Rhizosphere carboxylate concentrations of chickpea are affected by genotype and soil type

We investigated whether concentrations of carboxylates in the rhizosphere of chickpea (Cicer arietinum L.) roots were related to soil phosphorus levels. In a field experiment, cultivar Sona was grown at two P levels on eight soil types at three locations. There were large differences in extractable (0.2 mM CaCl₂) rhizosphere carboxylate concentrations amongst the locations. The effect of P fertiliser was variable and carboxylate concentrations depended on soil type. To examine the effect of soil P in more detail, a glasshouse experiment was carried out, in which three cultivars (Heera, Sona and Tyson) were grown at four P levels on one soil type. The biomass of chickpea plants increased with increasing P level of the soil, and the root mass ratio decreased at the highest soil P level. However, rhizosphere concentrations of the carboxylates malonate, malate and citrate did not differ significantly between P treatments. This implied that there was no simple relation between available P and root exudation rates, in contrast to earlier results in studies using hydroponics. Cultivars differed in carboxylate concentration pattern: Sona and Tyson showed a tendency towards increased rhizosphere carboxylate concentrations at the second harvest, whereas the carboxylate concentration of Heera tended to decrease. It is hypothesised that chickpea roots always exude a basal level of carboxylates into the rhizosphere. They only increase carboxylate exudation considerably when the P availability is extremely low, which may occur in soils that strongly bind P.     

Root Cluster Formation and Citrate Exudation of White Lupin （Lupinus albus L.） as Related to Phosphorus Availability

A split-root system was used to investigate whether the external or internal P concentration controls root cluster formation and citrate exudation in white lupin (Lupinus albus L.) grown under controlled conditions. In spite of low P concentrations in the shoots and roots of the -P plant, its dry weight was not reduced compared with the P plant. Supplying external P (0.25 mmol/L) to one root half resulted in an increase in P concentration not only in the shoot, but also in the P-deprived root half, indicating P cycling within the plants. Omitting P from both split-root pots stimulated root cluster formation in both root halves,whereas P supply to one root half stimulated root cluster formation at the beginning of the treatment. Neither P supply to just one root half continuously nor resupply of P to one root half after 19 d of P starvation inhibited root cluster formation on the P-deprived side, although the concentration of P in this root half and shoot increased markedly. The results indicate that root cluster formation in L. albus is controlled by both shoot and root P concentrations. The rates of citrate exudation by both root halves with P deficiency were higher than those of the one root half supplied with P only. In the treatment with one root half supplied with P, the rates of citrate exudation by either the P-supplied or -deprived root halves were almost the same,regardless of P concentration in the roots. The results suggest that internal P concentration controls root cluster formation and citrate exudation in white lupin, but these processes may be regulated by different mechanisms.     

Root Cluster Formation and Citrate Exudation of White Lupin (Lupinus albus L.) as Related to Phosphorus Availability

A split-root system was used to investigate whether the external or internal P concentration controls root cluster formation and citrate exudation in white lupin (Lupinus albus L.) grown under controlled conditions. In spite of low P concentrations in the shoots and roots of the -P plant, its dry weight was not reduced compared with the P plant. Supplying external P (0.25 mmol/L) to one root halfresulted in an increase in P concentration not only in the shoot, but also in the P-deprived root half, indicating P cycling within the plants. Omitting P from both split-root pots stimulated root cluster formation in both root halves,whereas P supply to one root halfstimulated root cluster formation at the beginning of the treatment. Neither P supply to just one root half continuously nor resupply of P to one root half after 19 d of P starvation inhibited root cluster formation on the P-deprived side, although the concentration of P in this root half and shoot increased markedly. The results indicate that root cluster formation in L. albus is controlled by both shoot and root P concentrations. The rates of citrate exudation by both root halves with P deficiency were higher than those of the one root half supplied with P only. In the treatment with one root half supplied with P, the rates of citrate exudation by either the P-supplied or -deprived root halves were almost the same,regardless of P concentration in the roots. The results suggest that internal P concentration controls root cluster formation and citrate exudation in white lupin, but these processes may be regulated by different mechanisms.     

