The expansion of maize root-cap mucilage during hydration. 3. Changes in water potential and water content

Root-cap mucilage from aerial nodal roots of maize has been found to have water potential values of −11 MPa or lower when air dried. The value approaches 0 MPa within 2 min of hydration in distilled water. In this time the expanding gel absorbs only about 0.3% of the water content of fully expanded mucilage. It is concluded that the root-cap mucilage per se has almost no capacity to retain water in the rhizosphere. Any function that it may play in the slowing of root desiccation would be indirect. For example, mucilage might decrease pore size between and within soil aggregates by pulling the particles together in a cycle of nocturnal efflux of water from the root surface, and diumal dyring during transpiration.     

Effect of maize canopy and water repellency on moisture patterns in a Dutch black plaggen soil

Man-made raised sandy soils in the Netherlands are classified as brown or black plaggen soils. When dry, the brown soils are wettable, but the black soils are water repellent. For one growing season, transects were sampled in a maize cropped black plaggen soil at the Heino experimental farm. Due to interception and stemflow, water was concentrated near the roots of the maize. Between the maize rows, higher soil water contents were found in microdepressions, due to rainwater dripping to the ground from overhanging leaves. Redistribution of soil water from wet to dry areas was restricted by the water repellency of the dry sand. As a consequence, there was a distinct variation in soil moisture content. These irregular wetting patterns did not induce preferential downward flow, but widened over time; because the dry, water repellent subsoil impeded and resisted infiltration into the deeper subsoil.     

根际土壤通透性对玉米水分和养分吸收的影响

为改善作物根系生长微环境,探索根际土壤通透性对作物水分和养分吸收的影响,在3个灌溉水平下（灌水量分别为每次600、400和200 ml）,采用盆栽玉米不通气、每隔2 d通1次气、每隔4 d通1次气等处理方法,研究了根际土壤通透性对盆栽玉米生理指标及水分和养分吸收的影响.结果表明：在相同灌水条件下,通气处理促进了玉米株高、叶面积的增长,提高了叶绿素含量;促进了玉米对土壤养分的吸收;显著提高了玉米的根系活力,灌水量为每次600 ml时,拔节期每隔4 d通气处理的玉米根系活力最大（8.24 mg·g^-1·h^-1）,较不通气处理（4.94 mg·g^-1·h^-1）提高了66.7%;根际土壤通气处理对盆栽玉米蒸腾的影响不显著,说明根际通气可提高玉米对水分与养分的吸收利用效率,促进植株生长发育.     

The dynamics of protons,aluminium, and calcium in the rhizosphere of maize cultivated in tropical acid soils: experimental study and modelling

The goals of this work were to understand the dynamics of H⁺, Al and Ca in the rhizosphere of maize cultivated in tropical acid soils, and to evaluate the contribution of the dissolution kinetics of the Al-hydroxides to Al dynamics. The study of the dissolution kinetics was based on a comparison between experimental and simulated data, using a model of the chemical processes in the rhizosphere. Two Oxisols, pH 5.1 and 4.6, and one Ultisol, pH 5.2, were studied. An Al-tolerant maize variety (Zea mays L.) was grown for 14 days on a 3-mm thick soil layer. The composition of the soil and the soil solution, together with the concentration of Al in the roots, were determined throughout the experiment. The results showed that root growth (i) decreased the soil solution pH, up to one pH unit, (ii) increased Al concentration in the soil solution, (iii) increased exchangeable Al, and (iv) decreased exchangeable Ca. Soil solution pH, exchangeable Al, and exchangeable Ca were closely linked. Exchangeable Al increased 1.5 – 3.0 times, due to the dissolution of easily mobilised Al components. In addition, Al accumulation in roots depended mainly on Al in the soil solution. Modelling the interactions between H⁺, Al, and Ca proved that the main factor determining Al in the soil solution was the kinetic reactivity of the easily mobilised Al components. These components, probably poorly crystallised Al-hydroxides, are key players in the functioning of the rhizosphere in tropical acid soils.     

