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PubMed2004

Proton-transfer-reaction mass spectrometry as a new tool for real time analysis of root-secreted volatile organic compounds in Arabidopsis.

Plant roots release about 5% to 20% of all photosynthetically-fixed carbon, and as a result create a carbon-rich environment for numerous rhizosphere organisms, including plant pathogens and symbiotic microbes. Although some characterization of root exudates has been achieved, especially of secondary metabolites and proteins, much less is known about volatile organic compounds (VOCs) released by roots. In this communication, we describe a novel approach to exploring these rhizosphere VOCs and their induction by biotic stresses. The VOC formation of Arabidopsis roots was analyzed using proton-transfer-reaction mass spectrometry (PTR-MS), a new technology that allows rapid and real time analysis of most biogenic VOCs without preconcentration or chromatography. Our studies revealed that the major VOCs released and identified by both PTR-MS and gas chromatography-mass spectrometry were either simple metabolites, ethanol, acetaldehyde, acetic acid, ethyl acetate, 2-butanone, 2,3,-butanedione, and acetone, or the monoterpene, 1,8-cineole. Some VOCs were found to be produced constitutively regardless of the treatment; other VOCs were induced specifically as a result of different compatible and noncompatible interactions between microbes and insects and Arabidopsis roots. Compatible interactions of Pseudomonas syringae DC3000 and Diuraphis noxia with Arabidopsis roots resulted in the rapid release of 1,8-cineole, a monoterpene that has not been previously reported in Arabidopsis. Mechanical injuries to Arabidopsis roots did not produce 1,8-cineole nor any C6 wound-VOCs; compatible interactions between Arabidopsis roots and Diuraphis noxia did not produce any wound compounds. This suggests that Arabidopsis roots respond to wounding differently from above-ground plant organs. Trials with incompatible interactions did not reveal a set of compounds that was significantly different compared to the noninfected roots. The PTR-MS method may open the way for functional root VOC analysis that will complement genomic investigations in Arabidopsis.

Animals↗
PubMed2007

Characterization of a selenate-resistant Arabidopsis mutant. Root growth as a potential target for selenate toxicity.

Screening an Arabidopsis (Arabidopsis thaliana) T-DNA mutant library for selenate resistance enabled us to isolate a selenate-resistant mutant line (sel1-11). Molecular and genetic characterization showed that the mutant contained a lesion in the SULTR1;2 gene that encodes a high affinity root sulfate transporter. We showed that SULTR1;2 is the only gene among 13 mutated genes of the Arabidopsis sulfate transporter family whose mutation conferred selenate resistance to Arabidopsis. The selenate resistance phenotype of the sel1-11 mutant was mirrored by an 8-fold increase of root growth in the presence of selenate as shown by the calculated lethal concentration values. The impairment of SULTR1;2 activity in sel1-11 resulted in a reduced (35)S-sulfate uptake capacity by both roots and calli and a reduced sulfate and selenate content in root, shoot, and calli. Comparing sulfate-to-selenate ratios instead of absolute sulfate and selenate contents in roots and shoots enabled us to gain better insight into the mechanism of selenate toxicity in Arabidopsis. Roots of the sel1-11 mutant line showed a higher sulfate to selenate ratio than that of wild-type roots, while there were no significant differences in sulfate to selenate ratios in shoots of wild-type and mutant lines. These results indicated that the mechanism that confers the selenate resistance phenotype to the sel1-11 line takes place rather in the roots. It might be in part the result of a lower selenate uptake and of a protective effect of sulfate against the toxic effects of selenate on root growth. These results revealed in plants a central and specific role of the transporter SULTR1;2 in selenate sensitivity; they further suggested that root growth and potentially the root tip activity might be a specific target of selenate toxicity in Arabidopsis.

Anion Transport Proteins↗
PubMed2006

Extracellular proteins in pea root tip and border cell exudates.