Chickpea and white lupin rhizosphere carboxylates vary with soil properties and enhance phosphorus uptake

Chickpea and white lupin roots are able to exude large amounts of carboxylates, but the resulting concentrations in the rhizosphere vary widely. We grew chickpea in pots in eleven different Western Australian soils, all with low phosphorus concentrations. While final plant mass varied more than two-fold and phosphorus content almost five-fold, there were only minor changes in root morphological traits that potentially enhance phosphorus uptake (e.g., the proportion of plant mass allocated to roots, or the length of roots per unit root mass). In contrast, the concentration of carboxylates (mainly malonate, citrate and malate, extracted using a 0.2 mM CaCl₂ solution) varied ten-fold (averaging 2.3 mol g^–1 dry rhizosphere soil, approximately equivalent to a soil solution concentration of 23 mM). Plant phosphorus uptake was positively correlated with the concentration of carboxylates in the rhizosphere, and it was consistently higher in soils with a smaller capacity to sorb phosphorus. Phosphorus content was not correlated with bicarbonate-extractable phosphorus or any other single soil trait. These results suggest that exuded carboxylates increased the availability of phosphorus to the plant, however, the factors that affected root exudation rates are not known. When grown in the same six soils, three commonly used Western Australian chickpea cultivars had very similar rhizosphere carboxylate concentrations (extracted using a 0.2 mM CaCl₂ solution), suggesting that there is little genetic variation for this trait in chickpea. Variation in the concentration of carboxylates in the rhizosphere of white lupin did not parallel that of chickpea across the six soils. However, in both species the proportion of citrate decreased and that of malate increased at lower soil pH. We conclude that patterns of variation in root exudates need to be understood to optimise the use of this trait in enhancing crop phosphorus uptake.     

Morphological plasticity of wheat and barley roots in response to spatial variation in soil strength

Root systems of individual crop plants may encounter large variations in mechanical impedance to root penetration. Split-root experiments were conducted to compare the effects of spatial variation in soil strength on the morphological plasticity of wheat and barley roots, and its relationship to shoot growth. Plants of spring barley (Hordeum vulgare cv Prisma) and spring wheat (Triticum aestivum cv Alexandria) were grown for 12 days with their seminal roots divided between two halves of a cylinder packed with sandy loam soil. Three treatment combinations were imposed: loose soil where both halves of the cylinder were packed to 1.1 g cm^–3 (penetrometer resistance 0.3 MPa), dense soil where both halves were packed to 1.4 g cm^–3 (penetrometer resistance 1 MPa), and a split-root treatment where one half was packed to 1.1 and the other to 1.4 g cm^–3. In barley, uniform high soil strength restricted the extension of main seminal root axes more than laterals. In the split-root treatment, the length of laterals and the dry weight of main axes and laterals were increased in the loose soil half and reduced in the dense soil half compared with their respective loose and dense-soil controls. No such compensatory adjustments between main axis and laterals and between individual seminal roots were found in wheat. Variation in soil strength had no effect on the density of lateral roots (number per unit main axis length) in either barley or wheat. The nature and extent of wheat root plasticity in response to variation in soil strength was very different from that in response to changes in N-supply in previous experiments. In spite of the compensatory adjustments in growth between individual seminal roots of barley, the growth of barley shoots, as in wheat, was reduced when part of the root system was in compacted soil.     

Impact of phosphorus mineral source (Al–P or Fe–P) and pH on cluster-root formation and carboxylate exudation in Lupinus albus L.

Lupinus albus L. were grown in rhizoboxes containing a soil amended with sparingly available Fe–P or Al–P (100 μg P g⁻¹ soil/resin mixture). Root halves of individual plants were supplied with nutrient solution (minus P) buffered at either pH 5.5 or 7.5, to assess whether the source of mineral-bound P and/or pH influence cluster-root growth and carboxylate exudation. The P-amended soil was mixed 3:1 (w/w) with anion-exchange resins to allow rapid fixation of carboxylates. Treatments lasted 10 weeks. Forty percent and 30% of the root mass developed as cluster roots in plants grown on Fe–P and Al–P respectively, but cluster-root growth was the same on root-halves grown at pH 5.5 or 7.5. Mineral-bound P source (Al– or Fe–P) had no influence on the types of carboxylates measured in soil associated with cluster roots—citrate (and trace amounts of malate and fumarate) was the only major carboxylate detected. The [citrate] in the rhizosphere of cluster roots decreased with increased shoot P status (suggesting a systemic effect) and also, only for plants grown on Al–P, with decreased pH in the root environment (suggesting a local effect). In a separate experiment using anion exchange resins pre-loaded with malate or citrate, we measured malate (50%) and citrate (79%) recovery after 30 days in soil. We therefore, also conclude that measurements of [citrate] and [malate] at the root surface may be underestimated and would be greater than the 40- and 1.6-μmol g⁻¹ root DM, respectively estimated by us and others because of decomposition of carboxylates around roots prior to sampling.     

