Influence of soil temperature during root appearance on the trajectory of nodal roots of field grown maize

Trajectories of maize nodal roots were studied to test the hypothesis that roots which appear on a common internode have similar geometrical characteristics, and to assess the effect of soil temperature on root trajectory. Treatments consisted of three sowing dates, a comparison between mulched and non-mulched soil, a replication of one sowing date for two years in two locations, and a comparison between two cultivars at one sowing date. All these sources of variation, except the cultivar, had an appreciable effect on the trajectories of roots which appeared on the first four internodes. The horizontal component of the trajectory differed significantly between treatments, ranging from 93 to 700 mm in roots which appeared on the second internode, and from 71 to 569 for those on the third internode. The original hypothesis had, therefore, to be rejected. Mean soil temperature during the 100°C.days after root appearance accounted for the differences in trajectory between location, year, sowing date and mulch treatments, and for the differences between internodes within each location. The critical period during which temperature affected root trajectory probably began at root appearance, and ended between 50 and 100°C days after root appearance, i.e. when the root was less than 100 mm long.     

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.     

Vertical distribution of maize roots in relation to permanent soil characteristics

The vertical distribution of maize roots was studied in four contrasting soils, (arenosols, luvisols, planosols and vertisols) by using in-situ root mapping on vertical planes. The relationship between root contact frequency and depth was different for each soil, with a relatively low field-to-field variability within each soil type. The general aspect of this relationship did not change appreciably for three years in arenosols, with a low colonization in sandy layers probably being due to mechanical barriers. The relationship was consistently non-monotonic in luvisols and planosols, because of the sparse colonization of sandy layers. In planosols, these layers were traversed by some primary roots, which were essentially clustered in animal burrows. The distribution of root contact frequency was closer to an exponential function in vertisols. In these soils rooting depth and colonization of deep soil layers showed a marked increase during two dry years compared with a wet year. This was probably due to a denser net of shrinkage cracks and slickensides, where roots were essentially located in dry years. These results raise the possibility of modelling the decrease in root distribution with depth using soil information and climatic characteristics.     

Effect of mutual shading on the emergence of nodal roots and the root/shoot ratio of maize

The effect of mutual shading on the root/shoot ratio and on the number of nodal roots of maize was studied. Plants of two varieties (Dea and LG2281) were grown in individual pots of 9 L, at three plant densities: 7.5, 11 and 15 plants m^–2. A control experiment was carried out in order to study if root growth was affected by the small size of the pots. Maize plants (cv Dea) were grown at a low plant density (7.5 plants m^–2) in pots of two different volumes (9 and 25 L respectively). In both experiments plants were watered every two hours with a nutrient solution. Some plants were sampled at five dates in the main experiment and the following data were recorded: foliar stage; root, stem and leaf dry weight; number of root primordia and number of emerged roots per phytomer. The final sampling date occurred at silking.Results of the control experiment showed that the root biomass was lower in small pots but the number of nodal roots per phytomer was not affected.Results of the main experiment showed that the total plant biomass and the root/shoot ratio were lower at high plant density. The number of emerged roots was strongly reduced on the upper phytomer (P₈). This reduction was mainly due to a lower percentage of root primordia which elongated. A proposed interpretation is that the number of roots which emerge on upper phytomers is controlled by carbohydrate availability.     

Effect of soil water content on the gravitropic behavior of nodal roots in maize

The direction of root growth is an important factor that determines the spatial distribution of roots in the soil. The influence of soil water content on the direction of growth of maize nodal roots was studied both in the field and in the greenhouse. In the field experiment, the one plot was regularly irrigated (I-plot) and the other non-irrigated (N-plot). In the greenhouse experiment, three water treatments were conducted on plants grown in pots: continuously wet (CW), early drying (ED), and late drying (LD). The direction of root growth was quantified by the angle from the vertical, measured at 1 cm intervals for 10 cm from the first five internodes. Nodal roots grew more vertically in the N-plot and ED treatment than those in the I-plot and CW treatment. This was due to the decrease of the initial angle and/or the liminal angle. It is therefore thought that two events regulate the growth direction of nodal roots under dry soil conditions: gravitropic bending at root emergence from the stem and the later establishment of the angle of growth. Nodal roots appearing after rewatering in the ED treatment grew in a similar direction as those in the CW treatment. It follows from this that the water content of the surrounding soil has a direct effect on the direction of growth. Nodal roots that emerged in rapidly drying soil in the LD treatment ceased growing after showing negative gravitropism. The possible mechanisms determining the growth direction of nodal roots in drier soils are discussed.     