Newly generated plant tissue is inherently sensitive to infection. Yet, when pea (Pisum sativum) roots are inoculated with the pea pathogen, Nectria haematococca, most newly generated root tips remain uninfected even though most roots develop lesions just behind the tip in the region of elongation. The resistance mechanism is unknown but is correlated spatially with the presence of border cells on the cap periphery. Previously, an array of >100 extracellular proteins was found to be released while border cell separation proceeds. Here we report that protein secretion from pea root caps is induced in correlation with border cell separation. When this root cap secretome was proteolytically degraded during inoculation of pea roots with N. haematococca, the percentage of infected root tips increased from 4% +/- 3% to 100%. In control experiments, protease treatment of conidia or roots had no effect on growth and development of the fungus or the plant. A complex of >100 extracellular proteins was confirmed, by multidimensional protein identification technology, to comprise the root cap secretome. In addition to defense-related and signaling enzymes known to be present in the plant apoplast were ribosomal proteins, 14-3-3 proteins, and others typically associated with intracellular localization but recently shown to be extracellular components of microbial biofilms. We conclude that the root cap, long known to release a high molecular weight polysaccharide mucilage and thousands of living cells into the incipient rhizosphere, also secretes a complex mixture of proteins that appear to function in protection of the root tip from infection.

14-3-3 Proteins↗
PubMed1996

Arabidopsis thaliana sku mutant seedlings show exaggerated surface-dependent alteration in root growth vector.

Roots of wild-type Arabidopsis thaliana seedlings in the Wassilewskija (WS) and Landsberg erecta (Ler) ecotypes often grow aslant on vertical agar surfaces. Slanted root growth always occurs to the right of the gravity vector when the root is viewed through the agar surface, and is not observed in the Columbia ecotype. Right-slanted root growth is surface-dependent and does not result directly from directional environmental stimuli or gradients in the plane of skewing. We have isolated two partially dominant mutations in WS (sku1 and sku2) that show an exaggerated right-slanting root-growth phenotype on agar surfaces. The right-slanting root-growth phenotype of wild-type and mutant roots is not the result of diagravitropism or of an alteration in root gravitropism. It is accompanied by a left-handed rotation of the root about its axis within the elongation zone, the rate of which positively correlates with the degree of right-slanted curvature. Our data suggest that the right-slanting root growth phenotype results from an endogenous structural asymmetry that expresses itself by a directional root-tip rotation.

Arabidopsis↗
PubMed1999

A maize glycine-rich protein is synthesized in the lateral root cap and accumulates in the mucilage.

The root cap functions in the perception of gravity, the protection of the root apical meristem, and facilitation of the passage of roots through the soil, but the genes involved in these functions are poorly understood. Here we report the isolation of a root-specific gene from the cap of maize (Zea mays L.) primary root by cDNA subtraction and differential screening. The gene zmGRP4 (Z. mays glycine rich protein 4) encodes a member of the glycine-rich proteins with a putative signal peptide at the amino terminus. The deduced molecular mass of mature zmGRP4 is 14.4 kD. In situ-hybridization analysis has shown zmGRP4 to be strongly expressed in the lateral root cap and weakly expressed in the root epidermis. A polyclonal antibody raised against recombinant zmGRP4 detected a protein of 36 kD in the insoluble protein fraction extracted from the root tip and the root proper, indicating posttranslational modification(s) of zmGRP4. Immunohistochemical analysis showed the accumulation of zmGRP4 in the mucilage that covers the root tip. These results indicate that lateral root-cap cells secrete modified zmGRP4 into the mucilage to which the protein may contribute to its characteristic physical properties.

Amino Acid Sequence↗
PubMed2000

Basipetal auxin transport is required for gravitropism in roots of Arabidopsis.

