Root hairs are the most important root trait for rhizosheath formation of barley (Hordeum vulgare), maize (Zea mays) and Lotus japonicus (Gifu)

Background and AimsRhizosheaths are defined as the soil adhering to the root system after it is extracted from the ground. Root hairs and mucilage (root exudates) are key root traits involved in rhizosheath formation, but to better understand the mechanisms involved their relative contributions should be distinguished.MethodsThe ability of three species [barley (Hordeum vulgare), maize (Zea mays) and Lotus japonicus (Gifu)] to form a rhizosheath in a sandy loam soil was compared with that of their root-hairless mutants [bald root barley (brb), maize root hairless 3 (rth3) and root hairless 1 (Ljrhl1)]. Root hair traits (length and density) of wild-type (WT) barley and maize were compared along with exudate adhesiveness of both barley and maize genotypes. Furthermore, root hair traits and exudate adhesiveness from different root types (axile versus lateral) were compared within the cereal species.Key ResultsPer unit root length, rhizosheath size diminished in the order of barley > L. japonicus > maize in WT plants. Root hairs significantly increased rhizosheath formation of all species (3.9-, 3.2- and 1.8-fold for barley, L. japonicus and maize, respectively) but there was no consistent genotypic effect on exudate adhesiveness in the cereals. While brb exudates were more and rth3 exudates were less adhesive than their respective WTs, maize rth3 bound more soil than barley brb. Although both maize genotypes produced significantly more adhesive exudate than the barley genotypes, root hair development of WT barley was more extensive than that of WT maize. Thus, the greater density of longer root hairs in WT barley bound more soil than WT maize. Root type did not seem to affect rhizosheath formation, unless these types differed in root length.ConclusionsWhen root hairs were present, greater root hair development better facilitated rhizosheath formation than root exudate adhesiveness. However, when root hairs were absent root exudate adhesiveness was a more dominant trait.     

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.     

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.     

Heterogeneity and Glycan Masking of Cell Wall Microstructures in the Stems of Miscanthus x giganteus,and Its Parents M. sinensis and M. sacchariflorus

Plant cell walls, being repositories of fixed carbon, are important sources of biomass and renewable energy. Miscanthus species are fast growing grasses with a high biomass yield and they have been identified as potential bioenergy crops. Miscanthus x giganteus is the sterile hybrid between M. sinensis and M. sacchariflorus, with a faster and taller growth than its parents. In this study, the occurrence of cell wall polysaccharides in stems of Miscanthus species has been determined using fluorescence imaging with sets of cell wall directed monoclonal antibodies. Heteroxylan and mixed linkage-glucan (MLG) epitopes are abundant in stem cell walls of Miscanthus species, but their distributions are different in relation to the interfascicular parenchyma and these epitopes also display different developmental dynamics. Detection of pectic homogalacturonan (HG) epitopes was often restricted to intercellular spaces of parenchyma regions and, notably, the high methyl ester LM20 HG epitope was specifically abundant in the pith parenchyma cell walls of M. x giganteus. Some cell wall probes cannot access their target glycan epitopes because of masking by other polysaccharides. In the case of Miscanthus stems, masking of xyloglucan by heteroxylan and masking of pectic galactan by heteroxylan and MLG was detected in certain cell wall regions. Knowledge of tissue level heterogeneity of polysaccharide distributions and molecular architectures in Miscanthus cell wall structures will be important for both understanding growth mechanisms and also for the development of potential strategies for the efficient deconstruction of Miscanthus biomass.     

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.     

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.     

Evaluation of a novel tool for sampling root exudates from soil-grown plants compared to conventional techniques

The release of low molecular weight (LMW) organic compounds (e.g. organic acids, amino acids, sugars, etc.) by living plant roots significantly contributes to the development of chemical, physical as well as microbial rhizosphere gradients. Suitable and accurate sampling procedures are crucial for enhancing our understanding of the dynamics of related rhizosphere processes. Here we compare common sampling techniques with a novel tool for root exudate collection that allows non-destructive and repetitive sampling from soil-grown roots. Root exudates from Zea mays L. were collected using the following techniques: (i) hydroponic growth and sampling, (ii) soil growth and hydroponic sampling and (iii) rhizoboxes fitted with a novel in situ root exudate collecting tool. Furthermore, rhizosphere soil solution for the analysis of exudates and microbial metabolites was sampled using micro-suction cups (iv). The effect of different sampling solutions (deionised water and 0.5 mM CaCl₂) on organic acid and amino acid exudation patterns was also investigated. The novel exudate collecting tool was successfully tested for root exudate sampling. Results showed that particularly amino acid exudation rates were significantly affected by growth conditions and sampling procedures, while organic acid exudation patterns varied less across the different sampling setups. Despite qualitative and quantitative differences, exudation rates were in the same order of magnitude across the different sampling procedures. Soil solution concentrations obtained from micro-suction-cup sampling at defined distance to the root surface showed no distinct gradient, highlighting the importance of soil microorganisms in regulating the soil solution concentration of LMW C compounds either via microbial degradation or the release of microbial metabolites. The exudate collector offers new opportunities to assess root exudation rates and composition from soil-grown plants and thus enhances our knowledge of fundamental rhizosphere processes.     

