Plant and algal cell walls: diversity and functionality

Background

Although plants and many algae (e.g. the Phaeophyceae, brown, and Rhodophyceae, red) are only very distantly related they are united in their possession of carbohydrate-rich cell walls, which are of integral importance being involved in many physiological processes. Furthermore, wall components have applications within food, fuel, pharmaceuticals, fibres (e.g. for textiles and paper) and building materials and have long been an active topic of research. As shown in the 27 papers in this Special Issue, as the major deposit of photosynthetically fixed carbon, and therefore energy investment, cell walls are of undisputed importance to the organisms that possess them, the photosynthetic eukaryotes (plants and algae). The complexities of cell wall components along with their interactions with the biotic and abiotic environment are becoming increasingly revealed.

Scope

The importance of plant and algal cell walls and their individual components to the function and survival of the organism, and for a number of industrial applications, are illustrated by the breadth of topics covered in this issue, which includes papers concentrating on various plants and algae, developmental stages, organs, cell wall components, and techniques. Although we acknowledge that there are many alternative ways in which the papers could be categorized (and many would fit within several topics), we have organized them as follows: (1) cell wall biosynthesis and remodelling, (2) cell wall diversity, and (3) application of new technologies to cell walls. Finally, we will consider future directions within plant cell wall research. Expansion of the industrial uses of cell walls and potentially novel uses of cell wall components are both avenues likely to direct future research activities. Fundamentally, it is the continued progression from characterization (structure, metabolism, properties and localization) of individual cell wall components through to defining their roles in almost every aspect of plant and algal physiology that will present many of the major challenges in future cell wall research.     

Some properties of the walls of metaxylem vessels of maize roots,including tests of the wettability of their lumenal wall surfaces

Background and Aims

Since the proposal of the cohesion theory there has been a paradox that the lumenal surface of vessels is rich in hydrophobic lignin, while tension in the rising sap requires adhesion to a hydrophilic surface. This study sought to characterize the strength of that adhesion in maize (Zea mays), the wettability of the vessel surface, and to reconcile this with its histochemical and physical nature.

Methods

Wettability was assessed by emptying the maize root vessels of sap, perfusing them with either water or oil, and examining the adhesion (as revealed by contact angles) of the two liquids to vessel walls by cryo-scanning electron microscopy. The phobicity of the lumenal surface was also assessed histochemically with hydrophilic and hydrophobic probes.

Key Results

Pit borders in the lumen-facing vessel wall surface were wetted by both sap/water and oil. The attraction for oil was weaker: water could replace oil but not vice versa. Pit apertures repelled oil and were strongly stained by hydrophilic probes. Pit chambers were probably hydrophilic. Oil never entered the pits. When vessels were emptied and cryo-fixed immediately, pit chambers facing away from the vessels were always sap-filled. Pit chambers facing vessel lumens were either sap- or gas-filled. Sap from adjoining tracheary elements entering empty vessels accumulated on the lumenal surface in hemispherical drops, which spread out with decreasing contact angles to fill the lumen.

Conclusions

The vessel lumenal surface has a dual nature, namely a mosaic of hydrophilic and hydrophobic patches at the micrometre scale, with hydrophilic predominating. A key role is shown, for the first time, of overarching borders of pits in determining the dual nature of the surface. In gas-filled (embolized) vessels they are hydrophobic. When wetted by sap (vessels refilling or full) they are hydrophilic. A hypothesis is proposed to explain the switch between the two states.     

Auxin transport in maize roots in response to localized nitrate supply

Background and Aims

Roots typically respond to localized nitrate by enhancing lateral-root growth. Polar auxin transport has important roles in lateral-root formation and growth; however, it is a matter of debate whether or how auxin plays a role in the localized response of lateral roots to nitrate.

Methods

Treating maize (Zea mays) in a split-root system, auxin levels were quantified directly and polar transport was assayed by the movement of [³H]IAA. The effects of exogenous auxin and polar auxin transport inhibitors were also examined.