Effect of cyclic wetting and drying of a soil on root hair growth of maize roots

The objective of this study was to investigate the effect of cyclic soil wetting and drying on maize (Zea mays L.) root hair growth. Three soils, Chalmers silty clay loam (Typic Haplaquolls), Raub silt loam (Aquic Argiudolls) and Aubbeenaubbee sandy loam (Aric Ochraqualfs) and two soil moisture contents, −175 (M₀) and −7.5 kPa (M₁), were used to study root hair growth in a controlled-climate chamber. Increasing soil moisture after 7d from M₀ and M₁ resulted in a cessation of root hair growth behind the root cap while drying the soil after 7d from M₁ and M₀ promoted root hair growth on new but not old or existing roots. By maintaining liquid continuity under cyclic wetting and drying of a soil, root hairs may be of far greater significance to the nutrition of the plant than originally thought. Journal Paper No. 11023, Purdue Univ. Agric. Exp. Stn., W. Lafayette, IN 47907. Contribution from the Dep. of Agron.     

Plagiogravitropism of maize roots

The direction of root growth can be studied by analyzing the trajectories of roots growing in soil. Both the primary seminal root and nodal roots of maize attain a preferred, or liminal, angle of growth that deviates from the vertical. These roots are said to be plagiogravitropic. Experiments using plants grown in soil-filled boxes revealed that the primary seminal root is truly plagiogravitropic. It shows both positive and negative gravitropism in response to gravity stimuli and tends to maintain its direction even after growing around obstacles. These are experimental results suggesting that plagiogravitropic growth is controlled by internal factors. The orientation of the grain affects the establishment of the liminal angle of the primary seminal root, and both the position of their node of origin and the root diameter are closely related to the plagiogravitropic behaviour of nodal roots. Several external factors are also known to influence plagiogravitropism. Low soil water content causes a decrease in the angle of growth and soil mechanical resistance suppresses the gravitropic curvature. Plagiogravitropic behaviour of both seminal and nodal roots plays a significant role in shaping the root system.     

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     

Partitioning of reduced nitrogen derived from exogenous nitrate in maize roots: Initial priority for protein synthesis

Summary When roots of five day-old maize seedlings were exposed to¹⁵N-nitrate, a constant (25–29%) proportion of the reduced¹⁵N derived from the entering¹⁵N-nitrate accumulated as insoluble¹⁵N nitrogen. Constancy was established by two hours and lasted through 12 hours at ambient¹⁵N-nitrate concentrations of 0.05 mM to 20.0 mM. Even when little¹⁵N nitrate had been reduced (<2 moles), there was a linear relationship between accumulation of insoluble¹⁵N (but not accumulation or translocation of soluble reduced¹⁵N) and total reduced¹⁵N. It is proposed that protein synthesis from the entering nitrate occurs in close association with nitrate reduction.Paper No. 9764 of the Journal Series of the North Carolina Agricultural Research Service, Raleigh, NC, 27695-7619, USA. This research was supported by Grant No. PCM-8118661 from the National Science Foundation.Use of trade names in this publication does not imply endorsement by the North Carolina Agricultural Research Service of the product's name or criticisms of similar ones not mentioned.     

