Acclimative changes in root epidermal cell fate in response to Fe and P deficiency: a specific role for auxin?

Summary Root hair formation and the development of transfer cells in the rhizodermis was investigated in various existing auxinrelated mutants ofArabidopsis thaliana and in the tomato mutantdiageotropica. Wild-type Arabidopsis plants showed increased formation of root hairs when the seedlings were cultivated in Fe- or P-free medium. These extranumerary hairs were located in normal positions and in positions normally occupied by nonhair cells, e.g., over periclinal walls of underlying cortical cells. Defects in auxin transport or reduced auxin sensitivity inhibited the formation of root hairs in response to Fe deficiency completely but did only partly affect initiation and elongation of hairs in P-deficient roots. Application of the ethylene precursor 1-aminocyclopropane-1-carboxylic acid or the auxin analog 2,4-dichlorophenoxyacetic acid did not rescue the phenotype of the auxin-resistantaxr2 mutant under control and Fe-deficient conditions, indicating that functionalAXR2 product is required for translating the Fe deficiency signal into the formation of extra hairs. The development of extra hairs inaxr2 roots under P-replete conditions was not affected by auxin antagonists, suggesting that this process is independent of auxin signaling. In roots of tomato, growth under Fe-deficient conditions induced the formation of transfer cells in the root epidermis. Transfer cell frequency was enhanced by application of 2,4-dichlorophenoxyacetic acid but was not inhibited by the auxin transport inhibitor N-1-naphthylphthalamic acid. In thediageotropica mutant, which displays reduced sensitivity to auxin, transfer cells appeared to develop in both Fe-sufficient and Fe-deficient roots. Similar to the wild type, no reduction in transfer cell frequency was observed after application of the above auxin transport inhibitor. These data suggest that auxin has no primary function in inducing transfer cell development; the formation of transfer cells, however, appears to be affected by the hormonal balance of the plants.Abbreviations ACC 1-aminocyclopropane-1-carboxylic acid - TIBA triiodobenzoic acid - NPA N-1-naphthylphthalamic acid - STS silver thiosulfate     

Role of hormones in the induction of iron deficiency responses in Arabidopsis roots

In "strategy I" plants, several alterations in root physiology and morphology are induced by Fe deficiency, although the mechanisms by which low Fe levels are translated into reactions aimed at alleviating Fe shortage are largely unknown. To prove whether changes in hormone concentration or sensitivity are involved in the adaptation to suboptimal Fe availability, we tested 45 mutants of Arabidopsis defective in hormone metabolism and/or root hair formation for their ability to increase Fe(III) chelate reductase activity and to initiate the formation and enlargement of root hairs. Activity staining for ferric chelate reductase revealed that all mutants were responsive to Fe deficiency, suggesting that hormones are not necessary for the induction. Treatment of wild-type plants with the ethylene precursor 1-aminocyclopropane-1-carboxylic acid caused the development of root hairs in locations normally occupied by non-hair cells, but did not stimulate ferric reductase activity. Ectopic root hairs were also formed in -Fe roots, suggesting a role for ethylene in the morphological responses to Fe deficiency. Ultrastructural analysis of rhizodermal cells indicated that neither Fe deficiency nor 1-aminocyclopropane-1-carboxylic acid treatment caused transfer-cell-like alterations in Arabidopsis roots. Our data indicate that the morphological and physiological components of the Fe stress syndrome are regulated separately.     

