Second-order Lateral Roots Research Articles

Cluster-root formation, carboxylate exudation and proton release of Lupinus pilosus Murr. as affected by medium pH and P deficiency

The study examined the interactive effect of pH and P supply on cluster-root formation, carboxylate exudation and proton release by an alkaline-tolerant lupin species (Lupinus pilosus Murr.) in nutrient solution. The plants were exposed to 1 (P1, deficient) and 50 µM P (P50, adequate) for 34 days in nutrient solution at either pH 5.6 or 7.8. Plant biomass was not influenced by pH at P1, but at P50 shoot and root dry weights were 23 and 18% higher, respectively, at pH 7.8 than at pH 5.6. There was no significant difference in plant biomass between two P treatments regardless of medium pH. Phosphorus deficiency increased significantly the number of the second-order lateral roots compared with the P50 treatment. Both total root length and specific root length of plants grown at pH 5.6 were higher than those at pH 7.8 regardless of P supply. Cluster roots were formed at P1, but cluster-root number was 2-fold higher at pH 7.8 than pH 5.6. Roots released 16 and 31% more protons at pH 5.6 and 7.8, respectively, in P1 than in P50 treatments, and the rate of proton release followed the similar pattern. At pH 5.6, citrate exudation rate was 0.39 µmol g−1 root DW h−1 at P1, but was under the detection limit at P50; at pH 7.8, it was 2.4-fold higher in P1 than in P50 plants. High pH significantly increased citrate exudation rate in comparison to pH 5.6. The uptake of anions P and S was inhibited at P1 and high pH increased cations Na, Mg and Ca uptake. The results suggested that enhanced cluster-root formation, proton release and citrate exudation may account for the mechanism of efficient P acquisition by alkaline-tolerant L. pilosus well adapted to calcareous soils. Cluster-root formation and citrate exudation in L. pilosus can be altered by medium pH and P deficiency. Phosphorus deficiency-induced proton release may be associated with the reduced anion uptake, but high pH-induced proton release may be partly attributed to increased cation uptake.

Root system architecture of Quercus pubescens trees growing on different sloping conditions.

Plant roots' growth direction has important implications for plant development and survival; moreover it plays an effective and vital role in stabilizing weathered soil on a steep slope. The aim of this work was to assess the influence of slope on the architecture of woody root systems. Five mature, single-stemmed Quercus pubescens trees growing on a steep slope and five on a shallow slope were excavated to a root diameter of 1 cm. A very precise numeric representation of the geometry and topology of structural root architecture was gained using a low-magnetic-field digitizing device (Fastrak, Polhemus). Several characteristics of root architecture were extracted by macros, including root volume, diameter, length, number, spatial position and branching order. The diameter at breast height (dbh) was the best predictor of the root volume but had no correlation with length and number of roots. The slope affected the root volume for each branching order, and the basal cross-sectional area (CSA), number and length of the first-order roots. Number and length of the second- and third-order laterals were closely related in both conditions, although this relationship was closer in the shallow trees, suggesting the influence of a genetic control. Sloping trees showed a clustering tendency of the first- and second-order lateral roots in the up-slope direction, suggesting that the laterals rather than the taproots provide much of the anchorage. In a steep-slope condition, the taproot tapering was positively correlated with the asymmetry magnitude of first-order roots, indicating compensation between taproot and main lateral roots' clustering tendency. These results suggest that on a slope, on clayey soils, root asymmetry appears to be a consequence of several environmental factors such as inclination, shallow-slides and soil compactness. In addition, this adaptive growth seems to counteract the turning moment induced by the self-loading forces acting in slope conditions, and as a consequence improves the tree stability.

A pteridosperm stem with clusters of shoot-borne roots from the Namurian B (Upper Carboniferous) of Hagen-Vorhalle (Germany)

Two compression specimens of a narrow pteridosperm stem with transverse sclerotic plates in the cortex are described from the upper Namurian B (Upper Carboniferous) of Hagen-Vorhalle (Germany). The pattern formed on the stem by the cortical transverse sclerotic plates permits assignment of the specimens to the mariopterids, genus Karinopteris or Mariopteris. Elongate clusters of densely spaced, up to 40 cm long shoot-borne roots with first- and second-order lateral roots extend from one side of the stem. Based on comparisons with modern angiosperms, the pteridosperm’s shoot-borne roots are considered with regard to their significance as adaptations to special physiological and mechanical requirements of scrambling/climbing plants. They may have (a) compensated for the stem’s narrow xylem area by providing additional connections with the ground, perhaps in order to maintain sufficient water and nutrient supply, (b) formed a loose interaction with support plants, and/or (c) contributed to the formation and condensation of thickets by intertwining with parts of neighboring individuals.