Key Results

Auxin levels in roots decreased more in the nitrate-fed compartment than in the nitrate-free compartment and nitrate treatment appeared to inhibit shoot-to-root auxin transport. However, exogenous application of IAA only partially reduced the stimulatory effect of localized nitrate, and auxin level in the roots was similarly reduced by local applications of ammonium that did not stimulate lateral-root growth.

Conclusions

It is concluded that local applications of nitrate reduced shoot-to-root auxin transport and decreased auxin concentration in roots to a level more suitable for lateral-root growth. However, alteration of root auxin level alone is not sufficient to stimulate lateral-root growth.     

Spatial distributions of tissue expansion and cell division rates are related to irradiance and to sugar content in the growing zone of maize roots

We have investigated the way in which the radiation absorbed by leaves affects the rate of elongation of maize ( Zea mays L.) roots. In five repeated growth chamber experiments, plants previously grown at a photon irradiance of 23 mol m^–2 d^–1 received either 7 or 34 mol m^–2 d^–1 from day 10 to day 20 after germination. The elongation rate of primary roots steadily decreased for 4 d after reduction in irradiance and then stabilized at 60% of that in plants at high irradiance. The elongating zone was slightly shorter after 2 d at low irradiance, and was further reduced after 8 d. The concentrations of sucrose and glucose in the elongating zone were greatly decreased after 2 d at low irradiance and the gradient of both sugars was suppressed. The longer period at low irradiance affected neither sugar content nor gradient. In the same way, cell production rate was reduced after 2 d at low irradiance and was not appreciably decreased afterwards. The root zone with cell division was shorter in plants at low irradiance, but cell division rate remained nearly constant temporally and spatially, and was unaffected by the irradiance treatment. Our results suggest that primary events after a reduction in irradiance were a change in cell flux and sugar content in the elongating zone. Change in elongation rate was slower and probably the result of a time-related developmental effect, which may be related to the change in cell production.     

Early local differentiation of the cell wall matrix defines the contact sites in lobed mesophyll cells of Zea mays

Background and Aims

The morphogenesis of lobed mesophyll cells (MCs) is highly controlled and coupled with intercellular space formation. Cortical microtubule rings define the number and the position of MC isthmi. This work investigated early events of MC morphogenesis, especially the mechanism defining the position of contacts between MCs. The distributions of plasmodesmata, the hemicelluloses callose and (1 → 3,1 → 4)-β-d-glucans (MLGs) and the pectin epitopes recognized by the 2F4, JIM5, JIM7 and LM6 antibodies were studied in the cell walls of Zea mays MCs.

Methods

Matrix cell wall polysaccharides were immunolocalized in hand-made sections and in sections of material embedded in LR White resin. Callose was also localized using aniline blue in hand-made sections. Plasmodesmata distribution was examined by transmission electron microscopy.

Results

Before reorganization of the dispersed cortical microtubules into microtubule rings, particular bands of the longitudinal MC walls, where the MC contacts will form, locally differentiate by selective (1) deposition of callose and the pectin epitopes recognized by the 2F4, LM6, JIM5 and JIM7 antibodies, (2) degradation of MLGs and (3) formation of secondary plasmodesmata clusterings. This cell wall matrix differentiation persists in cell contacts of mature MCs. Simultaneously, the wall bands between those of future cell contacts differentiate with (1) deposition of local cell wall thickenings including cellulose microfibrils, (2) preferential presence of MLGs, (3) absence of callose and (4) transient presence of the pectins identified by the JIM5 and JIM7 antibodies. The wall areas between cell contacts expand determinately to form the cell isthmi and the cell lobes.

Conclusions

The morphogenesis of lobed MCs is characterized by the early patterned differentiation of two distinct cell wall subdomains, defining the sites of the future MC contacts and of the future MC isthmi respectively. This patterned cell wall differentiation precedes cortical microtubule reorganization and may define microtubule ring disposition.     