Root excretion of carboxylic acids and protons in phosphorus-deficient plants

Phosphorus deficiency-induced metabolic changes related to exudation of carboxylic acids and protons were compared in roots of wheat (Triticum aestivum L. cv Haro), tomato (Lycopersicon esculentum L., cv. Moneymaker), chickpea (Cicer arietinum) and white lupin (Lupinus albus L. cv. Amiga), grown in a hydroponic culture system. P deficiency strongly increased the net release of protons from roots of tomato, chickpea and white lupin, but only small effects were observed in wheat. Release of protons coincided with increased exudation of carboxylic acids in roots of chickpea and white lupin, but not in those of tomato and wheat. P deficiency-induced exudation of carboxylic acids in chickpea and white lupin was associated with a larger increase of carboxylic acid concentrations in the roots and lower accumulation of carboxylates in the shoot tissue compared to that in wheat and tomato. - Citric acid was one of the major organic acids accumulated in the roots of all investigated species in response to P deficiency, and this was associated with increased activity and enzyme protein levels of PEP carboxylase, which is required for biosynthesis of citrate. Accumulation of citric acid was most pronounced in the roots of P-deficient white lupin, chickpea and tomato. Increased PEP carboxylase activity in the roots of these plants coincided with decreased activity of aconitase, which is involved in the breakdown of citric acid in the TCA cycle. In the roots of P-deficient wheat plants, however, the activities of both PEP carboxylase and aconitase were enhanced, which was associated with little accumulation of citric acid. The results suggest that P deficiency-induced exudation of carboxylic acids depends on the ability to accumulate carboxylic acids in the root tissue, which in turn is determined by biosynthesis, degradation and partitioning of carboxylic acids or related precursors between roots and shoot. In some plant species such as white lupin, there are indications for a specific transport mechanism (anion channel), involved in root exudation of extraordinary high amounts of citric acid. This revised version was published online in June 2006 with corrections to the Cover Date.     

Convergence of a specialized root trait in plants from nutrient-impoverished soils: phosphorus-acquisition strategy in a nonmycorrhizal cactus

In old, phosphorus (P)-impoverished habitats, root specializations such as cluster roots efficiently mobilize and acquire P by releasing large amounts of carboxylates in the rhizosphere. These specialized roots are rarely mycorrhizal. We investigated whether Discocactus placentiformis (Cactaceae), a common species in nutrient-poor campos rupestres over white sands, operates in the same way as other root specializations. Discocactus placentiformis showed no mycorrhizal colonization, but exhibited a sand-binding root specialization with rhizosheath formation. We first provide circumstantial evidence for carboxylate exudation in field material, based on its very high shoot manganese (Mn) concentrations, and then firm evidence, based on exudate analysis. We identified predominantly oxalic acid, but also malic, citric, lactic, succinic, fumaric, and malonic acids. When grown in nutrient solution with P concentrations ranging from 0 to 100 μM, we observed an increase in total carboxylate exudation with decreasing P supply, showing that P deficiency stimulated carboxylate release. Additionally, we tested P solubilization by citric, malic and oxalic acids, and found that they solubilized P from the strongly P-sorbing soil in its native habitat, when the acids were added in combination and in relatively low concentrations. We conclude that the sand-binding root specialization in this nonmycorrhizal cactus functions similar to that of cluster roots, which efficiently enhance P acquisition in other habitats with very low P availability.     

The effect of manganese supply on exudation of carboxylates by roots of lucerne (Medicago sativa)