Root System Development of Japonica-Indica Hybrid Cultivars. 2. Branching habit.

Root system development of japonica-indica hybrid rice cultivars (J-IH : Milyang 23, Suwon 258) was compared with that of japonica-type cultivars (JT: Nipponbare, Koshihikari) with reference to branching habit from the viewpoint of developmental morphology. The four cultivars were grown in 45-liter polyethylene containers with paddy field soil and the plants with their root systems were carefully dug up at the grain-filling stage. After careful washing, the main stems of each cultivar were divided into each phytomer and the number, diameter and length of both crown and lateral roots were recorded together with their node position. From these data, the root dimensions (total number of lateral roots, total roots surface area and total root volume) were estimated for each crown root. The results indicated that lateral root development of J-IH was more vigorous than that of JT, especially in the higher phytomers formed after the panicle initiation stage. Such vigorous root branching in J-IH was mainly attributed to an increase in the number of second-order lateral roots. Detailed examination showed that the L-type first-order lateral roots were longer and the density of both L-type first-order and S-type second-order lateral roots were higher in J-IH than in JT. Thus, the root system of J-IH was characterized by long and dense L-type first-order lateral roots and large volume of S-type second-order lateral roots due to their high density. Results of this study suggest that the root system of J-IH increases the total length and surface area, by developing many S-type sedond order lateral roots, which may be related to their high-yield of J-IH.

Root and Shoot Characteristics of Peach Trees with Different Growth Habits

Shoot and root characteristics of four peach tree [Prunus persica (L.) Batsch (Peach Group)] growth habits (compact, dwarf, pillar, and standard) were studied. In compact trees, leaf number (1350/tree) was twice, but leaf area (6 cm2/leaf) was half that of pillar and standard trees. The number of lateral branches in compact trees (34) was nearly three times more than in pillar and standard trees. Leaf area index (total one-side leaf area per tree divided by the canopy cross-sectional area of the tree) of pillar trees was greater than compact, dwarf, and standard trees (13 compared with 4, 4, and 3, respectively) due to a narrower crown diameter. Dwarf trees were distinct with few leaves (134/tree) and less than half the roots of the other growth habits. Compact trees produced more higher order lateral (HOL) roots than pillar and standard trees. More second order lateral (SOL) roots were produced by compact than standard trees (1.2 vs. 0.8 SOL roots per centimeter first order lateral root). Pillar trees had higher shoot: root dry weight (DW) ratios (2.4) than compact and standard trees (1.7 for both) due to lower root DWs. Root topology was similar among compact, pillar, and standard peach trees but root axes between branch junctions (links) were significantly longer in compact trees. Compact trees had more and longer HOL roots in roots originating near the root collar (stem-root junction) (i.e., more fibrous roots) and this appeared to correlate with more lateral branches in the canopy. These results indicate significant differences in root as well as shoot architecture among growth habits that can affect their use as scion or rootstock cultivars.

169 Root Characteristics of Peach Trees with Different Shoot Growth Habits

Peach trees (Prunus persica L.) with diverse shoot growth habits have been developed, but little is known about their root systems. Characterizing shoot and root systems can improve basic understanding of peach tree growth and be important in the development of rootstocks and own-rooted trees. This research determined shoot and root characteristics of four peach tree growth habits (compact, dwarf, pillar, and standard). Seed from four peach growth habits were planted in 128-L containers, grown outside during the 1998 growing season, and then harvested. Compact tree leaf number (1350/tree) was twice, but leaf area (6 cm2/leaf) was half, that of pillar and standard trees. The number of lateral branches in compact trees (34) was nearly three-times more than in pillar and standard trees. The leaf area index (LAI) of pillar trees was greater than compact and standard trees (13 compared with 4 and 3, respectively) due to a narrower crown diameter. Dwarf tree shoots were distinct with few leaves (134 per tree) and a large LAI of 76. Compact trees grew more higher-order lateral roots than pillar and standard trees. More second-order lateral (SOL) roots were produced by compact than standard trees (1.2 vs. 0.8 SOL roots/cm first-order lateral root). Pillar trees had higher shoot-to-root dry weight ratios (2.4) than compact and standard trees (1.7 for both) due to smaller root dry weights. The results indicate fundamental differences in root characteristics among the peach tree growth habits. Compact trees had more higher order lateral roots in roots originating near the root collar (i.e., more fibrous roots), and this correlated with more lateral branches in the canopy. Shoot weights were the same among pillar, compact, and standard trees but root weights were less in pillar trees, resulting in greater shoot-to-root dry weight ratios. These results indicate significant differences in root as well as shoot architecture among growth habits that can affect their use as scion or rootstock varieties.