Environmentally induced plasticity of root hair development in Arabidopsis

Postembryonic development of plants is dependent on both intrinsic genetic programs and environmental factors. The plasticity of root hair patterning in response to environmental signals was investigated in the Columbia-0 wild type and 19 Arabidopsis mutants carrying lesions in various parts of the root hair developmental pathway by withholding phosphate or iron (Fe) from the nutrient medium. In the aging primary root and in laterals of the wild type, the number of root hairs increased in response to phosphate and Fe deficiency in a manner typical of each growth type. Although an increase in root hair density in -phosphorus plants was mainly achieved by the formation of extra hairs over both tangential and radial wall of underlying cortical cells, roots of -Fe plants were characterized by a high percentage of extra hairs with two tips. Root hair patterning and hair length was differentially affected by the presence or absence of phosphate and Fe among the genotypes under investigation, pointing to separate cascades of gene activation under all three growth conditions. Divergence in root hair patterning was most pronounced among mutants with defects in genes that affect the first stages of differentiation, suggesting that nutritional signals are perceived at an early stage of epidermal cell development. During elongation of the root hairs, no differences in the requirement of gene products between the growth types were obvious. The role of genes involved in root hair development in the aging primary root of Arabidopsis under the various growth conditions is discussed.     

Modulation of the root epidermal phenotype by hormones,inhibitors and iron regime

Patterning of epidermal cells is subject to genetic regulation but also influenced by environmental stimuli. To adapt to unfavorable environmental conditions plants have developed various mechanisms to increase the plasma membrane's surface area of epidermal root cells, for example through the formation of root hairs and differentiation of rhizodermal transfer cells. Mechanisms controlling cell fate speciation in the rhizodermis were investigated by application of hormones and hormone antagonists. In addition, the effect of Fe deficiency on root epidermal patterning and Fe(III)-reduction activity was examined. In the iron-hyperaccumulating pea mutants dgl and brz and in the Arabidopsis mutant man1 Fe(III)-reduction activity was found to be up-regulated under both high and low iron supply. In contrast, morphological responses such as the development of transfer cells and extranumerary root hairs was repressed by a high iron concentration in the external medium. All morphological responses can be mimicked by exogenous application of the ethylene precursor 1-aminocyclopropane-1-carboxylic acid (ACC) or the auxin analog 2,4-dichlorophenoxyacetic acid (2,4-D). Conversely, Fe(III)-reduction rates were not influenced or only slightly affected by the hormone treatment. Application of inhibitors of ethylene synthesis, ethylene action or auxin transport was effective only in inhibiting the formation of extra root hairs, indicating that these hormones are not required for transfer cell formation or expression of Fe(III) reduction. These data suggest that the Fe reductase induced by iron stress does not depend on the formation of transfer cells and further imply separate regulatory pathways for the two responses. The data are compatible with a model in which root reduction activity is modulated by a shoot-borne signal coordinating iron uptake with the shoot demand, while the epidermal phenotype is primarily dependent on the intracellular iron concentration of root cells.     

Auxin modulates the enhanced development of root hairs in Arabidopsis thaliana (L.) Heynh. under elevated CO(2)

Root hairs may play a critical role in nutrient acquisition of plants grown under elevated CO(2) . This study investigated how elevated CO(2) enhanced the development of root hairs in Arabidopsis thaliana (L.) Heynh. The plants under elevated CO(2) (800 μL L(-1)) had denser and longer root hairs, and more H-positioned cells in root epidermis than those under ambient CO(2) (350 μL L(-1)). The elevated CO(2) increased auxin production in roots. Under elevated CO(2) , application of either 1-naphthoxyacetic acid (1-NOA) or N-1-naphthylphthalamic acid (NPA) blocked the enhanced development of root hairs. The opposite was true when the plants under ambient CO(2) were treated with 1-naphthylacetic acid (NAA), an auxin analogue. Furthermore, the elevated CO(2) did not enhance the development of root hairs in auxin-response mutants, axr1-3, and auxin-transporter mutants, axr4-1, aux1-7 and pin1-1. Both elevated CO(2) and NAA application increased expressions of caprice, triptychon and rho-related protein from plants 2, and decreased expressions of werewolf, GLABRA2, GLABRA3 and the transparent testa glabra 1, genes related to root-hair development, while 1-NOA and NPA application had an opposite effect. Our study suggests that elevated CO(2) enhanced the development of root hairs in Arabidopsis via the well-characterized auxin signalling and transport that modulate the initiation of root hairs and the expression of its specific genes.     