Cortical microtubule patterning in roots of Arabidopsis thaliana primary cell wall mutants reveals the bidirectional interplay with cell expansion

Cell elongation requires directional deposition of cellulose microfibrils regulated by transverse cortical microtubules. Microtubules respond differentially to suppression of cell elongation along the developmental zones of Arabidopsis thaliana root apex. Cortical microtubule orientation is particularly affected in the fast elongation zone but not in the meristematic or transition zones of thanatos and pom2–4 cellulose-deficient mutants of Arabidopsis thaliana. Here, we report that a uniform phenotype is established among the primary cell wall mutants, as cortical microtubules of root epidermal cells of rsw1 and prc1 mutants exhibit the same pattern described in thanatos and pom2–4. Whether cortical microtubules assume transverse orientation or not is determined by the demand for cellulose synthesis, according to each root zone''s expansion rate. It is suggested that cessation of cell expansion may provide a biophysical signal resulting in microtubule reorientation.     

Rate of differentiation and emergence of nodal maize roots

The timing of root production is one of the parameters required for modelling the root system architecture. The objectives of this study are (1) to describe the rate of appearance of adventitious root primordia of maize and their rate of emergence out of the stem; (2) to test equations for the prediction of the rank of the phytomer on which root emergence occurs, in a wide range of field situations.Maize, cultivar Dea, was grown in controlled conditions and in the field in 1987, 1988, 1989 and 1991. Plants were regularly sampled and the following data were recorded: foliar stage, number of root primordia and number of emerged roots per phytomer. Root primordia were counted in transverse thin sections in the stem.At a single plant level, root primordia differentiation occurred sequentially on the successive phytomers, with no overlapping between two phytomers. The same was true for root emergence. Roots belonging to the same phytomer emerged at approximately the same time.At a plant population level, there was a linear relationship between the rank of the phytomer on which root primordia were differentiated and cumulated degree-days after sowing. A linear relationship was also observed between the rank of the phytomer on which roots were emerging and cumulated degree-days or foliar stage. In the range of field situations tested (several years, sowing dates and planting densities), both equations gave an accurate prediction of the timing of root emergence during the plant cycle.     

Spatial and temporal profiles of cytokinin biosynthesis and accumulation in developing caryopses of maize

Background and Aims

Cytokinins are a major group of plant hormones and are associated with various developmental processes. Developing caryopses of maize have high levels of cytokinins, but little is known about their spatial and temporal distribution. The localization and quantification of cytokinins was investigated in maize (Zea mays) caryopsis from 0 to 28 d after pollination together with the expression and localization of isopentenyltransferase ZmIPT1 involved in cytokinin biosynthesis and ZmCNGT, the gene putatively involved in N9-glucosylation.

Methods

Biochemical, cellular and molecular approaches resolved the overall cytokinin profiles, and several gene expression assays were used for two critical genes to assess cytokinin cell-specific biosynthesis and conversion to the biologically inactive form. Cytokinins were immunolocalized for the first time in maize caryopses.

Key Results

During the period 0–28 d after pollination (DAP): (1) large quantities of cytokinins were detected in the maternal pedicel region relative to the filial tissues during the early stages after fertilization; (2) unpollinated ovules did not accumulate cytokinins; (3) the maternal nucellar region showed little or no cytokinin signal; (4) the highest cytokinin concentrations in filial endosperm and embryo were detected at 12 DAP, predominantly zeatin riboside and zeatin-9-glucoside, respectively; and (5) a strong cytokinin immuno-signal was detected in specific cell types in the pedicel, endosperm and embryo.

Conclusions

The cytokinins of developing maize caryopsis may originate from both local syntheses as well as by transport. High levels of fertilization-dependent cytokinins in the pedicel suggest filial control on metabolism in the maternal tissue; they may also trigger developmental programmed cell death in the pedicel.     

An integrated method for quantifying root architecture of field-grown maize

Background and Aims

A number of techniques have recently been developed for studying the root system architecture (RSA) of seedlings grown in various media. In contrast, methods for sampling and analysis of the RSA of field-grown plants, particularly for details of the lateral root components, are generally inadequate.