Auxin transport has been reported to occur in two distinct polarities, acropetally and basipetally, in two different root tissues. The goals of this study were to determine whether both polarities of indole-3-acetic acid (IAA) transport occur in roots of Arabidopsis and to determine which polarity controls the gravity response. Global application of the auxin transport inhibitor naphthylphthalamic acid (NPA) to roots blocked the gravity response, root waving, and root elongation. Immediately after the application of NPA, the root gravity response was completely blocked, as measured by an automated video digitizer. Basipetal [(3)H]IAA transport in Arabidopsis roots was inhibited by NPA, whereas the movement of [(14)C]benzoic acid was not affected. Inhibition of basipetal IAA transport by local application of NPA blocked the gravity response. Inhibition of acropetal IAA transport by application of NPA at the root-shoot junction only partially reduced the gravity response at high NPA concentrations. Excised root tips, which do not receive auxin from the shoot, exhibited a normal response to gravity. The Arabidopsis mutant eir1, which has agravitropic roots, exhibited reduced basipetal IAA transport but wild-type levels of acropetal IAA transport. These results support the hypothesis that basipetally transported IAA controls root gravitropism in Arabidopsis.

Arabidopsis↗
PubMed2000

Genotypical differences in aluminum resistance of maize are expressed in the distal part of the transition zone. Is reduced basipetal auxin flow involved in inhibition of root elongation by aluminum?

Short-term Al treatment (90 microM Al at pH 4.5 for 1 h) of the distal transition zone (DTZ; 1-2 mm from the root tip), which does not contribute significantly to root elongation, inhibited root elongation in the main elongation zone (EZ; 2.5-5 mm from the root tip) to the same extent as treatment of the entire maize (Zea mays) root apex. Application of Al to the EZ had no effect on root elongation. Higher genotypical resistance to Al applied to the entire root apex, and specifically to the DTZ, was expressed by less inhibition of root elongation, Al accumulation, and Al-induced callose formation, primarily in the DTZ. A characteristic pH profile along the surface of the root apex with a maximum of pH 5.3 in the DTZ was demonstrated. Al application induced a substantial flattening of the pH profile moreso in the Al-sensitive than in the Al-resistant cultivar. Application of indole-3-acetic acid to the EZ but not to the meristematic zone significantly alleviated the inhibition of root elongation induced by the application of Al to the DTZ. Basipetal transport of exogenously applied [(3)H]indole-3-acetic acid to the meristematic zone was significantly inhibited by Al application to the DTZ in the Al-sensitive maize cv Lixis. Our results provide evidence that the primary mechanisms of genotypical differences in Al resistance are located within the DTZ, and suggest a signaling pathway in the root apex mediating the Al signal between the DTZ and the EZ through basipetal auxin transport.

Aluminum↗
PubMed2001

Gibberellin biosynthesis mutations and root development in pea.

Dwarf mutants of pea (Pisum sativum), with impaired gibberellin (GA) biosynthesis in the shoot, were studied to determine whether the roots of these genotypes had altered elongation and GA levels. Mutations na, lh-2, and ls-1 reduced GA levels in root tips and taproot elongation, although in lh-2 and ls-1 roots the reduction in elongation was small (less than 15%). The na mutation reduced taproot length by about 50%. The roots of na plants elongated in response to applied GA(1) and recombining na with mutation sln (which blocks GA catabolism) increased GA(1) levels in root tips and completely restored normal root development. In shoots, Mendel's le-1 mutation impairs the 3beta-hydroxylation of GA(20) to the bioactive GA(1), resulting in dwarfism. However, GA(1) and GA(20) levels were normal in le-1 roots, as was root development. The null mutation le-2 also did not reduce root GA levels or elongation. The results support the theory that GAs are important for normal root elongation in pea, and indicate that a 3beta-hydroxylase gene other than LE operates in pea roots.

Gibberellins↗
PubMed2005

Control of root cap formation by MicroRNA-targeted auxin response factors in Arabidopsis.

