Influence of root diameter on the penetration of seminal roots into a compacted subsoil

A field experiment was conducted to evaluate the influence of root diameter on the ability of roots of eight plant species to penetrate a compacted subsoil below a tilled layer. The soil was a fine sandy loam red-brown earth with a soil strength of about 3.0 MPa (at water content of 0.13 kg kg^-1, corresponding to 0.81 plastic limit) at the base of a tilled layer. Relative root diameter (RRD), which was calculated as the ratio of the mean diameters of roots of plants grown in compacted soil to the mean diameters of those from uncompacted soil, was used to compare the sensitivity of roots to thicken under mechanical stress.Diameters of root tips of plants grown in soil with a compacted layer were consistently larger than those from uncompacted soil. Tap-rooted species generally had bigger diameters and RRDs than fibrous-rooted species. A higher proportion of thicker roots penetrated the strong layer at the interface than thinner roots. There were differences between plant species in the extent to which root diameter increased in response to the compaction. The roots which had larger RRD also tended to have higher penetration percentage.The results suggest that the size of a root has a significant influence on its ability to penetrate strong soil layers. It is suggested that this could be related to the effects which root diameter may have on root growth pressure and on the mode of soil deformation during penetration.     

Penetration of cover crop roots through compacted soils

Tap-rooted species may penetrate compacted soils better than fibrous-rooted species and therefore be better adapted for use in “biological tillage”. We evaluated penetration of compacted soils by roots of three cover crops: FR (forage radish: Raphanus sativus var. longipinnatus, cv. ‘Daikon’), rapeseed (Brassica napus, cv. ‘Essex’), two tap-rooted species in the Brassica family, and rye (cereal rye: Secale cereale L., cv. ‘Wheeler’), a fibrous-rooted species. Three compaction levels (high, medium and no compaction) were created by wheel trafficking. Cover crop roots were counted by the core-break method. At 15–50 cm depth under high compaction, FR had more than twice and rapeseed had about twice as many roots as rye in experiment 1; FR had 1.5 times as many roots as rye in experiment 2. Under no compaction, little difference in root vertical penetration among three cover crops existed. Rapeseed and rye root counts were negatively related to soil strength by linear and power functions respectively, while FR roots showed either no (Exp.1) or positive (Exp. 2) relationship with soil strength. We conclude that soil penetration capabilities of three cover crops were in the order of FR > rapeseed > rye.     

Soil strength influences wheat root interactions with soil macropores

Deep rooting is critical for access to water and nutrients found in subsoil. However, damage to soil structure and the natural increase in soil strength with depth, often impedes root penetration. Evidence suggests that roots use macropores (soil cavities greater than 75 μm) to bypass strong soil layers. If roots have to exploit structures, a key trait conferring deep rooting will be the ability to locate existing pore networks; a trait called trematotropism. In this study, artificial macropores were created in repacked soil columns at bulk densities of 1.6 g cm⁻³ and 1.2 g cm⁻³, representing compact and loose soil. Near isogenic lines of wheat, Rht-B1a and Rht-B1c, were planted and root–macropore interactions were visualized and quantified using X-ray computed tomography. In compact soil, 68.8% of root–macropore interactions resulted in pore colonization, compared with 12.5% in loose soil. Changes in root growth trajectory following pore interaction were also quantified, with 21.0% of roots changing direction (±3°) in loose soil compared with 76.0% in compact soil. These results indicate that colonization of macropores is an important strategy of wheat roots in compacted subsoil. Management practices to reduce subsoil compaction and encourage macropore formation could offer significant advantage in helping wheat roots penetrate deeper into subsoil.     

Root cap removal increases root penetration resistance in maize (Zea mays L)

The root cap assists the passage of the root through soil by means of its slimy mucilage secretion and by the sloughing of its outer cells. The root penetration resistance of decapped primary roots of maize (Zea mays L. cv. Mephisto) was compared with that of intact roots in loose (dry bulk density 1.0 g cm-3; penetration resistance 0.06 MPa) and compact soil (1.4 g cm-3; penetration resistance 1.0 MPa), to evaluate the contribution of the cap to decreasing the impedance to root growth. Root elongation rate and diameter were the same for decapped and intact roots when the plants were grown in loose soil. In compacted soil, however, the elongation rate of decapped roots was only about half that of intact roots, whilst the diameter was 30% larger. Root penetration resistances of intact and decapped seminal axis were 0.31 and 0.52 MPa, respectively, when the roots were grown in compacted soil. These results indicated that the presence of a root cap alleviates much of the mechanical impedance to root penetration, and enables roots to grow faster in compacted soils.     

Penetration of very strong soils by seedling roots of different plant species

The abilities of seedling roots of twenty-two plant species to penetrate a strong growth medium were compared under controlled conditions. Seedlings were grown for 10 days in compression chambers filled with siliceous sandy soil at 0.2 kg kg^–1 water content and mean penetrometer resistance of 4.2 MPa. Root elongation and thickening were measured after growth. The results show that soil strength reduced the elongation of roots of all plant species by over 90% and caused the diameters of the roots to increase compared with control plants grown in vermiculite (0 MPa resistance).Differences in both root elongation and root diameter were observed among plant species. Generally, the roots of dicotyledons (with large diameters) penetrated the strong medium more than graminaceous monocotyledons (with smaller diameters). There was a significant positive correlation (r=0.78, p<0.05) between root diameter and elongation over all the species in the stressed plants. The species were ranked according to the relative root elongation and relative root thickening. Based on this ranking, lupin (Lupinus angustifolius), medic (Medicago scutelata) and faba bean (Vicia faba) were the species with the greatest thickening and elongation while wheat (Triticum aestivum), rhodesgrass (Chloris gayana) and barley (Hordeum vulgare) had the least. The weight of the seeds did not seem to influence either the thickening or elongation of the roots.     