Methods

An integrated methodology was developed that includes a custom-made root-core sampling system for extracting intact root systems of individual maize plants, a combination of proprietary software and a novel program used for collecting individual RSA information, and software for visualizing the measured individual nodal root architecture.

Key Results

Example experiments show that large root cores can be sampled, and topological and geometrical structure of field-grown maize root systems can be quantified and reconstructed using this method. Second- and higher order laterals are found to contribute substantially to total root number and length. The length of laterals of distinct orders varies significantly. Abundant higher order laterals can arise from a single first-order lateral, and they concentrate in the proximal axile branching zone.

Conclusions

The new method allows more meaningful sampling than conventional methods because of its easily opened, wide corer and sampling machinery, and effective analysis of RSA using the software. This provides a novel technique for quantifying RSA of field-grown maize and also provides a unique evaluation of the contribution of lateral roots. The method also offers valuable potential for parameterization of root architectural models.     

Cell wall-bound trans- and cis-ferulic acids in growing maize roots

The levels of cell wall-bound trans - and cis -ferulic acids in roots of dark grown Zea mays cv. LG11 plants were measured. They were quantified after alkaline hydrolysis of purified cell walls by reversed phase HPLC using trans -cinnamic acid as internal standard. The total amount of ferulic acid ( trans - and cis -ferulic acid) in the root base was 3–4 times higher than in the root tip. Cis -ferulic acid represented between 2% (tip) and 18% (base) of the total ferulic acid content. The total content of trans - and cis -ferulic acids was approximately the same in the stele and the cortex, but the level of cis -ferulic acid in the stele was 5–6 times higher than in the cortex. Trans - and cis -ferulic acid levels as well as the percentage of cis -ferulic acid in the elongation zone were steady between 48 and 96 h after the beginning of germination. Slowly growing roots contained more wall-bound ferulic acids, particularly cis -ferulic acid, than fast growing roots. This relationship was found in the differentiation zone but not in the elongation zone. The importance of cell wall-bound trans - and cis -ferulic acids is discussed in the context of root growth and differentiation.     

Lateral root development in the maize (Zea mays) lateral rootless1 mutant

Background and Aims

The maize lrt1 (lateral rootless1) mutant is impaired in its development of lateral roots during early post-embryonic development. The aim of this study was to characterize, in detail, the influences that the mutation exerts on lateral root initiation and the subsequent developments, as well as to describe the behaviour of the entire plant under variable environmental conditions.

Methods

Mutant lrt1 plants were cultivated under different conditions of hydroponics, and in between sheets of moist paper. Cleared whole mounts and anatomical sections were used in combination with both selected staining procedures and histochemical tests to follow root development. Root surface permeability tests and the biochemical quantification of lignin were performed to complement the structural data.

Key Results

The data presented suggest a redefinition of lrt1 function in lateral roots as a promoter of later development; however, neither the complete absence of lateral roots nor the frequency of their initiation is linked to lrt1 function. The developmental effects of lrt1 are under strong environmental influences. Mutant primordia are affected in structure, growth and emergence; and the majority of primordia terminate their growth during this last step, or shortly thereafter. The lateral roots are impaired in the maintenance of the root apical meristem. The primary root shows disturbances in the organization of both epidermal and subepidermal layers. The lrt1-related cell-wall modifications include: lignification in peripheral layers, the deposition of polyphenolic substances and a higher activity of peroxidase.

Conclusions

The present study provides novel insights into the function of the lrt1 gene in root system development. The lrt1 gene participates in the spatial distribution of initiation, but not in its frequency. Later, the development of lateral roots is strongly affected. The effect of the lrt1 mutation is not as obvious in the primary root, with no influences observed on the root apical meristem structure and maintenance; however, development of the epidermis and cortex are impaired.     