Some low-molecular-weight carboxylates commonly found in plant root exudates have the potential to increase the availability of Mn in the rhizosphere. Release of various compounds into the rhizosphere by plant roots may also be a mechanism by which certain species and genotypes are able to tolerate conditions of low Mn availability better than others. Lucerne (Medicago sativa L.) plants of Salado, a genotype tolerant to Mn deficiency, and Sirosal, an intolerant genotype, were grown in solution culture with 0, 5 or 500 nM Mn (Mn-0, Mn-5 and Mn-500). Exudates of whole root systems were collected at 14, 24 and 36 d and analysed by HPLC. Oxalate, tartarate, L-malate, lactate, malonate, maleate, citrate and succinate were detected and quantified in exudates under all Mn treatments. Malonate, citrate and succinate accounted for the majority of carboxylates in the exudates. Exudation increased with plant age, but amounts of individual carboxylates remained constant in proportion to the total amount exuded. A significant increase in exudation of all carboxylates other than malonate and maleate resulted from omission of Mn from nutrient solutions. Salado exuded more oxalate, tartarate, L-malate, lactate, citrate and succinate than Sirosal at Mn-0, and more citrate and succinate than Sirosal at Mn-5. Genotypic differences in carboxylate exudation under Mn-0 were associated with production of roots with diameter <100 μm. Plant Mn concentrations and growth rates suggested carboxylate exudation differences were not the sole factor responsible for differential tolerance to Mn deficiency in the lucerne genotypes.     

Shoot P status regulates cluster-root growth and citrate exudation in Lupinus albus grown with a divided root system

The present study was carried out to investigate whether the P concentration in the roots or the shoots controls the growth and citrate exudation of cluster roots in white lupin (Lupinus albus L). Foliar P application indicated that low P concentration in the shoots enhanced cluster‐root growth and citrate‐exudation rate more so than low P concentration in the roots. In the split‐root study, the P concentration in the shoots increased with increased P supply (1, 25 or 75 mmol m^?3 P), to the ‘privileged’ root halves. Roots ‘deprived’ of P invariably had the same low P concentrations, whereas those in the ‘privileged’ roots increased with increasing P supply (1, 25 or 75 mmol m^?3 P). Nevertheless, the proportion of the total root mass allocated to cluster roots, and the citrate‐exudation rates from the root halves were always similar on both root halves, irrespective of P supply, and decreased with increasing shoot P concentrations. Peak citrate exudation rates from developing cluster roots were significantly faster from cluster roots on the ‘deprived’ root halves when the ‘privileged’ half was exposed to 1 mmol m^?3 P as compared with 25 or 75 mmol m^?3 P. The possibility that changes in the concentrations of P fractions in the root halves influenced cluster‐root growth and citrate exudation was discounted, because there were no significant differences in insoluble organic P, ester‐P and inorganic P among all ‘deprived’ root halves. The results indicate that cluster‐root proportions and citrate exudation rates were regulated systemically by the P status of the shoot, and that P concentrations in the roots had little influence on growth and citrate exudation of cluster roots in L. albus.     

Root cap influences root colonisation by Pseudomonas fluorescens SBW25 on maize

We investigated the influence of root border cells on the colonisation of seedling Zea mays roots by Pseudomonas fluorescens SBW25 in sandy loam soil packed at two dry bulk densities. Numbers of colony forming units (CFU) were counted on sequential sections of root for intact and decapped inoculated roots grown in loose (1.0 mg m(-3)) and compacted (1.3 mg m(-3)) soil. After two days of root growth, the numbers of P. fluorescens (CFU cm(-1)) were highest on the section of root just below the seed with progressively fewer bacteria near the tip, irrespective of density. The decapped roots had significantly more colonies of P. fluorescens at the tip compared with the intact roots: approximately 100-fold more in the loose and 30-fold more in the compact soil. In addition, confocal images of the root tips grown in agar showed that P. fluorescens could only be detected on the tips of the decapped roots. These results indicated that border cells, and their associated mucilage, prevented complete colonization of the root tip by the biocontrol agent P. fluorescens, possibly by acting as a disposable surface or sheath around the cap.     

Root cap removal increases root penetration resistance in maize (Zea mays L)

The root cap assists the passage of the root through soil by means of its slimy mucilage secretion and by the sloughing of its outer cells. The root penetration resistance of decapped primary roots of maize (Zea mays L. cv. Mephisto) was compared with that of intact roots in loose (dry bulk density 1.0 g cm-3; penetration resistance 0.06 MPa) and compact soil (1.4 g cm-3; penetration resistance 1.0 MPa), to evaluate the contribution of the cap to decreasing the impedance to root growth. Root elongation rate and diameter were the same for decapped and intact roots when the plants were grown in loose soil. In compacted soil, however, the elongation rate of decapped roots was only about half that of intact roots, whilst the diameter was 30% larger. Root penetration resistances of intact and decapped seminal axis were 0.31 and 0.52 MPa, respectively, when the roots were grown in compacted soil. These results indicated that the presence of a root cap alleviates much of the mechanical impedance to root penetration, and enables roots to grow faster in compacted soils.     

