genotypes of barley. The root hair and rhizosheath development of five root hair genotypes of barley (Hordeum vulgare L.) was compared in soils with penetrometer resistances ranging from 0.03 to 4.45 MPa (dry bulk densities 1.2–1.7 g cm^?3). A “short” (SRH) and “long” root hair (LRH) genotype was selected to further investigate whether differentiation of these genotypes was related to SWC or SPS when grown in washed graded sand. In low-strength soil (<1.43 MPa), root hairs of the LRH genotype were on average 25 % longer than that of the SRH genotype. In high-strength soil, root hair length of the LRH genotype was shorter than that in low-strength soil and did not differ from that of the SRH genotype. Root hairs were shorter in wetter soils or soils with smaller particles, and again SRH and LRH did not differ in hair length. Longer root hairs were generally, but not always, associated with larger rhizosheaths, suggesting that mucilage adhesion was also important. The root hair growth of barley was found to be highly responsive to soil properties and this impacted on the expression of phenotypic differences in root hair length. While root hairs are an important trait for phosphorus acquisition in dense soils, the results highlight the importance of selecting multiple and potentially robust root traits to improve resource acquisition in agricultural systems.     

What are the implications of variation in root hair length on tolerance to phosphorus deficiency in combination with water stress in barley (Hordeum vulgare)?

Background and Aims

Phosphorus commonly limits crop yield and is frequently applied as fertilizer; however, supplies of quality rock phosphate for fertilizer production are diminishing. Plants have evolved many mechanisms to increase their P-fertilizer use efficiency, and an understanding of these traits could result in improved long-term sustainability of agriculture. Here a mutant population is utilized to assess the impact of root hair length on P acquisition and yield under P-deficient conditions alone or when combined with drought.

Methods

Mutants with various root hair phenotypes were grown in the glasshouse in pots filled with soil representing sufficient and deficient P treatments and, in one experiment, a range of water availability was also imposed. Plants were variously harvested at 7 d, 8 weeks and 14 weeks, and variables including root hair length, rhizosheath weight, biomass, P accumulation and yield were measured.

Key Results

The results confirmed the robustness of the root hair phenotypes in soils and their relationship to rhizosheath production. The data demonstrated that root hair length is important for shoot P accumulation and biomass, while only the presence of root hairs is critical for yield. Root hair presence was also critical for tolerance to extreme combined P deficit and drought stress, with genotypes with no root hairs suffering extreme growth retardation in comparison with those with root hairs.

Conclusions

The results suggest that although root hair length is not important for maintaining yield, the presence of root hairs is implicit to sustainable yield of barley under P-deficient conditions and when combined with extreme drought. Root hairs are a trait that should be maintained in future germplasm.     

Aluminium tolerance of root hairs underlies genotypic differences in rhizosheath size of wheat (Triticum aestivum) grown on acid soil

We found significant genetic variation in the ability of wheat (Triticum aestivum) to form rhizosheaths on acid soil and assessed whether differences in aluminium (Al(3+) ) tolerance of root hairs between genotypes was the physiological basis for this genetic variation. A method was developed to rapidly screen rhizosheath size in a range of wheat genotypes. Backcrossed populations were generated from cv Fronteira (large rhizosheath) using cv EGA-Burke (small rhizosheath) as the recurrent parent. A positive correlation existed between rhizosheath size on acid soil and root hair length. In hydroponic experiments, root hairs of the backcrossed lines with large rhizosheaths were more tolerant of Al(3+) toxicity than the backcrossed lines with small rhizosheaths. We conclude that greater Al(3+) tolerance of root hairs underlies the larger rhizosheath of wheat grown on acid soil. Tolerance of the root hairs to Al(3+) was largely independent of the TaALMT1 gene which suggests that different genes encode the Al(3+) tolerance of root hairs. The maintenance of longer root hairs in acid soils is important for the efficient uptake of water and nutrients.     