The plant root cap mediates the direction of root tip growth and protects internal cells. Root cap cells are continuously produced from distal stem cells, and the phytohormone auxin provides position information for root distal organization. Here, we identify the Arabidopsis thaliana auxin response factors ARF10 and ARF16, targeted by microRNA160 (miR160), as the controller of root cap cell formation. The Pro(35S):MIR160 plants, in which the expression of ARF10 and ARF16 is repressed, and the arf10-2 arf16-2 double mutants display the same root tip defect, with uncontrolled cell division and blocked cell differentiation in the root distal region and show a tumor-like root apex and loss of gravity-sensing. ARF10 and ARF16 play a role in restricting stem cell niche and promoting columella cell differentiation; although functionally redundant, the two ARFs are indispensable for root cap development, and the auxin signal cannot bypass them to initiate columella cell production. In root, auxin and miR160 regulate the expression of ARF10 and ARF16 genes independently, generating a pattern consistent with root cap development. We further demonstrate that miR160-uncoupled production of ARF16 exerts pleiotropic effects on plant phenotypes, and miR160 plays an essential role in regulating Arabidopsis development and growth.

Arabidopsis↗
PubMed2004

Prevalence of apical periodontitis and frequency of root-filled teeth in an adult Spanish population.

AIM: To estimate the prevalence of teeth with apical periodontitis (AP) and root-filled teeth in an adult Spanish population. METHODOLOGY: A total of 180 subjects, aged 37.1 +/- 15.7 years, presenting as new patients to the Faculty of Dentistry, Seville, Spain, were examined. All participants underwent a full-mouth radiographic survey (14 periapical radiographs). The frequency of root canal treatment and the periapical status of all teeth, using the periapical index (PAI) score, were assessed. An intraobserver agreement test on PAI scores produced a Cohen's kappa of 0.77 (substantial agreement). Results were analysed statistically using the Chi-square test. RESULTS: Apical periodontitis in one or several teeth was found in 110 subjects (61.1% prevalence), and 73 (40.6% prevalence) had at least one root-filled tooth. Among subjects with root-filled teeth, 48 (65.8%) had AP affecting at least one root-filled tooth. A total of 4453 teeth were examined, of these 186 (4.2%) had AP. The total number of root-filled teeth was 93 (2.1%), of which 60 (64.5%) had AP. Among non-root filled teeth, only 2.9% had AP. The prevalence of AP in connection with molar teeth was higher (5.5%) than for premolar (4.5%) and anterior teeth (3.2%; P < 0.01). More premolar and molar teeth were root-filled (2.8 and 2.7%, respectively) than anterior teeth (1.3%; P < 0.01). The prevalence of AP increased with age. CONCLUSIONS: The prevalence of AP in root-filled and untreated teeth, and the frequency of root-filled teeth were comparable to those reported in previous similar studies carried out in European countries. The prevalence of root-filled teeth with AP was found to be higher compared to that demonstrated in other epidemiological studies.

Adolescent↗
PubMed2006

Root canal treatment in Denmark is most often carried out in carious vital molar teeth and retreatments are rare.

AIM: To study the reasons given by a representative sample of Danish general dental practitioners (GDPs) for undertaking root canal treatment and, to investigate their confidence in performing root canal treatment on molar teeth. METHODOLOGY: A questionnaire was sent to 600 Danish GDPs randomly selected from the roster of the Danish Dental Association. They were asked to recall various factors about their experience of the last root filling they completed, including the reason for treatment and the pulp diagnosis. Self-assessments on 100-mm visual analogue scales (VAS) were reported concerning the confidence in performing root canal treatment of a molar. End-point definitions were 'very easy' (0) and 'very difficult' (100), respectively. Time reports of molar treatments were given in categorized groups. RESULTS: The most frequent reason for performing root canal treatment was caries within the tooth involved (55%). The majority of treatments involved teeth with vital pulps (54%). Retreatments were carried out in 2% of the cases. The confidence in performing root canal treatment varied but was relatively high, expressed as VAS-values below 50. The creation of an aseptic working field was regarded as the most difficult procedure followed by root canal preparation. Fifty-six percent of the responders stated a time frame of 46-75 min to complete root filling in a molar tooth. CONCLUSIONS: Root canal treatment in Denmark was reported to be undertaken most often because of caries. Treatment was typically performed in molar teeth with vital pulp. Even though apical periodontitis was frequently noted in root filled teeth, retreatments were rare. From a subjective perspective root canal treatment was not considered to be very difficult and was carried out relatively rapidly.