Abscisic acid concentration,root pH and anatomy do not explain growth differences of chickpea (Cicer arietinum L.) and lupin (Lupinus angustifolius L.) on acid and alkaline soils

The ABA concentrations of leaves, roots, soils and transport fluids of chickpea and lupin plants growing in acid (pH=4.8) and alkaline (pH=8.0) soils and an acid soil with an alkaline subsoil and an alkaline soil with an acid subsoil were measured with the aim of explaining the poor growth of narrow-leafed lupins in alkaline soil. The ABA concentration in the leaves was higher in lupin than chickpea, but did not differ when the plants were grown in alkaline compared to acid soil. The ABA concentration of the roots and xylem sap of lupin did not differ significantly when grown in acid or alkaline soil. Chickpea roots and xylem sap had, however, lower ABA concentrations in acid soil. The ABA concentration in the soil solution was higher in the acid than in the alkaline soil. Roots of lupin and chickpea showed no suberization of the hypodermis or exodermis whether grown aeroponically or hydroponically and the pH of the cytoplasm did not change significantly when root cells of lupin and chickpea were exposed to external pHs of 4.8 or 8.0. The chickpea roots had greater suberization of the endodermal cells adjacent to radial xylem rays and maintained a slightly higher vacuolar pH than lupin in both acid and alkaline external media, but these small differences are insufficient to explain the reductions in lupin growth in alkaline soil.     

Carboxylate release of wheat, canola and 11 grain legume species as affected by phosphorus status

The capacity of plant roots to increase their carboxylate exudation at a low plant phosphorus (P) status is an adaptation to acquire sufficient P at low soil P availability. Our objective was to compare crop species in their adaptive response to a low-P availability, in order to gain knowledge to be used for improving crop P-acquisition efficiency from soils that are low in P or that have a high capacity to retain P. In the present screening study we compared 13 crop species, grown in sand at either 3 or 300 μM of P, and measured root mass ratio, cluster-root development, rhizosphere pH and carboxylate composition of root exudates. Root mass ratio decreased with increasing P supply for Triticum aestivum L., Brassica napus L., Cicer arietinum L. and Lens culinaris Medik., and increased only for Pisum sativum L., while the Lupinus species and Vicia faba L. were not responsive. Lupinus species that had the potential to produce root clusters either increased or decreased biomass allocation to clusters at 300 μM of P compared with allocation at 3 μM of P. All Lupinus species acidified their rhizosphere more than other species did, with average pH decreasing from 6.7 (control) to 4.3 for Lupinus pilosus L. and 5.9 for Lupinus atlanticus L.; B. napus maintained the most alkaline rhizosphere, averaging 7.4 at 300 μM of P. Rhizosphere carboxylate concentrations were lowest for T. aestivum, B. napus, V. faba, and L. culinaris than for the other species. Exuded carboxylates were mainly citrate and malate for all species, with the exception of L. culinaris and C. arietinum, which produced mainly citrate and malonate. Considerable variation in the concentration of exuded carboxylates and protons was found, even with a genus. Cluster-root forming species did not invariably have the highest concentrations of rhizosphere carboxylates. Lupinus species varied both in P-uptake and in the sensitivity of their cluster-root development to external P supply. Given the carbon cost of cluster roots, a greater plasticity in their formation and exudation (i.e. reduced investment in cluster roots and exudation at higher soil P, a negative feedback response) is a desirable trait for agricultural species that may have variable access to readily available P.     

Growth Medium and Phosphorus Supply Affect Cluster Root Formation and Citrate Exudation by Lupinus albus Grown in a Sand/Solution Split-Root System

