Sloughing of cap cells and carbon exudation from maize seedling roots in compacted sand

Sloughing of root cap cells and exudation of mucilage plays an important role in the penetration of compacted soils by roots. For the first time we have quantified the rate of sloughing of root cap cells in an abrasive growth medium that was compacted to create mechanical impedance to root growth. The number of maize ( Zea mays ) root cap cells sloughed into sand increased as a result of compaction, from 1930 to 3220 d⁻¹ per primary root. This represented a 12-fold increase in the number of cells sloughed per mm root extension (from 60 to >700). We estimated that the whole of the cap surface area was covered with detached cells in compacted sand, compared with c . 7% of the surface area in loose sand. This lubricating layer of sloughed cells and mucilage probably decreases frictional resistance to soil penetration. The total carbon deposited by the root was estimated at c . 110 μg g⁻¹ sand d⁻¹. Sloughed cells accounted for <10% of the total carbon, the vast majority of carbon being contained in mucilage exudates.     

Influence of silicon pretreatment on aluminium toxicity in maize roots

The influence of Si pretreatment on Al toxicity in an Al sensitive maize variety (Zea mays L. var. BR 201 F) was investigated using root elongation rates (RER) and hematoxylin staining as stress indicators. Plants pretreated with 1 mt M Si (+ Si) and then exposed for 24 h to Al in nutrient solution without concurrent Si supply in the rooting medium exhibited higher RER than plants that were not pretreated with Si (-Si). The ameliorative effect of Si was due to lower Al uptake and to the exclusion of Al from the root tips in + Si plants. Lower Al uptake in + Si plants was not a consequence of decreased Al availability in the bulk solution. The possible mechanisms of Si-induced increase of Al resistance are discussed     

Root hair formation and elongation of primary maize roots

In maize ( Zea mays L. cv. LG 11) roots cultured in humid air, the presence of hairs was not related to root growth. However, maximum hair length and length of the hair zone could be correlated to the elongation rate of the primary root. Under the growth conditions used, the emergence of root hairs always took place in the extending zone. In more basal regions, rhizodermal cells could not give rise to root hairs. Results were similar for roots preincubated in a buffer solution.     

Brassinosteroids affect ethylene production in the primary roots of maize (<Emphasis Type="Italic">Zea mays</Emphasis> L.)

To better understand the physiological roles of brassinosteroids (BRs) in the primary roots of maize, we examined their effect on ethylene production. Exogenously applied brassinolide (BL; 10^-9 to 10^-7 M) incrementally increased the level ethylene in a dose-dependent manner. This BL-induced production was enhanced in the presence of IAA, thereby implying a synergistic effect between BR and IAA. At 10^-7 M BL, the level of free ACC was increased, but that of conjugated ACC was diminished. Moreover, greater concentrations of BL proportionally increased ACC oxidase activity. In contrast, higher levels of IAA increased the endogenous content of conjugated ACC as well as ACC synthase activity. Based on these results, we conclude that BR activates ethylene production mainly via ACC oxidase, and interacts with IAA to produce ethylene. However, the functional site for ethylene production is different for each hormone.     

Spatial separation of light perception and growth response in maize root phototropism

Although the effects of gravity on root growth are well known and interactions between light and gravity have been reported, details of root phototropic responses are less documented. We used high-resolution image analysis to study phototropism in primary roots of Zea mays L. Similar to the location of perception in gravitropism, the perception of light was localized in the root cap. Phototropic curvature away from the light, on the other hand, developed in the central elongation zone, more basal than the site of initiation of gravitropic curvature. The phototropic curvature saturated at approximately 10 micromoles m-2 s-1 blue light with a peak curvature of 29 +/- 4 degrees, in part due to induction of positive gravitropism following displacement of the root tip from vertical during negative phototropism. However, at higher fluence rates, development of phototropic curvature is arrested even if gravitropism is avoided by maintaining the root cap vertically using a rotating feedback system. Thus continuous illumination can cause adaptation in the signalling pathway of the phototropic response in roots.     

Pattern of sulfate uptake during root elongation in maize: its correlation with productivity

The efficiency of sulfate uptake was evaluated in excised roots of 22 maize genotypes, 12 inbreds and 10 hybrids, in order to study the relationship between the kinetic characteristics of the uptake and the grain productivity. During root elongation, the uptake capacity showed a pulse which appeared when the root reached 1/3 to 1/2 of its final length. The size of the accumulated pool of sulfate was significantly correlated with the productivity. The kinetic parameters of the uptake, Vmax and Km, followed the same trend, showing pulses, whoxe maximum had the same position for Vmax and Km in each genotype. The variability with the genotype of the size and duration of the Vmax pulse was not strictly connected with that of Km. The main correlation between Vmax and Km patterns was the following; inbreds were generally characterized by low Vmax and low Km; hybrids by high Vmax and high Km. As a consequence, in most cases, the benefit of the heterotic stimulation of Vmax was contrasted by the loss of affinity of the transport system or the nutrients.     