Root:soil adhesion in the maize rhizosphere: the rheological approach

This study was designed to investigate the strength of attachment of plant seedling roots to the soil in which they were grown. The study also assessed the effects of differing soil textures and differing soil matric potentials upon the strength of the root:soil attachment. A device for growing roots upon a soil surface was designed, and was used to produce roots which were attached to the soil. In order to quantify root:soil adhesion, roots of maize seedlings, grown on the soil surface, were subsequently peeled off using a universal test machine, in conjunction with simultaneous time-lapse video observation. To clarify the partitioning of energy in the root:soil peeling test, separate mechanical tests on roots, and on two adherent remoulded topsoil balls were also carried out. The seedling root was characterised by a low bending stiffness. The energy stored in bending was negligible, compared to the root:soil adhesion energy. The mechanical properties of two adherent remoulded topsoil balls were a decrease of the soil:soil adhesion energy as the soil:soil plastic energy increased. These two parameters were therefore interdependent. Using a video-camera system, it was possible to separate the different processes occurring during the root:soil peeling test, in particular, the seed:soil adhesion and the root:soil soil adhesion. An interpretation of the complex and variable force:displacement curves was thus possible, enabling calculation of the root:soil interfacial rupture energy. At a given suction (10 kPa), the results of the peeling test showed a clear soil texture effect on the value of the root:soil interfacial rupture energy. In contrast, for the same silty topsoil, the effect of the soil water suction on the value of the interfacial rupture energy was very moderate. The root:soil interfacial rupture energy was controlled mainly by a product of microscopic soil specific surface area and the macroscopic contact surface area between the root and the soil. Biological and physical interactions contributing to root:soil adhesion such as root:soil interlocking mechanics were also analysed and discussed. This revised version was published online in June 2006 with corrections to the Cover Date.     

Manganese reduction in the rhizosphere of mycorrhizal and nonmycorrhizal maize

The influence of rhizosphere microorganisms and vesicular-arbuscular (VA) mycorrhiza on manganese (Mn) uptake in maize (Zea mays L. cv. Tau) plants was studied in pot experiments under controlled environmental conditions. The plants were grown for 7 weeks in sterilized calcareous soil in pots having separate compartments for growth of roots and of VA mycorrhizal fungal hyphae. The soil was left either uninoculated (control) or prior to planting was inoculated with rhizosphere microorganisms only (MO-VA) or with rhizosphere microorganisms together with a VA mycorrhizal fungus [Glomus mosseae (Nicol and Gerd.) Gerdemann and Trappe] (MO+VA). Mycorrhiza treatment did not affect shoot dry weight, but root dry weight was slightly inhibited in the MO+VA and MO-VA treatments compared with the uninoculated control. Concentrations of Mn in shoots decreased in the order MO-VA > MO+VA > control. In the rhizosphere soil, the total microbial population was higher in mycorrhizal (MO+VA) than nonmycorrhizal (MO-VA) treatments, but the proportion of Mn-reducing microbial populations was fivefold higher in the nonmycorrhizal treatment, suggesting substantial qualitative changes in rhizosphere microbial populations upon root infection with the mycorrhizal fungi. The most important microbial group taking part in the reduction of Mn was fluorescent Pseudomonas. Mycorrhizal treatment decreased not only the number of Mn reducers but also the release of Mn-solubilizing root exudates, which were collected by percolation from maize plants cultivated in plastic tubes filled with gravel quartz sand. Compared with mycorrhizal plants, the root exudates of nonmycorrhizal plants had two fold higher capacity for reduction of Mn. Therefore, changes in both rhizosphere microbial population and root exudation are probably responsible for the lower acquisition of Mn in mycorrhizal plants.     

Influence of maize mucilage on the diversity and activity of the denitrifying community