Manipulating exudate composition from root apices shapes the microbiome throughout the root system

Certain soil microorganisms can improve plant growth, and practices that encourage their proliferation around the roots can boost production and reduce reliance on agrochemicals. The beneficial effects of the microbial inoculants currently used in agriculture are inconsistent or short-lived because their persistence in soil and on roots is often poor. A complementary approach could use root exudates to recruit beneficial microbes directly from the soil and encourage inoculant proliferation. However, it is unclear whether the release of common organic metabolites can alter the root microbiome in a consistent manner and if so, how those changes vary throughout the whole root system. In this study, we altered the expression of transporters from the ALUMINUM-ACTIVATED MALATE TRANSPORTER and the MULTIDRUG AND TOXIC COMPOUND EXTRUSION families in rice (Oryza sativa L.) and wheat (Triticum aestivum L.) and tested how the subsequent release of their substrates (simple organic anions, including malate, citrate, and γ-amino butyric acid) from root apices affected the root microbiomes. We demonstrate that these exudate compounds, separately and in combination, significantly altered microbiome composition throughout the root system. However, the root type (seminal or nodal), position along the roots (apex or base), and soil type had a greater influence on microbiome structure than the exudates. These results reveal that the root microbiomes of important cereal species can be manipulated by altering the composition of root exudates, and support ongoing attempts to improve plant production by manipulating the root microbiome.

One-sentence summary: The root microbiome of rice and wheat can be manipulated by altering the activity of root transporters and exudates.     

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.     

Microorganisms and nematodes increase levels of secondary metabolites in roots and root exudates of Plantago lanceolata

Plant secondary metabolites play an important role in constitutive and inducible direct defense of plants against their natural enemies. While induction of defense by aboveground pathogens and herbivores is well-studied, induction by belowground organisms is less explored. Here, we examine whether soil microorganisms and nematodes can induce changes in levels of the secondary metabolites aucubin and catalpol (iridoid glycosides, IG) in roots and root exudates of two full-sib families of Plantago lanceolata originating from lines selected for low and high constitutive levels of IG in leaves. Addition of soil microorganisms enhanced the shoot and root biomass, and the concentration of aucubin in roots of both Plantago lines without affecting IG levels in the rhizosphere. By contrast, nematode addition tended to reduce the root biomass and enhanced the stalk biomass, and increased the levels of aucubin and catalpol in root exudates of both Plantago lines, without affecting root IG concentrations. The Plantago lines did not differ in constitutive levels of aucubin and total IG in roots, while the concentration of catalpol was slightly higher in roots of plants originally selected for low constitutive levels of IG in leaves. Root exudates of “high IG line” plants contained significantly higher levels of aucubin, which might be explained by their higher root biomass. We conclude that soil microorganisms can induce an increase of aucubin concentrations in the roots, whereas nematodes (probably plant feeders) lead to an enhancement of aucubin and catalpol levels in root exudates of P. lanceolata. A potential involvement of secondary metabolites in belowground interactions between plants and soil organisms is discussed.     

Increasing flavonoid concentrations in root exudates enhance associations between arbuscular mycorrhizal fungi and an invasive plant

Many invasive plants have enhanced mutualistic arbuscular mycorrhizal (AM) fungal associations, however, mechanisms underlying differences in AM fungal associations between introduced and native populations of invasive plants have not been explored. Here we test the hypothesis that variation in root exudate chemicals in invasive populations affects AM fungal colonization and then impacts plant performance. We examined flavonoids (quercetin and quercitrin) in root exudates of native and introduced populations of the invasive plant Triadica sebifera and tested their effects on AM fungi and plant performance. We found that plants from introduced populations had higher concentrations of quercetin in root exudates, greater AM fungal colonization and higher biomass. Applying root exudates more strongly increased AM fungal colonization of target plants and AM fungal spore germination when exudate donors were from introduced populations. The role of root exudate chemicals was further confirmed by decreased AM fungal colonization when activated charcoal was added into soil. Moreover, addition of quercetin into soil increased AM fungal colonization, indicating quercetin might be a key chemical signal stimulating AM fungal associations. Together these results suggest genetic differences in root exudate flavonoids play an important role in enhancing AM fungal associations and invasive plants’ performance, thus considering root exudate chemicals is critical to unveiling mechanisms governing shifting plant-soil microbe interactions during plant invasions.Subject terms: Population dynamics, Community ecology, Plant ecology     