Nutrient uptake estimates for woody species as described by the NST 3.0, SSAND, and PCATS mechanistic nutrient uptake models

The effect of soil acidity on root and rhizosheath development in wheat and barley seedlings was investigated in an acid Ferrosol soil to which various amounts of lime (CaCO₃) were applied to modify soil Al concentrations (pH (CaCl₂): 4.22 to 5.35 and Al (CaCl₂ extract): 17.7 to 0.4 mg kg^?1 soil; respectively), and Ferrosol soil from an adjacent location at the same site which had a higher Al concentration (pH 4.19; 29.2 mg kg^?1 Al). The cereal lines were selected on the basis of differences in their rate of root growth, Al-resistance and root hair morphology. Root morphology was assessed after 7 days of growth. The length of fine (mainly lateral) roots of Al-sensitive genotypes was more sensitive to soil Al concentrations than that of the coarse (mainly primary) roots. The experiments demonstrated that even where root growth was protected by expression of the TaALMT1 gene for Al-resistance, root-soil contact was diminished by soil acidity because root hair length (in many lines), and root hair density and rhizosheath formation (all lines) were adversely affected by soil acidity. In the case of Al-sensitive lines, fine root growth and rhizosheath mass were reduced over much the same range of soil Al concentrations (i.e. >3–6 mg kg^?1 Al). Although Al-resistant lines could maintain fine root length under these conditions, they were similarly unable to maintain rhizosheath mass. This finding may help to explain why Al-resistant wheats which yield relatively well in deep acid soils, may also benefit from application of lime to the surface layers of the soil.     

Rhizosheath formation and involvement in foxtail millet(Setaria italica) root growth under drought stress

The rhizosheath, a layer of soil particles that adheres firmly to the root surface by a combination of root hairs and mucilage, may improve tolerance to drought stress. Setaria italica(L.) P. Beauv.(foxtail millet), a member of the Poaceae family, is an important food and fodder crop in arid regions and forms a larger rhizosheath under drought conditions. Rhizosheath formation under drought conditions has been studied, but the regulation of root hair growth and rhizosheath size in response to soil moisture remains unclear. To address this question, in this study we monitored root hair growth and rhizosheath development in response to a gradual decline in soil moisture. Here, we determined that a soil moisture level of 10%–14%(w/w)stimulated greater rhizosheath production compared to other soil moisture levels. Root hair density and length also increased at this soil moisture level, which was validated by measurement of the expression of root hair-related genes.These findings contribute to our understanding of rhizosheath formation in response to soil water stress.     

Phosphorus (P) acquisition of cereal cultivars in the field at three levels of P fertilization

Low phosphorus (P) availability in soils and diminishing P reserves emphasize the need to create plants that are more efficient P users. Knowledge of P efficient germplasm among the existing cereal varieties may serve as the basis for improving soil P use by selection and breeding. We had identified some cereal cultivars (winter wheat: Kosack and Kraka; winter barley: Hamu and Angora; spring barley: Canut, Alexis, Salka, Zita;) which differed (p<0.05) in P depletion from thin slices (0.2 mm) of the rhizosphere soil under controlled conditions. In the present study, the same cultivars were studied under field conditions at three levels of P supply (no-P, 10 and 20 kg P ha^-1) and the differences in P uptake as found in the previous work were confirmed. Under both conditions, the variation between the cultivars was greatest in soil without P fertilizers (no-P) for about 30 years. The variation in P uptake with most cultivars disappeared when 10 kg P ha^-1 was applied. Root development did not differ between the cultivars much, but there was wide, consistent variation in their root hairs, regardless of growth media (solution, soil column and field). Increase in soil P level reduced the length of root hairs. The variation in root hairs between the cultivars was largest in no-P soil. When 10 kg P ha^-1 was applied, the root hair lengths did not differ between the cultivars. Barley cultivars with longer root hairs depleted more P from the rhizosphere soil and also absorbed more P in the field. The relationship between root hairs and phosphorus uptake of the wheat cultivars was less clear. The wide variation in P uptake among the barley cultivars in the field and its relationship to the root hair development confirms that root hair length may be a suitable plant characteristic to use as criterion for selecting barley cultivars for P efficiency, especially in low-P soils. This revised version was published online in June 2006 with corrections to the Cover Date.     