Stress induction of abscisic acid in maize roots

Moderate water stresses in the range 0 to −0.6 MPa applied with PEG 6000 to excised roots of Zea mays L. var. LG 11 induced increases of up to four-fold in the amount of abscisic acid (ABA) determined in the tissue after a 12 h period of xylem exudation. The ABA concentration of xylem exudate collected after a 2 h water stress also increased by up to four-fold. Salt stresses, induced with NaCl solutions, resulted in similar increases in the ABA concentrations. ABA concentrations in both root tissue and xylem exudate were highest 4 h after removal of the stress and then declined over a subsequent 8 h period. These results are interpreted in support of the concept that root-produced ABA may have a role in the fine control of the plant's water balance.     

Elongation by primary lateral roots and adventitious roots during conditions of hypoxia and high ethylene concentrations

Soil flooding results in unusually low oxygen concentrations and high ethylene concentrations in the roots of plants. This gas composition had a strongly negative effect on root elongation of two Rumex species. The effect of low oxygen concentrations was less severe when roots contained aerenchymatous tissues, such as in R. palustris Sm. R. thyrsiflorus Fingerh., which has little root porosity, was much more affected. Ethylene had an even stronger effect on root elongation than hypoxia, since very small concentrations (0.1 cm³ m^?3) reduced root extension in the two species, and higher concentrations inhibited elongation more severely than did anoxia in the culture medium. Thus, ethylene contributes strongly to the negative effects of flooding on root growth. An exception may be the highly aerenchymatous, adventitious roots of R. palustris. Aerenchyma in these roots provides a low-resistance diffusion pathway for both endogenously produced ethylene and shoot-derived oxygen. This paper shows that extension by roots of R. palustris in flooded soil depends almost completely on this shoot-derived oxygen, and that aerenchyma prevents accumulation of growth-inhibiting levels of ethylene in the root.     

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.     

Responses of maize roots to drying – limits of viability

Turgid pieces of mature maize roots were dried in air and progressive changes in their relative water content (RWC) determined. Viability was tested by reproducibility of the drying curves after dehydration to successively lower RWCs. After reaching a chosen RWC, the pieces were rehydrated (approximately 2 h), and a 2nd and 3rd dehydration curve measured. Each drying curve was characterized by two parameters (a scale parameter λ, and a shape parameter β) of a survivorship function, which is a linear function of time. The parameter λ is more informative, and does not change in successive dehydrations for RWC > 0·4, suggesting no irreversible damage to the roots. Damage and death were indicated by divergences of λ in successive dehydrations to RWC = 0·35–0·15. Cryo-analytical microscopy confirmed these data while indicating specifically death of 50 and 100% of cortical cells at RWC 0·30 and 0·15, respectively, and survival of 50% or more of sieve tubes, pericycle and vascular parenchyma cells at root RWC as low as 0·15. This pattern of stelar cell survival may allow roots to preserve their capacity for renewal of axial conductivity and branch root development following periods of severe water stress.     

Effect of ethylene antagonists on auxin-induced inhibition of intact primary root elongation in maize (Zeamays L.)

We tested that the hypothesis that root elongation might be controlled by altering the level of ethylene in intact primary roots of maize(Zea mays L.). We measured root elongation in a short period using a computerized root auxanometer. Compounds which regulate ethylene production were applied to intact primary roots in different time periods. Root elongation was stimulated by the treatment with ethylene antagonists such as Co²⁺, aminoethoxyvinylglycine (AVG) and L-canaline. This result suggested that root elongation was closely related to ethylene level of intact primary roots. Furthermore, IAA- and 1-aminocyclopropane-1-carboxylic acid (ACC)-induced inhibition of root elongation was reversed by treatment with Co²⁺. The application of ACC to roots which have been exposed to IAA and Co²⁺ have no significant effect on root elongation. However, the inhibition of root elongation by ACC in roots previously treated with IAA and AVG became manifest when the applied IAA concentrations were lower. These results were consistent with the hypothesis that the level of ethylene in intact roots functions to moderate root elongation, and suggested that auxin-induced inhibition of root elongation results from auxin induced promotion of ethylene production.     