Formation of transfer cells and H(+)-ATPase expression in tomato roots under P and Fe deficiency

In roots of tomato ( Lycopersicon esculentum Mill.), extranumerary root hairs and transfer cell-like wall ingrowth depositions in the rhizodermis were developed in response to P and Fe deficiency. Immunocytolocalization of the plasma membrane H(+)-ATPase in roots of P-deficient plants revealed no appreciable increase in H(+)-ATPase density relative to control plants. In transfer cells, immunogold labeling was considerably higher than in ordinary rhizodermal cells. H(+)-ATPase sites were asymmetrically distributed in cells with and without wall ingrowths under P-deficient conditions. A split-root study revealed that the frequency of transfer cells was higher in the low-P half of the root system, but the density of H(+)-ATPase molecules was enhanced only in the high-P half of the split roots, suggesting that formation of transfer cells was controlled directly by the external Pi concentration, whereas ATPase expression was regulated indirectly by the internal nutrient status of the plant. The role of hormones in the induction of transfer cells was investigated by treating plants with the ethylene precursor 1-aminocyclopropane-1-carboxylic acid (ACC) or various ethylene antagonists. Transfer cells were induced by ACC to an extent similar to that observed after P or Fe starvation, but inhibitors of either ethylene synthesis or action did not decrease their frequency. These results suggest that ethylene was not required for the induction of transfer cells but changes in ethylene levels appeared to modulate the number of cells forming wall ingrowths. In roots of ethylene-insensitive Never-ripe tomato plants the frequency of transfer cells was rather increased than decreased under most growth conditions relative to the wild type, indicating that ethylene responsiveness played no critical role in the differentiation of transfer cells and that the transduction of signals ultimately leading to their formation was independent of the ethylene signaling cascade.     

The effect of iron on the primary root elongation of Arabidopsis during phosphate deficiency

Root architecture differences have been linked to the survival of plants on phosphate (P)-deficient soils, as well as to the improved yields of P-efficient crop cultivars. To understand how these differences arise, we have studied the root architectures of P-deficient Arabidopsis (Arabidopsis thaliana Columbia-0) plants. A striking aspect of the root architecture of these plants is that their primary root elongation is inhibited when grown on P-deficient medium. Here, we present evidence suggesting that this inhibition is a result of iron (Fe) toxicity. When the Fe concentration in P-deficient medium is reduced, we observe elongation of the primary root without an increase in P availability or a corresponding change in the expression of P deficiency-regulated genes. Recovery of the primary root elongation is associated with larger plant weights, improved ability to take up P from the medium, and increased tissue P content. This suggests that manipulating Fe availability to a plant could be a valuable strategy for improving a plant's ability to tolerate P deficiency.     

Regulatory components involved in altering lateral root development in response to localized iron: Evidence for natural genetic variation

On the search for sparingly available nutrients, plants may alter their root architecture to improve soil exploration. So far, the examples for root system modifications induced by a heterogeneous availability of nutrients have been reported for the macronutrients nitrogen (N) and phosphorous (P). In an attempt to extend this type of knowledge to other nutrients, we recently provided evidence that Arabidopsis roots are able to sense a local availability of the micronutrient iron (Fe) and to respond with lateral root elongation into the Fe-containing patch. This specific root response was caused by enhanced elongation of cells leaving the root meristem and was dependent on an AUX1-mediated auxin accumulation in the lateral root apices. In this report, we compare mechanisms underlying this response with those known for other nutrients and show that a substantial genotypic variation exists among accessions of A. thaliana in the responsiveness of lateral roots toward localized Fe supplies.     

Physiological Effects of the Synthetic Strigolactone Analog GR24 on Root System Architecture in Arabidopsis: Another Belowground Role for Strigolactones?