We examined cluster root formation and root exudation by white lupin (Lupinus albus L. cv. Kiev Mutant) in response to growth medium and phosphorus supply in a sand/solution split-root system. The split-root system consisted of a nutrient solution compartment and a siliceous sand compartment. Phosphorus was applied at 1 (low-P plants) or 50 (high-P plants) μM as KH₂PO₄ to the solution compartment and at 10, 50 or 250 mg P kg⁻¹ as hydroxyapatite (Ca-P) to the sand compartment. In contrast to the high-P plants, P concentration and P uptake in the low-P plants increased with increasing P supply to the sand compartment. The NaHCO₃-extractable P was lower in the rhizosphere of the low-P plants than the high-P ones. The proton extrusion rate by the solution-grown roots of the low-P plants was higher than that of the high-P plants at the early growth stage. For the low-P plants, the proportion of dry root biomass allocated to cluster roots was higher in the solution compartment than that in the sand compartment. The citrate exudation increased in the sand compartment and decreased in the solution compartment with time, showing a lack of synchronization in citrate exudation by two root halves grown in different media. The cluster root proportion and citrate exudation in both compartments decreased with increasing shoot P concentration. An additional experiment with no P added to either root compartment showed that the proportion of cluster roots was about 9% lower in the sand than solution compartments. The results suggest that cluster root formation and citrate exudation can be significantly affected by the root growth medium in addition to being regulated by shoot P status. More P can be exploited from sparingly available Ca-P by the low-P plants than the high-P ones due to greater citrate exudation under P deficiency.     

Exudation of carboxylates in Australian Proteaceae: chemical composition

Roots of a wide range of plant species exude carboxylates, such as citrate, into the rhizosphere. In the present study, seedlings of a range of Australian Banksia, Hakea and Dryandra species (Proteaceae) were assayed for their exudation of carboxylates. All of these species (Hakea prostrata, Hakea undulata, Hakea petiolaris, Hakea baxteri, Banksia grandis, Banksia prionotes, Banksia occidentalis and Dryandra sessilis) form cluster roots when grown in nutrient solution with a low phosphate concentration. Exudation of carboxylates was studied for cluster roots and non‐cluster roots separately, and for the entire root system. Cluster roots of these Proteaceae exuded malate, malonate, lactate, acetate, maleate, citrate, fumarate, cis‐ and trans‐aconitate. The relative contributions of each of these carboxylates differed between species. Malate, malonate, lactate, citrate and trans‐aconitate, however, were invariably present in large proportions of total carboxylate exudation. Non‐cluster roots of H. prostrata exuded a spectrum of carboxylates (mainly malonate, lactate and citrate), which differed somewhat from the exudation pattern of cluster roots (mainly malate, malonate, lactate and citrate). The rate of exudation for cluster roots of the seven species was approximately 1·6 nmol g^?1 FM s^?1, which is considerably higher than that reported for a variety of crop and native species that do or do not form cluster roots. Contrary to what occurs in the cluster roots of Lupinus albus, which release carboxylates accompanied by protons so that the rhizosphere is acidified, the present Proteaceae exude the carboxylates as anions without concomitant proton release. The role of carboxylates in the mobilization of phosphate and other nutrients from soil is discussed.     

Functional significance of dauciform roots: exudation of carboxylates and acid phosphatase under phosphorus deficiency in Caustis blakei (Cyperaceae)

Caustis blakei produces an intriguing morphological adaptation by inducing dauciform roots in response to phosphorus (P) deficiency. We tested the hypothesis that these hairy, swollen lateral roots play a similar role to cluster roots in the exudation of organic chelators and ectoenzymes known to aid the chemical mobilization of sparingly available soil nutrients, such as P. Dauciform-root development and exudate composition (carboxylates and acid phosphatase activity) were analysed in C. blakei plants grown in nutrient solution under P-starved conditions. The distribution of dauciform roots in the field was determined in relation to soil profile depth and matrix. The percentage of dauciform roots of the entire root mass was greatest at the lowest P concentration ([P]) in solution, and was suppressed with increasing solution [P], while in the field dauciform roots were predominantely located in the upper soil horizons, and decreased with increasing soil depth. Citrate was the major carboxylate released in an exudative burst from mature dauciform roots, which also produced elevated levels of acid phosphatase activity. Malonate was the dominant internal carboxylate present, with the highest concentration in young dauciform roots. The high concentration of carboxylates and phosphatases released from dauciform roots, combined with their prolific distribution in the organic surface layer of nutrient-impoverished soils, provides an ecophysiological advantage for enhancing nutrient acquisition.     

Effects of external phosphorus supply on internal phosphorus concentration and the initiation,growth and exudation of cluster roots in Hakea prostrata R.Br.