Mycorrhiza and root hairs in barley enhance acquisition of phosphorus and uranium from phosphate rock but mycorrhiza decreases root to shoot uranium transfer

Some phosphate rocks (PR) contain high concentrations of uranium (U), which are potentially toxic via accumulation in soils and food chains, and plant uptake of U is likely to be influenced by characteristics of roots and associated microorganisms. The relative importance of root hairs and mycorrhiza in U uptake from PR was studied using a root hairless barley (Hordeum vulgare) mutant (Brb) and its wild type (WT). Both plant genotypes were grown in pots with Glomus intraradices BEG 87, or in the absence of mycorrhiza, and three P treatments were included: nil P, 2% (w/w) PR and 50 mg KH(2)PO(4)-P kg(-1) soil. Mycorrhiza markedly increased d. wts and P contents of Brb amended with nil P or PR, but generally depressed d. wts of WT plants, irrespective of P amendments. Mycorrhiza had contrasting effects on U contents in roots and shoots, in particular in Brb where mycorrhiza increased root U concentrations but decreased U translocation from roots to shoots. The experiment supports our understanding of arbuscular mycorrhiza as being multifunctional by not only improving the utilization of PR by the host plant but also by contributing to the phytostabilization of uranium.     

Comparative glycan profiling of Ceratopteris richardii ‘C-Fern’ gametophytes and sporophytes links cell-wall composition to functional specialization

Background and Aims

Innovations in vegetative and reproductive characters were key factors in the evolutionary history of land plants and most of these transformations, including dramatic changes in life cycle structure and strategy, necessarily involved cell-wall modifications. To provide more insight into the role of cell walls in effecting changes in plant structure and function, and in particular their role in the generation of vascularization, an antibody-based approach was implemented to compare the presence and distribution of cell-wall glycan epitopes between (free-living) gametophytes and sporophytes of Ceratopteris richardii ‘C-Fern’, a widely used model system for ferns.

Methods

Microarrays of sequential diamino-cyclohexane-tetraacetic acid (CDTA) and NaOH extractions of gametophytes, spores and different organs of ‘C-Fern’ sporophytes were probed with glycan-directed monoclonal antibodies. The same probes were employed to investigate the tissue- and cell-specific distribution of glycan epitopes.

Key Results

While monoclonal antibodies against pectic homogalacturonan, mannan and xyloglucan widely labelled gametophytic and sporophytic tissues, xylans were only detected in secondary cell walls of the sporophyte. The LM5 pectic galactan epitope was restricted to sporophytic phloem tissue. Rhizoids and root hairs showed similarities in arabinogalactan protein (AGP) and xyloglucan epitope distribution patterns.

Conclusions

The differences and similarities in glycan cell-wall composition between ‘C-Fern’ gametophytes and sporophytes indicate that the molecular design of cell walls reflects functional specialization rather than genetic origin. Glycan epitopes that were not detected in gametophytes were associated with cell walls of specialized tissues in the sporophyte.     

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.     

Turnover and distribution of root exudates of Zea mays

Decomposition and distribution of root exudates of Zea mays L. were studied by means of ¹⁴CO₂ pulse labeling of shoots on a loamy Haplic Luvisol. Plants were grown in two-compartment pots, where the lower part was separated from the roots by monofilament gauze. Root hairs, but not roots, penetrated through the gauze into the lower part of the soil. The root-free soil in the lower compartment was either sterilized with cycloheximide and streptomycin or remained non-sterile. In order to investigate exudate distribution, 3 days after the ¹⁴C labeling, the lower soil part was frozen and sliced into 15, one-mm thick layers using a microtome. Cumulative ¹⁴CO₂ efflux from the soil during the first 3 days after ¹⁴C pulse labeling did not change during plant growth and amounted to about 13–20% of the total recovered ¹⁴C (41–55% of the carbon translocated below ground). Nighttime rate of total CO₂ efflux was 1.5 times lower than during daytime because of tight coupling of exudation with photosynthesis intensity. The average CO₂ efflux from the soil with Zea mays was about 74 g C g^–1 day^–1 (22 g C m^–2 day^–1), although, the contribution of plant roots to the total CO₂ efflux from the soil was about 78%, and only 22% was respired from the soil organic matter. Zea mays transferred about 4 g m^–2 of carbon under ground during 26 days of growth. Three zones of exudate concentrations were identified from the distribution of the ¹⁴C-activity in rhizosphere profiles after two labeling periods: (1) 1–2 (3) mm (maximal concentration of exudates) 2) 3–5 mm (presence of exudates is caused by their diffusion from the zone 1); (3) 6–10 mm (very insignificant amounts of exudates diffused from the previous zones). At the distance further than 10 mm no exudates were found. The calculated coefficient of exudate diffusion in the soil was 1.9 × 10^–7 cm² s^–1.     