Analysis of Variance↗
PubMed2007

Cross-sectional evaluation of the periapical status and quality of root canal treatment in a selected population of urban Turkish adults.

AIM: To investigate the prevalence of apical periodontitis and the quality of root canal treatment in an adult Turkish population who visited a university dental clinic. SUMMARY: The periapical status of 8863 teeth belonging to 375 patients was evaluated by two pre-calibrated observers from panoramic radiographs. The presence of apical periodontitis, and the prevalence and quality of root fillings were recorded. The relationship between the radiographic quality of root fillings and apical periodontitis was examined by chi-squared statistical-analysis. Of 8863 teeth, 470 (5.3%) had undergone root canal treatment. Forty-seven per cent of all the subjects had at least one root-filled tooth. Periapical radiolucencies were visible on 4.2% of teeth examined, with 53.5% of root-filled teeth presenting apical periodontitis. Ninety-one per cent of root-filled teeth with periapical pathosis were determined to have inadequate root fillings. A statistically significant correlation was found between the quality of root fillings and the presence of apical periodontitis (P < 0.05). KEY LEARNING POINTS: The radiographic quality of root fillings correlated with periapical radiographic signs of pathosis in a selected population of Turkish adults. There is a need to improve the standards of root canal treatment in the study population.

Adolescent↗
PubMed2005

Intrinsic and environmental response pathways that regulate root system architecture.

Root system development is an important agronomic trait. The right architecture in a given environment allows plants to survive periods of water of nutrient deficit, and compete effectively for resources. Root systems also provide an optimal system for studying developmental plasticity, a characteristic feature of plant growth. This review proposes a framework for describing the pathways regulating the development of complex structures such as root systems: intrinsic pathways determine the characteristic architecture of the root system in a given plant species, and define the limits for plasticity in that species. Response pathways co-ordinate environmental cues with development by modulating intrinsic pathways. The current literature describing the regulation of root system development is summarized here within this framework. Regulatory pathways are also organized based on their specific developmental effect in the root system. All the pathways affect lateral root formation, but some specifically target initiation of the lateral root, while others target the development and activation of the lateral root primordium, or the elongation of the lateral root. Finally, we discuss emerging approaches for understanding the regulation of root system architecture.

Cell Cycle↗
PubMed2005

Fine root distribution and persistence under field conditions of three co-occurring Great Basin species of different life form.

Fine roots of an annual grass, a perennial grass and a perennial shrub were examined. Based on life histories and tissue composition, we expected the greatest root persistence for the shrub and shortest for the annual grass. Roots were observed with minirhizotrons over 2 yr for number, length and diameter changes. A Cox proportional hazard regression correlated root persistence with soil water, depth, diameter and date of production. In 2001, grass roots had similar persistence times, but shrub roots had the shortest. In 2002, the annual had the longest median root persistence, the perennial grass intermediate and the perennial shrub had the shortest. All species responded similarly to the magnitude of seasonal precipitation; root numbers increased with favorable soil moisture and disappeared with drying; fewer, thinner roots at greater soil depths were found in the drier year (2001). Root persistence increased with soil moisture, diameter and earlier appearance in the spring. Plasticity in root morphology and placement was influenced by water availability, yet persistence was surprisingly contrary to expectations.

Agropyron↗
PubMed2005

Conventional detection methodology is limiting our ability to understand the roles and functions of fine roots.