Analyses of mutants of three genes that influence root hair development in Zea mays (Gramineae) suggest that root hairs are dispensable

Root hairs are specialized epidermal cells that are thought to play an important role in plant nutrition by facilitating the absorption of water and nutrients. Three maize mutants with abnormal root hair morphologies (rthl, rth2, and rth3) have been isolated from Mutator transposon stocks. All three root hair mutant phenotypes are controlled by single recessive alleles. The rthl mutant initiates normal-looking root hair primordia that fail to elongate. The normal-looking root hair primordia of the rth2 mutant elongate to only approximately one-fifth to one-fourth the length of wild type root hairs. Like rth1 primordia, rth3 primordia undergo little elongation. However, unlike the relatively normal-looking rth1 primordia, rth3 primordia are distinctly abnormal when viewed through a scanning electron microscope. The rth1 mutant exhibits pleiotropic nutrient deficiencies, while the rth2 and rth3 mutants grow vigorously. This finding suggests that under some environmental conditions, root hairs are less important to plant growth than has been previously thought. The rthl, rth2, and rth3 genes have been mapped to chromosomes 1L, 5L, and 1S, respectively, via crosses with BA translocation stocks. The rth2 allele exhibits reduced transmission through the male gametophyte, but a normal rate of transmission through female gametophytes; rth1 and rth3 are transmitted at normal rates.     

Formation and Stabilization of Rhizosheaths of Zea mays L. (Effect of Soil Water Content)

Field observations have shown that rhizosheaths of grasses formed under dry conditions are larger, more coherent, and more strongly bound to the roots than those formed in wet soils. We have quantified these effects in a model system in which corn (Zea mays L.) primary roots were grown through a 30-cm-deep prepared soil profile that consisted of a central, horizontal, "dry" (9% water content) or "wet" (20% water content) layer (4 cm thick) sandwiched between damp soil (15-17% water content). Rhizosheaths formed in dry layers were 5 times the volume of the subtending root. In wet layers, rhizosheaths were only 1.5 times the root volume. Fractions of the rhizosheath soil were removed from individual roots by three successive treatments; sonication, hot water, and abrasion. Sonication removed 50 and 90% of the soil from rhizosheaths formed in dry and wet soils, respectively. After the heat treatment, 35% of the soil still adhered to those root portions where rhizosheaths had developed in dry soil, compared with 2% where sheaths had formed in wet soil. Root hairs were 4.5 times more abundant and were more distorted on portions of roots from dry layers than from wet layers. Drier soil enhanced adhesiveness of rhizosheath mucilages and stimulated the formation of root hairs; both effects stabilize the rhizosheath. Extensive and stable rhizosheaths may function in nutrient acquisition in dry soils.     

Roothairless5, which functions in maize (Zea mays L.) root hair initiation and elongation encodes a monocot‐specific NADPH oxidase

Root hairs are instrumental for nutrient uptake in monocot cereals. The maize (Zea mays L.) roothairless5 (rth5) mutant displays defects in root hair initiation and elongation manifested by a reduced density and length of root hairs. Map‐based cloning revealed that the rth5 gene encodes a monocot‐specific NADPH oxidase. RNA‐Seq, in situ hybridization and qRT‐PCR experiments demonstrated that the rth5 gene displays preferential expression in root hairs but also accumulates to low levels in other tissues. Immunolocalization detected RTH5 proteins in the epidermis of the elongation and differentiation zone of primary roots. Because superoxide and hydrogen peroxide levels are reduced in the tips of growing rth5 mutant root hairs as compared with wild‐type, and Reactive oxygen species (ROS) is known to be involved in tip growth, we hypothesize that the RTH5 protein is responsible for establishing the high levels of ROS in the tips of growing root hairs required for elongation. Consistent with this hypothesis, a comparative RNA‐Seq analysis of 6‐day‐old rth5 versus wild‐type primary roots revealed significant over‐representation of only two gene ontology (GO) classes related to the biological functions (i.e. oxidation/reduction and carbohydrate metabolism) among 893 differentially expressed genes (FDR <5%). Within these two classes the subgroups ‘response to oxidative stress’ and ‘cellulose biosynthesis’ were most prominently represented.     