In this study, the role of the recently identified class of phytohormones, strigolactones, in shaping root architecture was addressed. Primary root lengths of strigolactone-deficient and -insensitive Arabidopsis (Arabidopsis thaliana) plants were shorter than those of wild-type plants. This was accompanied by a reduction in meristem cell number, which could be rescued by application of the synthetic strigolactone analog GR24 in all genotypes except in the strigolactone-insensitive mutant. Upon GR24 treatment, cells in the transition zone showed a gradual increase in cell length, resulting in a vague transition point and an increase in transition zone size. PIN1/3/7-green fluorescent protein intensities in provascular tissue of the primary root tip were decreased, whereas PIN3-green fluorescent protein intensity in the columella was not affected. During phosphate-sufficient conditions, GR24 application to the roots suppressed lateral root primordial development and lateral root forming potential, leading to a reduction in lateral root density. Moreover, auxin levels in leaf tissue were reduced. When auxin levels were increased by exogenous application of naphthylacetic acid, GR24 application had a stimulatory effect on lateral root development instead. Similarly, under phosphate-limiting conditions, endogenous strigolactones present in wild-type plants stimulated a more rapid outgrowth of lateral root primordia when compared with strigolactone-deficient mutants. These results suggest that strigolactones are able to modulate local auxin levels and that the net result of strigolactone action is dependent on the auxin status of the plant. We postulate that the tightly balanced auxin-strigolactone interaction is the basis for the mechanism of the regulation of the plants' root-to-shoot ratio.     

A multidrug and toxic compound extrusion (MATE) transporter modulates auxin levels in root to regulate root development and promotes aluminium tolerance

MATE (multidrug and toxic compound extrusion) transporters play multiple roles in plants including detoxification, secondary metabolite transport, aluminium (Al) tolerance, and disease resistance. Here we identify and characterize the role of the Arabidopsis MATE transporter DETOXIFICATION30. AtDTX30 regulates auxin homeostasis in Arabidopsis roots to modulate root development and Al-tolerance. DTX30 is primarily expressed in roots and localizes to the plasma membrane of root epidermal cells including root hairs. dtx30 mutants exhibit reduced elongation of the primary root, root hairs, and lateral roots. The mutant seedlings accumulate more auxin in their root tips indicating role of DTX30 in maintaining auxin homeostasis in the root. Al induces DTX30 expression and promotes its localization to the distal transition zone. dtx30 seedlings accumulate more Al in their roots but are hyposensitive to Al-mediated rhizotoxicity perhaps due to saturation in root growth inhibition. Increase in expression of ethylene and auxin biosynthesis genes in presence of Al is absent in dtx30. The mutants exude less citrate under Al conditions, which might be due to misregulation of AtSTOP1 and the citrate transporter AtMATE. In conclusion, DTX30 modulates auxin levels in root to regulate root development and in the presence of Al indirectly modulates citrate exudation to promote Al tolerance.     

Growth promotion of Chinese cabbage and Arabidopsis by Piriformospora indica is not stimulated by mycelium-synthesized auxin

Piriformospora indica, an endophytic fungus of the order Sebacinales, interacts with the roots of a large variety of plant species. We compared the interaction of this fungus with Chinese cabbage (Brassica campestris subsp. chinensis) and Arabidopsis seedlings. The development of shoots and roots of Chinese cabbage seedlings was strongly promoted by P. indica and the fresh weight of the seedlings increased approximately twofold. The strong stimulation of root hair development resulted in a bushy root phenotype. The auxin level in the infected Chinese cabbage roots was twofold higher compared with the uncolonized controls. Three classes of auxin-related genes, which were upregulated by P. indica in Chinese cabbage roots, were isolated from a double-subtractive expressed sequence tag library: genes for proteins related to cell wall acidification, intercellular auxin transport carrier proteins such as AUX1, and auxin signal proteins. Overexpression of B. campestris BcAUX1 in Arabidopsis strongly promoted growth and biomass production of Arabidopsis seedlings and plants; the roots were highly branched but not bushy when compared with colonized Chinese cabbage roots. This suggests that BcAUX1 is a target of P. indica in Chinese cabbage. P. indica also promoted growth of Arabidopsis seedlings but the auxin levels were not higher and auxin genes were not upregulated, implying that auxin signaling is a more important target of P. indica in Chinese cabbage than in Arabidopsis. The fungus also stimulated growth of Arabidopsis aux1 and aux1/axr4 and rhd6 seedlings. Furthermore, a component in an exudate fraction from P. indica but not auxin stimulated growth of Chinese cabbage and Arabidopsis seedlings. We propose that activation of auxin biosynthesis and signaling in the roots might be the cause for the P. indica-mediated growth phenotype in Chinese cabbage.     