Inhibition of maize root growth by high nitrate supply is correlated with reduced IAA levels in roots

The plant root system is highly sensitive to nutrient availability and distribution in the soil. For instance, root elongation is inhibited when grown in high nitrate concentrations. To decipher the mechanism underlying the nitrate-induced inhibition of root elongation, the involvement of the plant hormone auxin in nitrate-dependent root elongation of maize was investigated. Root growth, nitrogen and nitrate concentrations, and indole-3-acetic acid (IAA) concentrations in roots and in phloem exudates of maize grown under varying nitrate concentrations were analyzed. Total N and nitrate concentrations in shoots and roots increased and elongation of primary, seminal and crown roots were inhibited with increasing external nitrate from 0.05 to 5 mM. High nitrate-inhibited root growth resulted primarily from the reduced cell elongation and not from changes in meristem length. IAA concentrations in phloem exudates reduced with higher nitrate supply. Inhibition of root growth by high nitrate was closely related to the reduction of IAA levels in roots, especially in the sections close to root tips. Exogenous NAA and IAA restored primary root growth in high nitrate concentrations. It is concluded that the inhibitory effect of high nitrate concentrations on root growth may be partly attributed to the decrease in auxin concentrations of roots.     

Inheritance of emergence time at low temperatures in segregating generations of maize

Summary North Carolina Design III and generation means analyses were used to study the inheritance of seedling emergence time and a related seedling growth parameter in crosses between 5–154, a line from CIMMYT Pool 5 with rapid seedling emergence under cool conditions, and two Corn Belt Dent lines of maize (Zea mays L.). The crosses were evaluated at low temperatures in controlled environment rooms. Additive genetic variances were larger than dominance variances in both crosses and estimates of the average levels of dominance were in the partial dominance range. Dominance was in the direction of rapid seedling emergence and rapid utilization of seed reserve. Estimates of minimum numbers of effective factors provided evidence for polygenic inheritance.     

Maize nodal root ramification: Absence of dormant primordia,root classification using histological parameters and consequences on sap conduction

Maize plants grown in field conditions were used to describe the histological organisation of the nodal roots, those of their laterals, and also to test the presence of critical stages where the subsequent capability for growth and development of young laterals was determined irreversibly. The absence of undeveloped primordia, which stop their development before boring through the nodal mother-root epidermis, proved that the number of laterals could not be regulated between the differentiation and the emission stage. Cross sections performed on nodal roots beared by the internodes 2, 4 and 6 and their long (>3 cm) and short (<3 cm) laterals showed that: u     

Tillage and N fertilization effects on maize root growth and root:shoot ratio

Two methods for estimating the size of the maize (Zea mays l.) root system from soil cores taken in the field were compared. The spatially weighed block method of estimation accounted for variation in root density by using 18 samples per plant which varied in distance from plant and soil depth. This method was compared to an estimation which averaged all of the 18 samples together. Both methods gave surprisingly similar estimates for total root growth. Increased root growth in the surface soil layers, due to tillage and N fertilization, did not impact on the estimation of total root growth. Total root length remained unchanged or increased with N fertilization, while root weight remained the same or decreased. Root mass per length decreased with N fertilization. The estimated size of the root system was used to calculate root:shoot weight ratios. The largest root:shoot ratio was found in the vegetative stage and decreased throughout the rest of the season. In this field experiment, the estimated size of the root system at 8 weeks after planting was not significantly different from the size at silking or harvest. Nitrogen fertilization significantly decreased the root:shoot weight ratio. However, tillage did not significantly change the ratio.     

Activities of proteinases in maize and soybean roots in response to anoxic stress

This study characterized the changes in proteinase activities in maize inbred line H60 and soybean cultivar Keller roots in response to anoxia. After 24 h of anoxia, crude protein extracts from both maize and soybean root tips (10 cm) were assayed for proteinase activities at pHs ranging from 4.5 to 10.2. In anoxic roots of both maize and soybean, activities of proteinases with alkaline pH optima increased, and activities of proteinases with acidic pH optima declined. Proteinases with neutral pH increased in anoxic maize roots, but declined in anoxic soybean roots. Whether the differences in proteinase activities in anaerobic maize and soybean roots contribute to the differental susceptibility of the two species requires further study.Journal Article No. 265-89.     