In order to understand the effect of the maize rhizosphere on denitrification, the diversity and the activity of the denitrifying community were studied in soil amended with maize mucilage. Diversity of the denitrifying community was investigated by polymerase chain reaction (PCR) amplification of total community DNA extracted from soils using gene fragments, encoding the nitrate reductase (narG) and the nitrous oxide reductase (nosZ), as molecular markers. To assess the underlying diversity, PCR products were cloned and 10 gene libraries were obtained for each targeted gene. Libraries containing 738 and 713 narG and nosZ clones, respectively, were screened by restriction fragment analysis, and grouped based on their RFLP (restriction fragment length polymorphism) patterns. In all, 117 and 171 different clone families have been identified for narG and nosZ and representatives of RFLP families containing at least two clones were sequenced. Rarefaction curves of both genes did not reach a clear saturation, indicating that analysis of an increasing number of clones would have revealed further diversity. Recovered NarG sequences were related to NarG from Actinomycetales and from Proteobacteria but most of them are not related to NarG from known bacteria. In contrast, most of the NosZ sequences were related to NosZ from alpha, beta, and gammaProteobacteria. Denitrifying activity was monitored by incubating the control and amended soils anaerobically in presence of acetylene. The N2O production rates revealed denitrifying activity to be greater in amended soil than in control soil. Altogether, our results revealed that mucilage addition to the soil results in a strong impact on the activity of the denitrifying community and minor changes on its diversity.     

Microbial community composition and functional diversity in the rhizosphere of maize

This study investigates the small-scale stratification of bacterial community composition and functional diversity in the rhizosphere of maize. Maize seedlings were grown in a microcosm with a horizontal mesh (53 M) creating a planar root mat and rhizosphere soil. An unplanted microcosm served as control. Thin slices of soil were cut at different distances from the mesh surface (0.2–5.0 mm) and analysed for bacterial community composition by PCR-DGGE (polymerase chain reaction-denaturing gradient gel electrophoresis) of 16S rDNA and tested for activities of different enzymes involved in C, N, P and S cycling. Bacterial community composition and microbial functional diversity were affected by the presence of the maize roots. The bacterial composition showed a clear gradient up to 2.2 mm from the root surface, while no such gradient was observed in the unplanted pot. Invertase and phosphatase activities were higher in the close vicinity of maize roots (0.2–0.8 mm), whereas xylanase activity was unaffected. This study shows that the changes in bacterial community composition and functional diversity induced by roots may extend several millimetres into the soil.     

Dynamics of fungal communities in bulk and maize rhizosphere soil in the tropics

The fungal population dynamics in soil and in the rhizospheres of two maize cultivars grown in tropical soils were studied by a cultivation-independent analysis of directly extracted DNA to provide baseline data. Soil and rhizosphere samples were taken from six plots 20, 40, and 90 days after planting in two consecutive years. A 1.65-kb fragment of the 18S ribosomal DNA (rDNA) amplified from the total community DNA was analyzed by denaturing gradient gel electrophoresis (DGGE) and by cloning and sequencing. A rhizosphere effect was observed for fungal populations at all stages of plant development. In addition, pronounced changes in the composition of fungal communities during plant growth development were found by DGGE. Similar types of fingerprints were observed in two consecutive growth periods. No major differences were detected in the fungal patterns of the two cultivars. Direct cloning of 18S rDNA fragments amplified from soil or rhizosphere DNA resulted in 75 clones matching 12 dominant DGGE bands. The clones were characterized by their HinfI restriction patterns, and 39 different clones representing each group of restriction patterns were sequenced. The cloning and sequencing approach provided information on the phylogeny of dominant amplifiable fungal populations and allowed us to determine a number of fungal phylotypes that contribute to each of the dominant DGGE bands. Based on the sequence similarity of the 18S rDNA fragment with existing fungal isolates in the database, it was shown that the rhizospheres of young maize plants seemed to select the Ascomycetes order Pleosporales, while different members of the Ascomycetes and basidiomycetic yeast were detected in the rhizospheres of senescent maize plants.     

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.     

Dynamics of soil water content in the rhizosphere

Water flow from soil to plants depends on the properties of the soil next to roots, the rhizosphere. Although several studies showed that the rhizosphere has different properties than the bulk soil, effects of the rhizosphere on root water uptake are commonly neglected. To investigate the rhizosphere’s properties we used neutron radiography to image water content distributions in soil samples planted with lupins during drying and subsequent rewetting. During drying, the water content in the rhizosphere was 0.05 larger than in the bulk soil. Immediately after rewetting, the picture reversed and the rhizosphere remained markedly dry. During the following days the water content of the rhizosphere increased and after 60 h it exceeded that of the bulk soil. The rhizosphere’s thickness was approximately 1.5 mm. Based on the observed dynamics, we derived the distinct, hysteretic and time-dependent water retention curve of the rhizosphere. Our hypothesis is that the rhizosphere’s water retention curve was determined by mucilage exuded by roots. The rhizosphere properties reduce water depletion around roots and weaken the drop of water potential towards roots, therefore favoring water uptake under dry conditions, as demonstrated by means of analytical calculation of water flow to a single root.     