Genetic variation in ZmTIP1 contributes to root hair elongation and drought tolerance in maize

Drought is a major abiotic stress that threatens maize production globally. A previous genome‐wide association study identified a significant association between the natural variation of ZmTIP1 and the drought tolerance of maize seedlings. Here, we report on comprehensive genetic and functional analysis, indicating that ZmTIP1, which encodes a functional S‐acyltransferase, plays a positive role in regulating the length of root hairs and the level of drought tolerance in maize. We show that enhancing ZmTIP1 expression in transgenic Arabidopsis and maize increased root hair length, as well as plant tolerance to water deficit. In contrast, ZmTIP1 transposon‐insertional mutants displayed the opposite phenotype. A calcium‐dependent protein kinase, ZmCPK9, was identified as a substrate protein of ZmTIP1, and ZmTIP1‐mediated palmitoylation of two cysteine residues facilitated the ZmCPK9 PM association. The results of this research enrich our knowledge about ZmTIP1‐mediated protein S‐acylation modifications in relation to the regulation of root hair elongation and drought tolerance. Additionally, the identification of a favourable allele of ZmTIP1 also provides a valuable genetic resource or selection target for the genetic improvement of maize.     

Dynamics of Aerenchyma distribution in the cortex of sulfate-deprived adventitious roots of maize

BACKGROUND AND AIMS: Aerenchyma formation in maize adventitious roots is induced in nutrient solution by the deprivation of sulfate (S) under well-oxygenated conditions. The aim of this research was to examine the extent of aerenchyma formation in the cortex of sulfate-deprived adventitious roots along the root axis, in correlation with the presence of reactive oxygen species (ROS), calcium levels and pH of cortex cells and root lignification. METHODS: The morphometry of the second whorl of adventitious (W2) roots, subject to S-deprivation conditions throughout development, was recorded in terms of root length and lateral root length and distribution. W2 roots divided into sectors according to the mean length of lateral roots, and cross-sections of each were examined for aerenchyma. In-situ detection of alterations in ROS presence, calcium levels and pH were performed by means of fluorescence microscopy using H(2)DCF-DA, fluo-3AM and BCECF, respectively. Lignification was detected using the Wiesner test. KEY RESULTS: S-deprivation reduced shoot growth and enhanced root proliferation. Aerenchyma was found in the cortex of 77 % of the root length, particularly in the region of emerging or developing lateral roots. The basal and apical sectors had no aerenchyma and no aerenchyma connection was found with the shoot. S-deprivation resulted in alterations of ROS, calcium levels and pH in aerenchymatous sectors compared with the basal non-aerenchymatous region. Lignified epidermal layers were located at the basal and the proximal sectors. S-deprivation resulted in shorter lateral roots in the upper sectors and in a limited extension of the lignified layers towards the next lateral root carrying sector. CONCLUSIONS: Lateral root proliferation is accompanied by spatially localized induced cell death in the cortex of developing young maize adventitious roots during S-deprivation.     

Expression of genetically distinct superoxide dismutases in the maize seedling during development

Immunoassays for the cytosolic and mitochondrial superoxide dismutases (SOD) of maize were developed and used to study the expression of these proteins in the maize seedling. The genetically distinct proteins, SOD-3 and SOD-4, are preferentially expressed in the scutellum, comprising approximately 1% of the total water-soluble protein of that tissue. SOD-2, SOD-3, and SOD-4 are synthesized in the scutellum during early sporophytic development, probably on cytosolic ribosomes. Two-dimensional gel electrophoresis of crude scutellar extracts indicates that significant changes occur in the protein composition of the maize scutellum following seed imbibition. Using the immunoassays, a maize line exhibiting a significant reduction in cyanide-sensitive SOD protein was identified.     

Aerenchyma formation in the wetland plant Juncus effusus is independent of ethylene

Flooded plant roots commonly form aerenchyma, which allows gas diffusion between shoots and roots. The programmed cell death involved in this induced aerenchyma formation is controlled by the plant hormone ethylene, as has been shown for maize (Zea mays). However, the role of ethylene is uncertain in wetland species that form constitutive aerenchyma (also under nonflooded conditions). The aim of this study is to shed light on the involvement of ethylene in constitutive aerenchyma formation in Juncus effusus. Plants of J. effusus and maize were treated with ethylene and inhibitors of ethylene action to determine the consequences for aerenchyma formation. Neither treatment with high ethylene concentrations nor with ethylene inhibitors resulted in changes in root aerenchyma in J. effusus. By contrast, ethylene increased aerenchyma development in maize unless ethylene action inhibitors were applied simultaneously. Similarly, root elongation was insensitive to ethylene treatment in J. effusus, but was affected negatively in maize. The data show that aerenchyma in J. effusus is highly constitutive and, in contrast to the inducible aerenchyma in maize, is not obviously controlled by ethylene.     