Nitric Oxide Functions as a Positive Regulator of Root Hair Development

The root epidermis is composed of two cell types: trichoblasts (or hair cells) and atrichoblasts (or non-hair cells). In lettuce (Lactuca sativa cv. Grand Rapids var. Rapidmor oscura) plants grown hydroponically in water, the root epidermis did not form root hairs. The addition of 10 µM sodium nitroprusside (SNP), a nitric oxide (NO) donor, resulted in almost all rhizodermal cells differentiated into root hairs. Treatment with the synthetic auxin 1-naphthyl acetic acid (NAA) displayed a significant increase of root hair formation (RHF) that was prevented by the specific NO scavenger carboxy-PTIO (cPTIO). In Arabidopsis, two mutants have been shown to be defective in NO production and to display altered phenotypes in which NO is implicated. Arabidopsis nos1 has a mutation in an NO synthase structural gene (NOS1), and the nia1 nia2 double mutant is null for nitrate reductase (NR) activity. We observed that both mutants were affected in their capacity of developing root hairs. Root hair elongation was significantly reduced in nos1 and nia1 nia2 mutants as well as in cPTIO-treated wild type plants. A correlation was found between endogenous NO level in roots detected by the fluorescent probe DAF-FM DA and RHF. In Arabidopsis, as well as in lettuce, cPTIO blocked the NAA-induced root hair elongation. Taken together, these results indicate that: (1) NO is a critical molecule in the process leading to RHF and (2) NO is involved in the auxin-signaling cascade leading to RHF.Key Words: auxin, nitric oxide, root hair, lettuce, arabidopsis, nos1 mutant, nia1, nia2 mutant     

Genotypic differences in the presence of hairs on roots and gynophores of peanuts (Arachis hypogaea L.) and their significance for phosphorus uptake

Root hairs substantially increase the surface area of plant roots with positive effects for phosphorus (P) uptake, but the ability of peanuts to form root hairs has been questioned. The aim was to examine hair development on roots and gynophores of a variety of peanut genotypes and to relate genotypic differences in hair formation to differences in P uptake. Five out of eighteen genotypes completely lacked hairs on both organs whereas others consistently developed hairs on roots and gynophores, although with considerable variation in hair density. The ability to form root hairs as well as root hair density concurred with the presence and density of hairs on gynophores, suggesting a possible connection between both developmental processes. The contribution of root hairs to P uptake was studied in three genotypes differing in hair density. The final amount of P taken up by roots did not differ between genotypes but two distinct P uptake strategies could be identified. The genotype lacking root hairs maintained P uptake due to the development of a large root system whereas densely covered roots of genotype 'Wasedairyu' were three times as efficient in extracting P from a P-deficient soil. Furthermore P uptake through gynophores contributed about 20% to the total P uptake of Wasedairyu but only insignificant amounts to other genotypes. The ability to form hairs on roots and gynophores can therefore be seen as an adaptation to low P availability and if combined with a large root system, could substantially increase the tolerance of peanuts to P deficiency.     