不同灌溉方式下土玉米根际硝态氮的分布

实验着重研究分根条件下常规灌溉、交替灌溉和固定灌溉玉米苗期根际硝态氮的分布, 研究结果表明不同灌水方式下,玉米根际硝态氮的分布不同.在这3种灌水方式的湿润区,NO-3-N的累积趋势为交替灌水＞固定灌水＞常规灌水.     

Kinetic modelling of phototropism in maize coleoptiles

Blue-light-induced phototropism of maize (Zea mays L.) coleoptiles was studied with a view to kinetic models. Red-light-grown plants were used to eliminate complication arising from the activation by blue light of phytochrome-mediated phototropism. In the first part, mathematical models were developed to explain the phototropic fluence-response data, which were obtained for the responses induced by a single unilateral pulse (30 s) and those induced by a unilateral pulse (30 s) given immediately after a bilateral pulse (30 s, fixed fluences). These data showed bell-shaped fluence-response curves, characteristic of first positive curvature. Modelling began with the assumptions that the light gradient plays a fundamental role in phototropism and that the magnitude of the response is determined by the gradient, or the concentration difference, in a photoproduct between the irradiated and the shaded sides of the tissue. Minimal mathematical models were then derived, by defining chemical kinetics of the photoreaction and introducing the minimum of parameters needed to correlate the incident fluencerate to the functional fluence-rates within the tissue, the functional fluence-rate to the rate constant of the photoreaction, and the photoproduct concentration difference to the curvature response. The models were tested using a curve-fitting computer program. The model obtained by assigning first-order kinetics to the photoreaction failed to explain the fluence-response data, whereas application of second-order kinetics led to a successful fit of the model to the data. In the second part, temporal aspects of the photosystem were examined. Experimental results showed that a high-fluence bilateral pulse eliminated the bell-shaped fluence-response curve for an immediate unilateral pulse, and that the curve gradually reappeared as the time for unilateral stimulation elapsed after the bilateral pulse. The model based on a second-order photoreaction could be extended to explain the results, with assumed changes in two components: the concentration of the reactant for the photoproduct, and the light-sensitivity of the reaction. The reactant concentration, computed with the curvefitting program, showed a gradual increase from zero to a saturation level. This increase was then modelled in terms of regeneration of the reactant from the photoproduct, with an estimated first-order rate constant of about 0.001·s^-1. The computed value for the constant reflecting the light-sensitivity showed a sharp decline after the high-fluence pulse, followed by a gradual return to the initial level. From these analytical results, the appearance of second positive curvature was predicted.Abbreviations FPC first positive curvature - SPC second positive curvature CIW-DPB publication No. 884     

Diversity and distribution of Burkholderia cepacia complex in the rhizosphere of rice and maize

A survey of Burkholderia cepacia complex (Bcc) species was conducted in agricultural fields within Hangzhou, China. Out of the 251 bacterial isolates recovered on the selective media from the rhizosphere of rice and maize, 112 of them were assigned to Bcc by PCR assays. The species composition of the Bcc isolates was analyzed by a combination of recA-restriction fragment length polymorphism assays, species-specific PCR tests and recA gene sequencing. The results revealed that the majority belong to B. cepacia, Burkholderia cenocepacia recA lineage IIIB, Burkholderia vietnamiensis and Burkholderia pyrrocinia. Burkholderia cenocepacia and B. vietnamiensis dominated the rhizosphere of maize and rice, respectively, indicating that species composition and abundance of Bcc may vary dramatically in different crop rhizospheres. In addition, one isolate (R456) formed a single discrete cluster within the phylogenetic analysis of the Bcc recA gene, and it may belong to a new genomovar.     