Root cortical aerenchyma inhibits radial nutrient transport in maize (Zea mays)

Background and Aims

Formation of root cortical aerenchyma (RCA) can be induced by nutrient deficiency. In species adapted to aerobic soil conditions, this response is adaptive by reducing root maintenance requirements, thereby permitting greater soil exploration. One trade-off of RCA formation may be reduced radial transport of nutrients due to reduction in living cortical tissue. To test this hypothesis, radial nutrient transport in intact roots of maize (Zea mays) was investigated in two radiolabelling experiments employing genotypes with contrasting RCA.

Methods

In the first experiment, time-course dynamics of phosphate loading into the xylem were measured from excised nodal roots that varied in RCA formation. In the second experiment, uptake of phosphate, calcium and sulphate was measured in seminal roots of intact young plants in which variation in RCA was induced by treatments altering ethylene action or genetic differences.

Key Results

In each of three paired genotype comparisons, the rate of phosphate exudation of high-RCA genotypes was significantly less than that of low-RCA genotypes. In the second experiment, radial nutrient transport of phosphate and calcium was negatively correlated with the extent of RCA for some genotypes.

Conclusions

The results support the hypothesis that RCA can reduce radial transport of some nutrients in some genotypes, which could be an important trade-off of this trait.     

The effects of lead,nickel, and strontium nitrates on cell division and elongation in maize roots

The effects of Pb, Sr, and Ni nitrates on the root growth, its cell division and elongation were studied. Two-day-old maize seedlings were incubated on the 35 μM Ni(NO₃)₂, 10 μM Pb(NO₃)₂, or 3 mM Sr(NO₃)₂ in the presence or absence of 3 mM Ca(NO₃)₂. Metal toxicity was evaluated after the inhibition of root growth for the first and second days of incubation in comparison with the roots kept on water or Ca(NO₃)₂ solution. The contents of metals were determined in the apical (the first centimeter from the tip) and basal (the third centimeter from the kernel) root parts by voltamperometry and atomic-absorption spectrophotometry. We measured the length of the meristem, the length of the fully elongated cells, counted the mitotic index (MI) in the meristem and the number of meristematic cells in the cortex row; we also calculated duration the cell cycle. In the absence of Ca(NO₃)₂, the metal content in the apical root region was higher than in basal one. In the presence of Ca(NO₃)₂, we observed reverse ratio most pronounced in the case of Pb and Sr. All metals tested markedly reduced MI in the cortex, which was determined by the increase in the cell cycle duration and accompanied by the meristem shortening. These metals affected differently cell division and elongation: Ni inhibited mainly cell division and to a lesser degree their elongation, whereas Sr and Pb affected both cell division and elongation; only Sr treatment resulted in the increased length of the fully elongated cells. In the presence of Ca, all studied growth indices changed less than in the absence of Ca, which was manifested in the less severe suppression of the root growth and was in agreement with the lower accumulation of the metals in the root tips. Possible causes for the heavy metal action on growth are discussed in connection with the specificity of their transport and accumulation.     

Effects of air-filled soil porosity and aeration on the initiation and growth of secondary roots of maize (Zea mays)

We studied the possibility whether the initiation of secondary roots is regulated by the air-filled porosity in soil, i.e. the availability of oxygen in the soil. Maize plants were grown in long PVC tubes (1 m long and 12 cm diameter) and were unwatered for different numbers of days so that variations of soil water content with depth were achieved on the same date with plants at the same age. The plants were harvested when their root systems were established in the whole soil column and watering had been withheld for 0, 15, 20, 25 days. A decrease of soil water content was significantly correlated with an increase of air-filled porosity in soil. The number of secondary lateral roots from segments of primary adventitious roots increased dramatically when soil water content decreased from field capacity to about 0.05 g water g^-1 dried soil. The total dried mass of roots at different soil depths was also positively correlated with soil air-filled porosity. It was observed that the elongation of the initiated secondary roots responded differently to the variations of soil air-filled porosity. The length of secondary roots increased initially when the soil was dried from field capacity to 0.18 g g^-1 dried soil (water potential at about−0.2 MPa, air-filled porosity 0.26 cm³ cm^-3), but was drastically reduced when the soil was dried further. Obviously elongation of secondary roots was inhibited when soil water potential began to deviate substantially from an optimum value. The present results suggested that the initiation of secondary roots was greatly promoted by the increase of air-filled soil porosity, i.e. availability of oxygen. This conclusion was further verified in a separate experiment where solution-cultured maize seedlings were subjected to different aeration treatments. An obvious increase in secondary root initiation was found in plants which were aerated with normal air (21% O₂) than in plants which were either not aerated or aerated with 5% O₂ air. ei]Section editor: B E Clothier     