We lack a thorough conceptual and functional understanding of fine roots. Studies that have focused on estimating the quantity of fine roots provide evidence that they dominate overall plant root length. We need a standard procedure to quantify root length/biomass that takes proper account of fine roots. Here we investigated the extent to which root length/biomass may be underestimated using conventional methodology, and examined the technical reasons that could explain such underestimation. Our discussion is based on original X-ray-based measurements and on a literature review spanning more than six decades. We present evidence that root-length recovery depends strongly on the observation scale/spatial resolution at which measurements are carried out; and that observation scales/resolutions adequate for fine root detection have an adverse impact on the processing times required to obtain precise estimates. We conclude that fine roots are the major component of root systems of most (if not all) annual and perennial plants. Hence plant root systems could be much longer, and probably include more biomass, than is widely accepted.

Brassica napus↗
PubMed2006

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.

Acid Phosphatase↗
PubMed2007

Impact of temperature on the relationship between respiration and nitrogen concentration in roots: an analysis of scaling relationships, Q10 values and thermal acclimation ratios.

* The impact of nitrogen (N) supply on the temperature response of root respiratory O(2) uptake (R) was assessed in several herbaceous species grown in solution culture. Warm-grown (25 : 20 degrees C, day:night) plants differing in root N concentration were shifted to 13 : 8 degrees C for 7 d to cold-acclimate. * Log-log plots of root R vs root N concentration both showed that R increased with increasing tissue N concentration, irrespective of the growth temperature. Although the regression slopes of the log-log plots did not differ between the warm-grown and cold-acclimated plants, cold-acclimated plants did exhibit a higher y-axis intercept than their warm-grown counterparts. This suggests that cold acclimation of root R is not entirely dependent on cold-induced increases in tissue N concentration and that scaling relationships (i.e. regression equations fitted to the log-log plots) between root R and N concentration are not fixed. * No systematic differences were found in the short-term Q(10) (proportional change in R per 10 degrees C change in temperature), or degree of cold acclimation (as measured by the proportional difference between warm- and cold-acclimated roots) among roots differing in root N concentration. The temperature response of root R is therefore insensitive to tissue N concentration. * The insensitivity of Q(10) values and acclimation to tissue N concentration raises the possibility that root R and its temperature sensitivity can be predicted for a range of N supply scenarios.

Acclimatization↗
PubMed1985

Healing following reimplantation of teeth subjected to root planing and citric acid treatment.

The aim of the present study was to examine the effect of citric acid treatment on periodontal healing around teeth which were extracted, root planed and then reimplanted. Maxillary incisors and mandibular incisors, premolars and molars of 5 monkeys were used. The teeth were divided into 3 experimental groups. In 1 group, the teeth were extracted and immediately reimplanted into their own sockets. In a 2nd group, the teeth were extracted, root planed to a level corresponding to 50-75% of the root length and then reimplanted. In the 3rd group, periodontal breakdown extending to 50-75% of the root length was first induced by placing orthodontic elastic ligatures around the teeth. They were then extracted and root planed and transplanted into the sockets of the contralateral, periodontally healthy teeth which had just been extracted. Half the number of the teeth of groups 2 and 3 were treated with citric acid before reimplantation or transplantation. The animals were sacrificed after 6 months of healing. The jaws were removed and histological specimens prepared for microscopic examination. With the exception of a limited coronal regrowth of new cementum in the apical part of the planed portion of a few roots, connective tissue attachment failed to reform on most root surfaces deprived of their periodontal ligament tissue. Healing was most frequently characterized by root resorption and ankylosis. These were the most predominant features of healing both on root surfaces which had been deprived of the ligament tissue by mechanical means or during a course of experimentally-induced periodontal disease, and occurred in citric acid as well as non-citric acid treated roots. The findings imply that the result of healing following tooth reimplantation or transplantation is determined by the type of cells that repopulate the wound area adjacent to the denuded root surface.

Animals↗