The rhd6 Mutation of Arabidopsis thaliana Alters Root-Hair Initiation through an Auxin- and Ethylene-Associated Process

Root-hair initiation in Arabidopsis thaliana provides a model for studying cell polarity and its role in plant morphogenesis. Root hairs normally emerge at the apical end of root epidermal cells, implying that these cells are polarized. We have identified a mutant, rhd6, that displays three defects: (a) a reduction in the number of root hairs, (b) an overall basal shift in the site of root-hair emergence, and (c) a relatively high frequency of epidermal cells with multiple root hairs. These defects implicate the RHD6 gene in root-hair initiation and indicate that RHD6 is normally associated with the establishment of, or response to, root epidermal cell polarity. Similar alterations in the site of root-hair emergence, although less extreme, were also discovered in roots of the auxin-, ethylene-, abscisic acid-resistant mutant axr2 and the ethylene-resistant mutant etr1. All three rhd6 mutant phenotypes were rescued when either auxin (indoleacetic acid) or an ethylene precursor (1-aminocyclopropane-1-carboxylic acid) was included in the growth medium. The rhd6 root phenotypes could be phenocopied by treating wild-type seedlings with an inhibitor of the ethylene pathway (aminoethoxyvinylglycine). These results indicate that RHD6 is normally involved in directing the selection or assembly of the root-hair initiation site through a process involving auxin and ethylene.     

Localized iron supply triggers lateral root elongation in Arabidopsis by altering the AUX1-mediated auxin distribution

Root system architecture depends on nutrient availability, which shapes primary and lateral root development in a nutrient-specific manner. To better understand how nutrient signals are integrated into root developmental programs, we investigated the morphological response of Arabidopsis thaliana roots to iron (Fe). Relative to a homogeneous supply, localized Fe supply in horizontally separated agar plates doubled lateral root length without having a differential effect on lateral root number. In the Fe uptake-defective mutant iron-regulated transporter1 (irt1), lateral root development was severely repressed, but a requirement for IRT1 could be circumvented by Fe application to shoots, indicating that symplastic Fe triggered the local elongation of lateral roots. The Fe-stimulated emergence of lateral root primordia and root cell elongation depended on the rootward auxin stream and was accompanied by a higher activity of the auxin reporter DR5-β-glucuronidase in lateral root apices. A crucial role of the auxin transporter AUXIN RESISTANT1 (AUX1) in Fe-triggered lateral root elongation was indicated by Fe-responsive AUX1 promoter activities in lateral root apices and by the failure of the aux1-T mutant to elongate lateral roots into Fe-enriched agar patches. We conclude that a local symplastic Fe gradient in lateral roots upregulates AUX1 to accumulate auxin in lateral root apices as a prerequisite for lateral root elongation.     

Acid phosphatase activity in phosphorus-deficient white lupin roots

White lupin ( Lupinus albus L.) develops proteoid roots when grown in phosphorus (P)-deficient conditions. These short, lateral, densely clustered roots are adapted to increase P availability. Previous studies from our laboratory have shown proteoid roots have higher rates of non-photosynthetic carbon fixation than normal roots and altered metabolism to support organic acid exudation, which serves to solubilize P in the rhizosphere. The present work indicates that proteoid roots possess additional adaptations for increasing P availability and possibly for conserving P in the plant. Roots from P-deficient (–P) plants had significantly greater acid phosphatase activity in both root extracts and root exudates than comparable samples from P-sufficient (+P) plants beginning 10 d after emergence. The increase in activity in –P plants was most pronounced in the proteoid regions. In contrast, no induction of phytase activity was found in –P plants compared to +P plants. The number of proteoid roots present was not affected by the source of phosphorus supplied, whether organic or inorganic forms. Adding molybdate to the roots increased the number of proteoid roots in plants supplied with organic P, but not inorganic P. Increased acid phosphatase activity was detected in root exudates in the presence of organic P sources. Native-polyacrylamide gel electrophoresis demonstrated that under P-deficient conditions, a unique isoform of acid phosphatase was induced between 10 and 12 d after emergence. This isoform was found not only within the root, but it comprised the major form exuded from proteoid roots of –P plants. The fact that exudation of proteoid-root-specific acid phosphatase coincides with proteoid root development and increased exudation of organic acids indicates that white lupin has several coordinated adaptive strategies to P-deficient conditions.     