Cadmium translocation by contractile roots differs from that in regular,non-contractile roots

Background and Aims Contractile roots are known and studied mainly in connection with the process of shrinkage of their basal parts, which acts to pull the shoot of the plant deeper into the ground. Previous studies have shown that the specific structure of these roots results in more intensive water uptake at the base, which is in contrast to regular root types. The purpose of this study was to find out whether the basal parts of contractile roots are also more active in translocation of cadmium to the shoot.Methods Plants of the South African ornamental species Tritonia gladiolaris were cultivated in vitro for 2 months, at which point they possessed well-developed contractile roots. They were then transferred to Petri dishes with horizontally separated compartments of agar containing 50 µmol Cd(NO₃)₂ in the region of the root base or the root apex. Seedlings of 4-d-old maize (Zea mays) plants, which do not possess contractile roots, were also transferred to similar Petri dishes. The concentrations of Cd in the leaves of the plants were compared after 10 d of cultivation. Anatomical analyses of Tritonia roots were performed using appropriately stained freehand cross-sections.Key Results The process of contraction required specific anatomical adaptation of the root base in Tritonia, with less lignified and less suberized tissues in comparison with the subapical part of the root. These unusual developmental characteristics were accompanied by more intensive translocation of Cd ions from the basal part of contractile roots to the leaves than from the apical–subapical root parts. The opposite effects were seen in the non-contractile roots of maize, with higher uptake and transport by the apical parts of the root and lower uptake and transport by the basal part.Conclusions The specific characteristics of contractile roots may have a significant impact on the uptake of ions, including toxic metals from the soil surface layers. This may be important for plant nutrition, for example in the uptake of nutrients from upper soil layers, which are richer in humus in otherwise nutrient-poor soils, and also has implications for the uptake of surface-soil pollutants.     

Towards modelling the flexible timing of shoot development: simulation of maize organogenesis based on coordination within and between phytomers

Background and Aims

Experimental evidence challenges the approximation, central in crop models, that developmental events follow a fixed thermal time schedule, and indicates that leaf emergence events play a role in the timing of development. The objective of this study was to build a structural development model of maize (Zea mays) based on a set of coordination rules at organ level that regulate duration of elongation, and to show how the distribution of leaf sizes emerges from this.

Methods

A model of maize development was constructed based on three coordination rules between leaf emergence events and the dynamics of organ extension. The model was parameterized with data from maize grown at a low plant population density and tested using data from maize grown at high population density.

Key Results

The model gave a good account of the timing and duration of organ extension. By using initial conditions associated with high population density, the model reproduced well the increase in blade elongation duration and the delay in sheath extension in high-density populations compared with low-density populations. Predictions of the sizes of sheaths at high density were accurate, whereas predictions of the dynamics of blade length were accurate up to rank 9; moderate overestimation of blade length occurred at higher ranks.

Conclusions

A set of simple rules for coordinated growth of organs is sufficient to simulate the development of maize plant structure without taking into account any regulation by assimilates. In this model, whole-plant architecture is shaped through initial conditions that feed a cascade of coordination events.     