Relative importance of an arbuscular mycorrhizal fungus (Rhizophagus intraradices) and root hairs in plant drought tolerance

Both arbuscular mycorrhizal (AM) fungi and root hairs play important roles in plant uptake of water and mineral nutrients. To reveal the relative importance of mycorrhiza and root hairs in plant water relations, a bald root barley (brb) mutant and its wild type (wt) were grown with or without inoculation of the AM fungus Rhizophagus intraradices under well-watered or drought conditions, and plant physiological traits relevant to drought stress resistance were recorded. The experimental results indicated that the AM fungus could almost compensate for the absence of root hairs under drought-stressed conditions. Moreover, phosphorus (P) concentration, leaf water potential, photosynthetic rate, transpiration rate, stomatal conductance, and water use efficiency were significantly increased by R. intraradices but not by root hairs, except for shoot P concentration and photosynthetic rate under the drought condition. Root hairs even significantly decreased root P concentration under drought stresses. These results confirm that AM fungi can enhance plant drought tolerance by improvement of P uptake and plant water relations, which subsequently promote plant photosynthetic performance and growth, while root hairs presumably contribute to the improvement of plant growth and photosynthetic capacity through an increase in shoot P concentration.     

Variation in phosphorus uptake efficiency by genotypes of cowpea (Vigna unguiculata) due to differences in root and root hair length and induced rhizosphere processes

The lengths of roots and root hairs and the extent of root-induced processes affect phosphorus (P) uptake efficiency by plants. To assess the influence of variation in the lengths of roots and root hairs and rhizosphere processes on the efficiency of soil phosphorus (P) uptake, a pot experiment with a low-P soil and eight selected genotypes of cowpea (Vigna unguiculata (L) WALP) was conducted. Root length, root diameter and root hair length were measured to estimate the soil volume exploited by roots and root hairs. The total soil P was considered as a pool of Olsen-P, extractable with 0.5 M NaHCO₃ at pH 8.5, and a pool of non-Olsen-P. Model calculations were made to estimate P uptake originated from Olsen-P in the root hair zone and the Olsen-P moving by diffusion into the root hair cylinder and non-Olsen-P uptake. The mean uptake rate of P and the mean rate of non-Olsen-P depletion were also estimated. The genotypes differed significantly in lengths of roots and root hairs, and in P uptake, P uptake rates and growth. From 6 to 85% of total P uptake in the soil volume exploited by roots and root hairs was absorbed from the pool of non-Olsen-P. This indicates a considerable activity of root-induced rhizosphere processes. Hence the large differences show that traits for more P uptake-efficient plants exist in the tested cowpea genotypes. This opens the possibility to breed for more P uptake-efficient varieties as a way to bring more sparingly soluble soil P into cycling in crop production and obtain capitalisation of soil P reserves.     

Cereal root exudates contain highly structurally complex polysaccharides with soil‐binding properties

Rhizosheaths function in plant?soil interactions, and are proposed to form due to a mix of soil particle entanglement in root hairs and the action of adhesive root exudates. The soil‐binding factors released into rhizospheres to form rhizosheaths have not been characterised. Analysis of the high‐molecular‐weight (HMW) root exudates of both wheat and maize plants indicate the presence of complex, highly branched polysaccharide components with a wide range of galactosyl, glucosyl and mannosyl linkages that do not directly reflect cereal root cell wall polysaccharide structures. Periodate oxidation indicates that it is the carbohydrate components of the HMW exudates that have soil‐binding properties. The root exudates contain xyloglucan (LM25), heteroxylan (LM11/LM27) and arabinogalactan‐protein (LM2) epitopes, and sandwich‐ELISA evidence indicates that, in wheat particularly, these can be interlinked in multi‐polysaccharide complexes. Using wheat as a model, exudate‐binding monoclonal antibodies have enabled the tracking of polysaccharide release along root axes of young seedlings, and their presence at root hair surfaces and in rhizosheaths. The observations indicate that specific root exudate polysaccharides, distinct from cell wall polysaccharides, are adhesive factors secreted by root axes, and that they contribute to the formation and stabilisation of cereal rhizosheaths.     