Ups and downs of tissue and planar polarity in plants

The polar orientation of cells within a tissue is an intensively studied research area in animal cells. The term planar polarity refers to the common polar arrangement of cells within the plane of an epithelium. In plants, the subcellular analysis of tissue polarity has been limited by the lack of appropriate markers. Recently, research on plant tissue polarity has come of age. Advances are based on studies of Arabidopsis patterning, cell polarity and auxin transport mutants employing the coordinated, polar localization of auxin transporters and the planar polarity of root epidermal hairs as markers. These approaches have revealed auxin transport and response, vesicular trafficking, membrane sterol and cytoskeletal requirements of tissue polarity. This review summarizes recent progress in research on vascular tissue and planar epidermal polarity in the Arabidopsis root and compares it to findings on planar polarity in animals and cell polarity in yeast.     

SCHIZORIZA controls an asymmetric cell division and restricts epidermal identity in the Arabidopsis root

The primary root of Arabidopsis has a simple cellular organisation. The fixed radial cell pattern results from stereotypical cell divisions that occur in the meristem. Here we describe the characterisation of schizoriza (scz), a mutant with defective radial patterning. In scz mutants, the subepidermal layer (ground tissue) develops root hairs. Root hairs normally only form on epidermal cells of wild-type plants. Moreover, extra periclinal divisions (new wall parallel to surface of the root) occur in the scz root resulting in the formation of supernumerary layers in the ground tissue. Both scarecrow (scr) and short root (shr) suppress the extra periclinal divisions characteristic of scz mutant roots. This results in the formation of a single layered ground tissue in the double mutants. Cells of this layer develop root hairs, indicating that mis-specification of the ground tissue in scz mutants is uncoupled to the cell division defect. This suggests that during the development of the ground tissue SCZ has two distinct roles: (1) it acts as a suppressor of epidermal fate in the ground tissue, and (2) it is required to repress periclinal divisions in the meristem. It may act in the same pathway as SCR and SHR.     

The role of cell wall components from groundnut roots in solubilizing sparingly soluble phosphorus in low fertility soils

Groundnuts have a superior ability to take up P from soils with low P fertility compared to sorghum and soybean. Previous experiments showed that this ability was neither attributable to better root development nor to root exudates capable of solubilizing Fe- and Al-bound P, the sparingly soluble P forms in soils. Direct "contact reactions" between cell wall components from these 3 plant species (groundnut, soybean and sorghum) and P-fixing Fe and Al minerals were examined. Cell wall preparations from groundnut roots showed a superior P solubilizing ability than those of soybean and sorghum. Cell wall activity of groundnut roots may thus at least partly explain the superior growth of this crop under P-deficient conditions. To characterize the active site responsible for P solubilization, effects of pH, heat, addition of cations, and digestion with enzymes (pectinase and cellulase) or HCl on P solubilization were investigated. Conclusion are 1) Solubilizing ability is not related to root CEC because soybean with higher root CEC showed an inferior solubilizing ability compared to groundnut. 2) The reaction site of cell-walls of groundnut roots is stable against heating and digestion with cellulase and pectinase. 3) Solubilizing ability was severely reduced by digestion with HCl. 4) Pre-treating cell walls with either Al3+, Fe3+, or Ga3+ decreased solubilizing ability but cations with lower valency such as Na+, K+, Ca2+ or Mg2+ had no effect. Soaking roots of groundnuts grown in solution culture in 0.5 M NaOH for 30 seconds prior to cell wall preparation led to a 30% reduction in solubilization of P from FePO4 without permanently damaging plants. This suggests that 5) the active component of the cell walls was located on the root epidermal cell surfaces. Based on these results a phosphorus solubilizing mechanism is proposed.