Possible processes releasing nonexchangeable potassium from the rhizosphere of maize

The source and the releasing processes of nonexchangeable K from the rhizosphere were evaluated by using a 0.01 M HCl sequential extraction that enables the detection of subtle depletion of nonexchangeable K in the rhizosphere. Rhizobox experiments were conducted in which maize (Zea mays L.) plants were grown on different K sources (non-allophanic Andosol, Fluvisol, biotite and orthoclase) for 17 days. Nonexchangeable K decreased significantly in the rhizosphere of Andosol, Fluvisol and biotite, but not of orthoclase. The width of depletion was about 0–1 mm from the root-accumulating compartment regardless of the K sources, and was much less than that of exchangeable (5–10 mm) and water-soluble (50 mm) K. Rhizosphere pHs were above 4.5 in any treatment. These results suggested that the main source of nonexchangeable K for maize was interlayer K in 2:1 type phyllosilicate, and that the releasing process involved was cation exchange of the K rather than mineral dissolution by protons. For the exchange of interlayer K, a decrease of solution K⁺ below a certain threshold is known as a prerequisite. But the concentration of solution K⁺ at the root compartment of Andosol or Fluvisol, estimated to be more than 100 M, was relatively higher than the known thresholds. Moreover, the significant release of nonexchangeable K from biotite occurred only at the root compartment where marked depletion of solution K⁺ was not observed. We therefore suggest that the release of interlayer K from the rhizosphere can occur even without a marked depletion of solution K⁺ through the following processes; (1) accumulation of cations such as Ca²⁺, Mg²⁺ or Na⁺ in the rhizosphere, (2) their adsorption on 2:1 type clay minerals near the edge of interlayer but inaccessible and nonexchangeable by NH₄ ⁺, (3) concomitant removal of the NH₄ ⁺-nonexchangable K even above the known thresholds of solution K⁺, which is followed by the expansion of the interlayer space, and (4) further removal of the deeper K by repetition of the above processes.     

Burkholderia cepacia complex: distribution of genomovars among isolates from the maize rhizosphere in Italy

Burkholderia cepacia is a 'complex' in which seven genomic species or genomovars have so far been identified. It appears that all seven B. cepacia genomovars are capable of causing infections in vulnerable persons; in particular, the importance of Burkholderia multivorans (genomovar II) and B. cepacia genomovar III among cystic fibrosis isolates, especially epidemic ones, has been emphasized. In order to acquire a better comprehension of the genomovar composition of environmental populations of B. cepacia, 120 strains were isolated from the rhizosphere of maize plants cultivated in fields located in northern, central and southern Italy. The identification of the different genomovars was accomplished by a combination of molecular polymerase chain reaction (PCR)-based techniques, such as restriction fragment length polymorphism (RFLP) analysis of 16S rDNA (ARDRA), genomovar-specific PCR tests and RFLP analyses based on polymorphisms in the recA gene whole-cell protein electrophoresis. ARDRA analysis allowed us to distinguish between all B. cepacia genomovars except B. cepacia genomovar I, B. cepacia genomovar III and Burkholderia ambifaria (genomovar VII). The latter genomovars were differentiated by means of recA PCR tests and RFLP analyses. Among the rhizospheric isolates of B. cepacia, we found only B. cepacia genomovar I, B. cepacia genomovar III, Burkholderia vietnamiensis (genomovar V) and B. ambifaria. B. cepacia genomovars I and III and B. ambifaria were recovered from all three fields, whereas B. vietnamiensis was detected only in the population isolated from the field located in central Italy. Among strains isolated from northern and southern Italy, the most abundant genomovars were B. ambifaria and B. cepacia genomovar III respectively; in contrast, the population isolated in central Italy showed an even distribution of strains among genomovars. These results indicate that it is not possible to differentiate clinical and environmental strains, or pathogenic and non-pathogenic strains, of the B. cepacia complex simply on the basis of genomovar status, and that the environment may serve as a reservoir for B. cepacia genomovar III infections in vulnerable humans.     

不同灌溉方式下(土娄)土玉米根际硝态氮的分布

实验着重研究分根条件下常规灌溉、交替灌溉和固定灌溉玉米苗期根际硝态氮的分布, 研究结果表明:不同灌水方式下,玉米根际硝态氮的分布不同.在这3种灌水方式的湿润区,NO-3-N的累积趋势为:交替灌水＞固定灌水＞常规灌水.