Uncovering Genes and Ploidy Involved in the High Diversity in Root Hair Density,Length and Response to Local Scarce Phosphate in Arabidopsis thaliana

Plant root hairs increase the root surface to enhance the uptake of sparingly soluble and immobile nutrients, such as the essential nutrient phosphorus, from the soil. Here, root hair traits and the response to scarce local phosphorus concentration were studied in 166 accessions of Arabidopsis thaliana using split plates. Root hair density and length were correlated, but highly variable among accessions. Surprisingly, the well-known increase in root hair density under low phosphorus was mostly restricted to genotypes that had less and shorter root hairs under P sufficient conditions. By contrast, several accessions with dense and long root hairs even had lower hair density or shorter hairs in local scarce phosphorus. Furthermore, accessions with whole-genome duplications developed more dense but phosphorus-insensitive root hairs. The impact of genome duplication on root hair density was confirmed by comparing tetraploid accessions with their diploid ancestors. Genome-wide association mapping identified candidate genes potentially involved in root hair responses tp scarce local phosphate. Knock-out mutants in identified candidate genes (CYR1, At1g32360 and RLP48) were isolated and differences in root hair traits in the mutants were confirmed. The large diversity in root hair traits among accessions and the diverse response when local phosphorus is scarce is a rich resource for further functional analyses.     

Interactions between root hair length and arbuscular mycorrhizal colonisation in phosphorus deficient barley (Hordeum vulgare)

Aims

Phosphorus (P) limits crop yield and P-fertilisers are frequently applied to agricultural soils. However, supplies of quality rock phosphate are diminishing. Plants have evolved mechanisms to improve P-acquisition and understanding these could improve the long-term sustainability of agriculture. Here we examined interactions between root hairs and arbuscular mycorrhizal (AM) colonisation in barley (Hordeum vulgare L.).

Methods

Barley mutants exhibiting different root hair phenotypes, wild type barley and narrowleaf plantain (Plantago lanceolata L.) were grown in the glasshouse in P-sufficient and P-deficient treatments and allowed to develop AM colonization from the natural soil community. Plants were harvested after 6 weeks growth and root hair length, AM-fungal colonisation, shoot biomass and P-accumulation measured.

Results

Under P-deficient conditions, root hair length and AM colonisation were negatively related suggesting that resources are allocated to root hairs rather than to AM fungi in response to P-deficiency. There was evidence that barley and narrowleaf plantain employed different strategies to increase P-acquisition under identical conditions, but root hairs were more effective.

Conclusions

This research suggests future barley breeding programmes should focus on maintaining or improving root hair phenotypes and that pursuing enhancements to AM associations under the prevalent agricultural conditions tested here would be ineffectual.     

Root hair phenotypes influence nitrogen acquisition in maize

Background and AimsThe utility of root hairs for nitrogen (N) acquisition is poorly understood.MethodsWe explored the utility of root hairs for N acquisition in the functional–structural model SimRoot and with maize genotypes with variable root hair length (RHL) in greenhouse and field environments.Key ResultsSimulation results indicate that long, dense root hairs can improve N acquisition under varying N availability. In the greenhouse, ammonium availability had no effect on RHL and low nitrate availability increased RHL, while in the field low N reduced RHL. Longer RHL was associated with 216 % increase in biomass and 237 % increase in plant N content under low-N conditions in the greenhouse and a 250 % increase in biomass and 200 % increase in plant N content in the field compared with short-RHL phenotypes. In a low-N field environment, genotypes with long RHL had 267 % greater yield than those with short RHL. We speculate that long root hairs improve N capture by increased root surface area and expanded soil exploration beyond the N depletion zone surrounding the root surface.ConclusionsWe conclude that root hairs play an important role in N acquisition. We suggest that root hairs merit consideration as a breeding target for improved N acquisition in maize and other crops.