The response of internal phosphorus concentration, cluster-root initiation, and growth and carboxylate exudation to different external P supplies was investigated in Hakea prostrata R.Br. using a split-root design. After removal of most of the taproot, equal amounts of laterals were allowed to grow in two separate pots fastened together at the top, so that the separate root halves could be exposed to different conditions. Plants were grown for 10 weeks in this system; one root half was supplied with 1 M P while the other halves were supplied with 0, 1, 25 or 75 M P. Higher concentrations of P supplied to one root half significantly increased the P concentration of those roots and in the shoots. The P concentrations in root halves supplied with 1 M P were invariably low, regardless of the P concentration supplied to the other root half. Cluster root initiation was completely suppressed on root halves supplied with 25 or 75 M P, whereas it continued on the other halves supplied with 1 M P indicating that cluster-root initiation was regulated by local root P concentration. Cluster-root growth (dry mass increment) on root halves supplied with 1 M P was significantly reduced when the other half was either deprived of P or supplied with 25 or 75 M P. Cluster-root growth was favoured by a low shoot P status at a root P supply that was adequate for increased growth of roots and shoots without increased tissue P concentrations. The differences in cluster-root growth on root halves with the same P supply suggest that decreased cluster-root growth was systemically regulated. Carboxylate-exudation rates from cluster roots on root halves supplied with 1 M P were the same, whether the other root half was supplied with 1, 25 or 75 M P, but were approximately 30 times faster when the other half was deprived of P. Estimates of root P-uptake rates suggest a rather limited capacity for down-regulating P uptake when phosphate was readily available.     

Root carboxylate exudation capacity under phosphorus stress does not improve grain yield in green gram

Key message

Genetic variability in carboxylate exudation capacity along with improved root traits was a key mechanism for P-efficient green gram genotype to cope with P-stress but it did not increase grain yield.

Abstract

This study evaluates genotypic variability in green gram for total root carbon exudation under low phosphorus (P) using ¹⁴C and its relationship with root exuded carboxylates, growth and yield potential in contrasting genotypes. Forty-four genotypes grown hydroponically with low (2 μM) and sufficient (100 μM) P concentrations were exposed to ¹⁴CO₂ to screen for total root carbon exudation. Contrasting genotypes were employed to study carboxylate exudation and their performance in soil at two P levels. Based on relative ¹⁴C exudation and biomass, genotypes were categorized. Carboxylic acids were measured in exudates and root apices of contrasting genotypes belonging to efficient and inefficient categories. Oxalic and citric acids were released into the medium under low-P. PDM-139 (efficient) was highly efficient in carboxylate exudation as compared to ML-818 (inefficient). In low soil P, the reduction in biomass was higher in ML-818 as compared to PDM-139. Total leaf area and photosynthetic rate averaged for genotypes increased by 71 and 41 %, respectively, with P fertilization. Significantly, higher root surface area and volume were observed in PDM-139 under low soil P. Though the grain yield was higher in ML-818, the total plant biomass was significantly higher in PDM-139 indicating improved P uptake and its efficient translation into biomass. The higher carboxylate exudation capacity and improved root traits in the later genotype might be the possible adaptive mechanisms to cope with P-stress. However, it is not necessary that higher root exudation would result in higher grain yield.     

Root exudation from Hordeum vulgare in response to localized nitrate supply

Root proliferation as a response to exploit zones of nutrient enrichment in soil has been demonstrated for a wide range of plant species. However, the effectiveness of this as a strategy to acquire nutrients is also dependent on interactions with the soil microbial community. Specifically, C-flow from roots modifies microbial activity and probably the balance between nutrient mineralization and immobilization processes in the rhizosphere. In this study, near-natural abundance 13C-labelling and gene-reporter methods were applied to determine the effects of uneven nitrate supply to roots of Hordeum vulgare on assimilate partitioning and root exudation. Plants were initially grown in uniform nitrate supply in split-root, sand microcosms after which one treatment continued to receive uniform supply, and the other received nitrate to one root compartment only. At the time of imposing the treatments, the CO2 supplied to the plants was switched to a cylinder source, providing a distinct delta13C-signature and allowing the fate of new assimilate within the plants to be determined. The labelling approach allowed quantification of the expected preferential allocation of new C-assimilate to roots in enriched nitrate, prior to any measurable effect on whole biomass or root architecture. Biosensor (lux-marked Pseudomonas fluorescens 10586 pUCD607) bioluminescence, quantified spatially by CCD imaging, demonstrated that root exudation was significantly increased for roots in enriched nitrate. This response of root exudation, being primarily associated with root apices and concurrent with enhanced assimilate supply, strongly suggests that C-flow from roots is an integral component of the proliferation response to nitrate.