Suberin lamella development in maize seedling roots grown in aerated and stagnant conditions

Hypoxia can stimulate the development of a suberized exodermis in aquatic plants; however, its influence on this aspect of terrestrial root development is sparsely documented. To determine the effects of hypoxia on maize (Zea mays cv. Seneca Horizon) roots, seedlings were grown in vermiculite (VERM), aerated hydroponics (AER), stagnant hydroponics with agar (STAG), or aerated hydroponics with agar (AERAG). The endo- and exodermis were examined for wall modifications. Lateral root emergence and aerenchyma formation were documented qualitatively. The endodermal Casparian band formation was unaffected by treatment. Endodermal and exodermal suberin lamella formation was earliest and most extensive in VERM. Suberization, especially in the exodermis of aerated treatments, was depressed in all hydroponic media. In comparison with AER, STAG exodermal lamellae were increased, but endodermal lamellae were decreased. Since the suberized exodermis forms a barrier to radial oxygen loss from roots to the medium, its stimulation in STAG roots (which also developed extensive aerenchyma) would help retain oxygen in the root. The reduction of endodermal lamellae should facilitate oxygen diffusion into the stele. Clearly, the response to environmental conditions is variable within individual cortical cell layers. Additionally, the observed patterns of lamellae, aerenchyma and lateral root development indicate a tight radial co-ordination of root development.     

Sequence of drought response of maize seedlings in drying soil

Leaf elongation in monocotyledonous plants is sensitive to drought. To better understand the sequence of events in plants subjected to soil drying, leaf elongation and transpiration of maize seedlings ( Zea mays L.) of 4 cultivars were monitored continuously and the diurnal courses of the root and leaf water relations were determined. Results from this study indicate the following sequence of drought response: Leaf elongation decreased before changes in the leaf water relations of non‐growing zones of leaf blades were detected and before transpiration decreased. Reductions in leaf elongation preceded changes in the root water potential (ψ_w). Root ψ_w was not a very sensitive indicator of soil dryness, whereas the root osmotic potential (ψ_s) and root turgor (ψ_p) were more sensitive indicators. The earliest events observed in drying soil were a significant increase in the largest root diameter class (1 720 to 1 960 gm) and a decrease in leaf elongation ( P = 0.08) 2 days after withholding water. Significant increases in root length were observed 2 days later. Soil drying increased the number of fine roots with diameters of <240 µm. Slight increases in soil strength did not affect leaf elongation in the drying soil.     

De-submergence-induced ethylene production in Rumex palustris: regulation and ecophysiological significance

Rumex palustris responds to total submergence by increasing the elongation rate of young petioles. This favours survival by shortening the duration of submergence. Underwater elongation is stimulated by ethylene entrapped within the plant by surrounding water. However, abnormally fast extension rates were found to be maintained even when leaf tips emerged above the floodwater. This fast post-submergence growth was linked to a promotion of ethylene production that is presumed to compensate for losses brought about by ventilation. Three sources of ACC contributed to post-submergence ethylene production in R. palustris: (i) ACC that had accumulated in the roots during submergence and was transported in xylem sap to the shoot when stomata re-opened and transpiration resumed, (ii) ACC that had accumulated in the shoot during the preceding period of submergence and (iii) ACC produced de novo in the shoot following de-submergence. This new production of ethylene was associated with increased expression of an ACC synthase gene (RP-ACS1) and an ACC oxidase gene (RP-ACO1), increased ACC synthase activity and a doubling of ACC oxidase activity, measured in vitro. Out of seven species of Rumex examined, a de-submergence upsurge in ethylene production was seen only in shoots of those that had the ability to elongate fast when submerged.     

The effect of local application of fertilizer on the content of cytokinins in the xylem sap of maize

Cytokinin content in xylem sap was higher in plants grown under local supply of fertilizers as compared to those grown under homogenous distribution of nutrients in soil. The separate assay of cytokinins in xylem exudate from split root system showed that roots, which were in contact with fertilizer mainly contributed to cytokinins transported from roots to shoots. This revised version was published online in July 2006 with corrections to the Cover Date.