Morphogenetic Effect of the Herbicide Cinch on Arabidopsis thaliana Root Development

Cinch is a morphogenetically active herbicide that inhibits primary root growth and induces abnormal ``nodule-like'' lateral roots on Arabidopsis thaliana seedlings. Using 200 nm Cinch, the early stages of lateral root formation occurred along the apical half of the root axis; but once emerged, they were inhibited from further growth. Second-order lateral roots formed at the base of stunted first-order lateral roots after 5 days of Cinch treatment. Results from Cinch experiments suggested that pericycle cells are determined in the meristem to be potential sites of lateral root formation, and the developmental transition point between emerged lateral roots and subsequent growth is inhibited. Results using 2,4-dichlorophenoxyacetic acid and 2,3,5-triiodobenzoic acid suggest that Cinch is not a chemical analog of auxin.

Développement et polymorphisme racinaires chez de jeunes semis d'hévéa (Hevea brasiliensis)

The major problems (bad anchorage, water stress, and root disease) with the cropping of the rubber tree (Hevea brasiliensis) are related to the architecture of its root system. So, the morphogenetic properties of the various roots that emerge during the development of the young seedlings are described to improve our understanding of the dynamics of root system architecture. In this study, the development of the root system of 12 seedlings grown for 40 days in root observation boxes has been recorded on a daily basis and analysed. Additional observations were also made on plants grown in pots or in the field for a 60-day period. This analysis of the sequential differentiation of the root system allowed us to distinguish five different root types according to their spatial and temporal location at emergence: (i) taproot, (ii) early first-order lateral roots, (iii) acropetal first-order lateral roots, (iv) late first-order lateral roots, (v) second-order lateral roots. Large variations and close relationship were found within the growth and branching characteristics on one hand, and within anatomical characteristics on the other (number of xylem poles, external diameter, and stele diameter). These between-type variations were also observed, to a lesser extent, among roots from the same type. The continuity of these development characteristics, which were shown to be acquired early, led us to characterize a morphogenetic gradient. This gradient is a reference scale to understand the morphological plasticity of the root system growing in other mediums. Key words: root system, development, morphogenesis, anatomy, root observation box, Hevea brasiliensis.

Modelling the branching growth fractal pattern of the maize root system

Using the technique of L-systems, a growth model of the maize root system is developed. From the observation of the root systems developed under various soil density in eight root boxes, a spatial hierarchy of growth rules was extracted. The rules were divided into three categories: a meta-rule for describing features of an entire root system, a branching growth rule and a tip elongation rule. Some variations in the entire features of the root system, such as the outline and the root distribution, were confirmed by observation, and then the respective meta-rules were re-defined. The branching properties of first- and second-order lateral roots were statistically almost equal in the observations, and this lead us to set up a single stochastic branching growth rule. Tip elongation movement was not observed here; its rule had to be assumed by reference to data in the literature. A single set of branching growth and tip elongation rules were coupled with the respective meta-rules corresponding to the root samples observed, where a small scale rule was loosely governed by a large scale rule. Computer simulations offered optimized drawings of the observed root systems, and they also reproduced a typical anisotropic power distribution of roots similar to those observed.