Balance between maternal and paternal alleles sets the timing of resource accumulation in the maize endosperm

Key aspects of seed development in flowering plants are held to be under epigenetic control and to have evolved as a result of conflict between the interests of the male and female gametes (kinship theory). Attempts to identify the genes involved have focused on imprinted sequences, although imprinting is only one mechanism by which male or female parental alleles may be exclusively expressed immediately post-fertilization. We have studied the expression of a subset of endosperm gene classes immediately following interploidy crosses in maize and show that departure from the normal 2 : 1 ratio between female and male genomes exerts a dramatic effect on the timing of expression of some, but not all, genes investigated. Paternal genomic excess prolongs the expression of early genes and delays accumulation of reserves, while maternal genomic excess foreshortens the expression period of early genes and dramatically brings forward endosperm maturation. Our data point to a striking interdependence between the phases of endosperm development, and are consonant with previous work from maize showing progression from cell proliferation to endoreduplication is regulated by the balance between maternal and paternal genomes, and from Arabidopsis suggesting that this ‘phasing’ is regulated by maternally expressed imprinted genes. Our findings are discussed in context of the kinship theory.     

Changes in cellular osmotic potential and mechanical properties of cell walls during light-induced inhibition of cell elongation in maize coleoptiles

The growth rate of maize ( Zea mays L. cv. Cross Bantam T51) coleoptiles in the dark was highest at the basal zone and decreased towards the tip. Growth was strongly inhibited by white fluorescent light (5 W m⁻²), especially in the basal zone of coleoptiles. Light irradiation caused an increase in the values of stress-relaxation parameters, the minimum stress-relaxation time and the relaxation rate and a decrease in the extensibility (strain/stress) of the cell walls at all zones. In addition, during growth, the accumulation of osmotic solutes was strongly inhibited by white light irradiation, resulting in an increased osmotic potential. The influences of white light on the mechanical properties of the cell wall and the osmotic potential of the tissue sap were most prominent in the basal zone. Significant correlations were observed between the increment of coleoptile length and the mechanical properties of the cell walls or the osmotic potential of the tissue sap and osmotic solutes content. Furthermore, light inhibited the outward bending of split coleoptile segments. These facts suggest that white light inhibits elongation of maize coleoptiles by modifying both the mechanical properties of the cell walls and cellular osmotic potential, which control the rate of water uptake.     

Trajectory of the nodal roots of maize in fields with low mechanical constraints

The trajectories of seventy three nodal roots of maize were studied in two fields with loose soil structure. Their projections on horizontal and vertical planes were traced. These roots tended to remain in a vertical plane. Trajectories were related to each other by an affine transformation. Thus, all the observed trajectories could be obtained by transformation of a common root archetype. The horizontal component of the trajectories was mainly in the first 0.4 m depth of soil, in the layer where soil structure was disturbed by ploughing. This horizontal component decreased with later appearance of roots (upper internodes), but differed between the two sites. The average soil temperature during the week following root appearance accounted for differences between internodes and sites. Lungley's algorithm, which is commonly used in modelling root trajectories, was tested. A general pattern could be simulated, but the model failed to fit the trajectories in the first 100 to 200 mm of soil. As a consequence, the initial angle between the stem and the root, which is a sensitive parameter in Lungley's model, did not account for differences between root trajectories. Laboratoire d'agronomie de Colmar     

Regulation of nitrate uptake by amino acids in maize cell suspension culture and intact roots

The effect of amino acids on nitrate transport was studied in Zea mays cell suspension cultures and in Zea mays excised roots. The inclusion of aspartic acid, arginine, glutamine and glycine (15mM total amino acids) in a complete cell-culture media containing 1.0 mM NO₃ ^- strongly inhibited nitrate uptake and the induction of accelerated uptake rates. The nitrate uptake rate increased sharply once solution amino acid levels fell below detection limits. Glutamine alone inhibited induction in the cell suspension culture. Maize seedlings germinated and grown for 7 days in a 15 mM mixture of amino acids also had lower nitrate uptake rates than seedlings grown in 0.5 mM Ca(NO₃)₂ or 1 mM CaCl₂. As amino acids are the end product of nitrate assimilation, the results suggest an end-product feed-back mechanism for the regulation of nitrate uptake.