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.     

Effect of larch (Larix gmelini Rupr.) root exudates on Manchurian walnut (Juglans mandshurica Maxim.) growth and soil juglone in a mixed-species plantation

The establishment and productivity of a Manchurian walnut (Juglans mandshurica Maxim.) plantation can be improved by inter-planting with larch (Larix gmelini Rupr.) in Northeast China, but the potential mechanism remains obscure. We carried out a series of experiments in a 20-year-old mixed-species plantation, as well as in Manchurian walnut and larch plantations. Manchurian walnut seedlings had difficulty surviving in the Manchurian walnut plantation because their growth was inhibited by their own soil and root exudates. In sharp contrast, Manchurian walnut seedlings grew well in larch and mixed-species plantations. Larch soil and root exudates greatly stimulated the growth of Manchurian walnut seedlings in controlled conditions. In particular, larch root exudates can increase the soil microbial populations, including bacteria, actinomycetes, azotobacter and cellulose-decomposing microorganisms; larch root exudates can also increase the enzyme activities of saccharase, urease, proteinase and polyphenol oxidase. Significant results led to a rapid degradation of the root-exuded phytotoxic juglone from Manchurian walnut. Manchurian walnut root exudates contained juglone at a high concentration of 121.3?±?6.6 mg g^?1, while juglone concentrations in the soil beneath Manchurian walnut trees ranged from 2.9–6.2 µg g^?1 soil. It appears from the results that juglone may be released from Manchurian walnut roots into the soil in a sufficient quantity but rapidly degrades due to interactions with soil factors. Furthermore, juglone was more resistant toward degradation in the Manchurian walnut soil (t _1/2 ?=?7.36?±?0.63 h) when compared to the larch soil (t _1/2 ?=?4.66?±?0.82 h). The results suggest that larch may improve the establishment and productivity of Manchurian walnut in a mixed-species plantation through the release of root exudates.     

Drought-induced changes in soil contact and hydraulic conductivity for roots of Opuntia ficus-indica with and without rhizosheaths

Water movement between roots and soil can be limited by incomplete root–soil contact, such as that caused by air gaps due to root shrinkage, and can also be influenced by rhizosheaths, composed of soil particles bound together by root exudates and root hairs. The possible occurrence of air gaps between the roots and the soil and their consequences for the hydraulic conductivity of the root–soil pathway were therefore investigated for the cactus t Opuntia ficus-indica, which has two distinct root regions: a younger, distal region where rhizosheaths occur, and an older, proximal region where roots are bare. Resin-embedded sections of roots in soil were examined microscopically to determine root–soil contact for container-grown plants kept moist for 21 days, kept moist and vibrated to eliminate air gaps, droughted for 21 days, or droughted and vibrated. During drought, roots shrank radially by 30% and root–soil contact in the bare root region of nonvibrated containers was reduced from 81% to 31%. For the sheathed region, the hydraulic conductivity of the rhizosheath was the least limiting factor and the root hydraulic conductivity was the most limiting; for the bare root region, the hydraulic conductivity of the soil was the least limiting factor and the hydraulic conductivity of the root–soil air gap was the most limiting. The rhizosheath, by virtually eliminating root–soil air gaps, facilitated water uptake in moist soil. In the bare root region, the extremely low hydraulic conductivity of the root–soil air gap during drought helped limit water loss from roots to a drier soil.     

Lead complexation behaviour of root exudates of salt marsh plant Salicornia europaea L

Abstract

Root exudates are considered to have an important role in mobility and bioavailability of heavy metals. High molecular weight (HMW) substances are the main components of root exudates, however, knowledge about their interactions with heavy metals is lacking. In the present study, Pb(II) complexation of the HMW fluorescent fractions in root exudates from Salicornia europaea L. was investigated using excitation emission matrix (EEM) fluorescence spectroscopy. Two protein-like fluorescence peaks were identified in the EEM spectrum of root exudates. The fluorescence of both peaks was clearly quenched by Pb(II). The values of conditional stability constants, log K_a, for these two protein-like fluorescence peaks were 4.14 and 3.79. This indicates that the fluorescent substances are strong Pb(II) complexing organic ligands.