Root Hairs and Root Lengths in Nine Warm-season Turfgrass Genotypes

Root hairs contributed variously to total root length, ranging from a low of 1% for `Emerald' zoysiagrass (Zoysia japonica Steud. x Z. tenuifolia Willd. ex Trin) and 5% for `Georgia Common' centipedegrass [Eremochloa ophiuroides (Munro.) Hack], to a high of 95% and 89% for `Texturf 10' and `FB 119' bermudagrasses [Cynodon dactylon (L.) Pers.], respectively. Genotypes ranking highest for root lengths with root hairs also ranked highest for root lengths without root hairs and for number of main roots per plant. In terms of root lengths with root hairs, first-order lateral roots contributed more to total root length than root lengths of either main roots or second-order lateral roots for all nine genotypes. Number and length of root hairs arising from either main or lateral roots were not significantly affected by their relative distance from the cap of the main root. `Texturf 10' and `FB 119' bermudagrasses ranked highest for root and root-hair extent.

Ontogeny of Alnus rubra – Alpova diplophloeus ectomycorrhizae. I. Light microscopy and scanning electron microscopy

Ectomycorrhizae synthesized between Alpova diplophloeus and Alnus rubra are of two morphological types: one with a mantle formed along the entire length of the lateral roots and the other, the clavate type, with the mantle confined to the apical portion of the laterals. The morphology of the mycorrhiza is dependent on the stage of lateral root elongation at the time of colonization by fungal hyphae. Clavate mycorrhizae form on lateral roots that have already elongated at the time of fungal colonization. Fungal hyphae interact with root hairs at the base of clavate mycorrhizae. Mantles of both types are fairly compact with few extramatrical hyphae. Hartig net hyphae, which branch profusely primarily in the radial direction, are confined to the epidermis and midway along the radial walls of the outer layer of cortical cells. Second-order lateral root primordia are initiated in the mature Hartig net zone. Cells in the outer layer of the cortex of mycorrhizal roots collapse during fixation, indicating the possible presence of a barrier in the cell wall blocking the ingress of fixative.

Effects of root severing treatments on loblolly pine

Six-year-old loblolly pine seedlings were subjected to root severing treatments varying from 0 to 100% of first-order lateral roots. Separate treatments severed surface-oriented or deep-oriented roots. Plant water status was monitored periodically for several months. After all measurements were taken, gross root system structure was determined by excavation. Treatment responses were evident on all dates of measurement. Relationships between percentage of root system cut and leaf conductance or water potential were stronger when surface-oriented roots were cut than when deep-oriented roots were cut. Severing surface-oriented first-order lateral (SOFOL) roots probably resulted in greater impact on plant water status than severing deep-oriented first-order lateral (DOFOL) roots because (i) SOFOL roots had both surface-oriented and deep-oriented second-order lateral roots that could tap both surface and subsurface soil horizons for soil moisture, and (ii) the deep-oriented second-order roots (originating from the SOFOL roots) were spatially distributed over a much larger area than the DOFOL roots and thus would have access to soil water in a larger volume of soil. For SOFOL roots the relationship between percentage cut and leaf conductance or transpiration was strongly negative; for DOFOL roots, no relationship between these variables was observed. Initially water potential decreased with the percentage of roots cut in both groups; in later measurements, water potential was affected more by severing SOFOL than DOFOL roots. Calculation of soil moisture depletion by depth indicated that both surface- and deep-oriented second-order lateral roots were important for water uptake. Severing SOFOL roots significantly decreased nitrogen, phosphorus, and potassium levels in needles of the first growth flush of the year. Levels of these elements in terminal buds were not affected by severing SOFOL roots, but were significantly reduced by severing DOFOL roots. Secondary xylem production was reduced proportionately to the amount of root system cross-sectional area severed.

Genetic variability of root growth in wheat (Triticum aestivum L.)

Ten wheat genotypes were examined for their pattern of root development in glass-fronted growth containers in the greenhouse. Differences between genotypes were recorded in the number of first-order lateral roots at 2,3 and 4 weeks' growth, the number of second-order lateral roots, the maximum depth of penetration, and the angle between the seminal root axes at 4 weeks' growth and the length of lateral root per 10 cm depth increment at 5 weeks' growth. Significant positive correlations existed between many of the seminal root parameters. Differences between genotypes in the length of nodal root axes were recorded at 7 and 8 weeks' growth. Nodal root length was influenced by tiller number and the rate of tiller initiation. Tiller development, and consequently root growth, was related to plant phasic development. Genetic modification of seminal root development appears possible, with the direction of selection depending on the definition of an optimum root development pattern.