Plasma-membrane-bound malate dehydrogenase activity in maize roots

Summary Plasma membranes were isolated and purified from 14-day-old maize roots (Zea mays L.) by two-phase partitioning at a 6.5% polymer concentration, and compared to isolated mitochondria, microsomes, and soluble fraction. Marker enzyme analysis demonstrated that the plasma membranes were devoid of cytoplasmic, mitochondrial, tonoplast, and endoplasmic-reticulum contaminations. Isolated plasma membranes exhibited malate dehydrogenase activity, catalyzing NADH-dependent reduction of oxaloacetate as well as NAD⁺-dependent malate oxidation. Malate dehydrogenase activity was resistant to osmotic shock, freeze-thaw treatment, and salt washing and stimulated by solubilization with Triton X-100, indicating that the enzyme is tightly bound to the plasma membrane. Malate dehydrogenase activity was highly specific to NAD⁺ and NADH. The enzyme exhibited a high degree of latency in both right-side-out (80%) and inside-out (70%) vesicle preparations. Kinetic and regulatory properties with ATP and P_i, as well as pH dependence of plasma-membrane-bound malate dehydrogenase were different from mitochondrial and soluble malate dehydrogenases. Starch gel electrophoresis revealed a characteristic isozyme form present in the plasma membrane isolate, but not present in the soluble, mitochondrial, and microsomal fractions. The results presented show that purified plasma membranes isolated from maize roots contain a tightly associated malate dehydrogenase, having properties different from mitochondrial and soluble malate dehydrogenases.Abbreviations FCR ferricyanide reductase - MDH malate dehydrogenase     

Hydrotropism and its interaction with gravitropism in roots

We have studied hydrotropism and its interaction with gravitropism in agravitropic roots of a pea mutant and normal roots of peas (Pisum sativum L.) and maize (Zea mays L.). The interaction between hydrotropism and gravitropism in normal roots of peas or maize were also examined by nullifying the gravitropic response on a clinostat and by changing the stimulus-angle for gravistimulation. Depending on the intensity of both hydrostimulation and gravistimulation, hydrotropism and gravitropism of seedling roots strongly interact with one another. When the gravitropic response was reduced, either genetically or physiologically, the hydrotropic response of roots became more unequivocal. Also, roots more sensitive to gravity appear to require a greater moisture gradient for the induction of hydrotropism. Positive hydrotropism of roots occurred due to a differential growth in the elongation zone; the elongation was much more inhibited on the moistened side than on the dry side of the roots. It was suggested that the site of sensory perception for hydrotropism resides in the root cap, as does the sensory site for gravitropism. Furthermore, an auxin inhibitor, 2,3,5-triiodobenzoic acid (TIBA), and a calcium chelator, ethyleneglycol-bis-(-aminoethylether)-N,N,N,N- tetraacetic acid (EGTA), inhibited both hydrotropism and gravitropism in roots. These results suggest that the two tropisms share a common mechanism in the signal transduction step.     

Osmotic induced stimulation of the reduction of the viability dye 2,3,5-triphenyltetrazolium chloride by maize roots and callus cultures

Live cells can reduce colorless 2,3,5-triphenyltetrazolium chloride (TTC) to a red insoluble compound, formazan. Maize (Zea mays) callus, when osmotically stressed by 0.53 mol/L mannitol, produced 7-times or more formazan than untreated control callus. This result was seen with all osmotica tested and could not be attributed to differences in TTC uptake rate or accumulation, increased respiration rate as measured by O2 uptake, or to de novo protein synthesis. Increased formazan production could be detected after 2.5 h of exposure to osmotic stress and leveled off after 48 h of exposure. The increased formazan production was only detected when callus was moved from high osmotic medium to low osmotic, TTC-containing medium. Abscisic acid increased TTC reduction only when added in combination with 0.53 mol/L mannitol. Incubation of maize seedling roots with 0.53 mol/L mannitol also increased formazan production as seen visually. Further studies are needed to determine the cause of the increased formazan production. These results show that TTC viability measurements must be carefully evaluated with appropriate controls to confirm their validity.