A comparison of penetrometer pressures and the pressures exerted by roots

Summary Previous work is reviewed in which the ratio of the pressures required for soil penetration by roots and penetrometers are compared. It appears that this ratio can vary from about 2 to 8 depending on conditions. However, there is very little experimental evidence and most of the work has been inferential.Direct measurements are reported for the stresses exerted by a 1 mm diameter penetrometer probe and by the roots of pea seedlings when penetrating Urrbrae fine sandy loam. Six soil conditions were used: (non-weathered remoulded soil cores + artificially weathered remoulded soil cores + undisturbed field clods) × (confined + unconfined cores or clods). The confinement treatment was to test for any effects of additional restraint to cylindrical root expansion. The weathering and field clod treatments were to test the hypothesis that root elongation is facilitated by tensile failure ahead of the root tip.The principal conclusions are as follows. The laboratory weathering treatment reduced the soil tensile strength by 25%. This resulted in a small but significant reduction in the pressure for root penetration into confined cores. Compared with remoulded non-weathered cores, field clods had a 2 to 3 fold greater penetrometer resistance and a 50% lower tensile strength. The force required for root penetration into unconfined field clods was only 10% greater than for unconfined non-weathered cores. For the former (which is closest to field conditions) the penetrometer had to exert a pressure 5.1 times greater than a root tip in order to penetrate the soil. Penetrometer penetration pressure was independent of probe diameter in the 1–2 mm range in the soil used. Core confinement restricts root radial expansion and modifies the penetration force of metal probes and plant roots.On the basis of the new results it is tentatively concluded that soil tensile failure can facilitate penetration by roots.     

Rooting characteristics of two widely distributed woody plant species growing in different karst habitats of southwest China

A series of studies claimed that deep root development of plant established in karst regions was facilitated by fractured bedrock beneath the shallow soils; however, bedrock is not a uniform medium for root proliferation. We hypothesized plant species that survived in different karst habitats had some other rooting characteristics rather than deep penetration. To test the hypothesis, coarse root systems of two widely distributed woody species (one tree and one shrub) growing in three typical rocky karst habitats (shallow soil, loose rocky soil and exposed rock) were excavated in karst region of southwest China. Root systems were investigated based on four parameters: maximum rooting depth, maximum radial extent, root tapering pattern and root curvature. In all the three habitats, maximum rooting depths were no deeper than 120 and 40 cm for the tree and shrub species, respectively. Maximum radial extents were extremely large compared with maximum rooting depth, indicating that rooting characteristics were dominated by horizontal extension rather than deep penetration. Roots of both species growing in shallow soil habitat tapered gradually and curved slightly, which was consistent with the specific characteristics of this habitat. On the contrary, roots of the tree species growing in the other two habitats tapered rapidly but curved slightly, while roots of the shrub species tapered gradually but curved strongly. It was speculated that limited depths and rapid tapering rates of the tree roots were likely compensated by their utmost radial extensions, while the shrub species might benefit from its root curvature as the associated root tropisms may increase the ability of root to encounter more water and contribute to potentially high resource absorption efficiency. Our results highlight the importance of taking shallow-rooted species into account in understanding the distribution of natural plant communities and predicting future vegetation dynamics in karst regions.     

Genotypic differences in root penetration ability of wheat through thin wax layers in contrasting water regimes and in the field

A significant proportion of arable land in south-western Australia is highly susceptible to subsoil compaction, which limits access of roots of wheat to water and nutrients at depth. Genotypic variation in the ability of roots to penetrate a hardpan has been reported for other cereals, using a pot technique, where a thin wax-layer of paraffin wax and petroleum jelly is placed in a soil column to simulate a hardpan. Previously we have modified and validated this technique for measuring root penetration ability of wheat seedlings under contrasting water regimes. Here we report on a series of five experiments (runs), two in well-watered and three in drought stress conditions, which evaluated seminal and nodal root penetration ability through thin wax layers among 24 Australian wheat cultivars and breeding lines (entries). These results were compared with observations on their rooting depths in two contrasting soil types in field trials, including a sandy duplex that contained a hardpan and a red clay that increased in soil strength with depth. Nodal roots ceased growth early under soil water deficit, and water uptake was instead dependant on seminal roots under conditions imposed in the pots. Plants were then reliant on the ability of seminal roots to penetrate the wax layer. Eight entries had superior root penetration ability in both well-watered and drought stressed conditions. Roots of three other entries, which failed to penetrate the wax layers, died under drought stress conditions. In field trials, there was a significant interaction between site and entry for maximum root depth. Our results from the pot studies and field trials indicate that there exists genotypic variation in root traits that are required to penetrate uniformly hard soil, dry soil or soil containing a hardpan. As four of the eight superior entries also showed superior root penetration ability at both sites in the field, there was an overall consistency, but there were exceptions at individual field sites. Factors likely to result in such exceptions were discussed, and topics for further research identified.     

Growth of seedling roots in response to external osmotic stress by polyethylene glycol 20,000

Seedling roots of ten plant species were grown in siliceous sand wetted with solutions of polyethylene glycol (PEG) of MW=20,000 with osmotic potentials of 0.0, ? 0.25, ? 0.5 and ? 1.0 MPa. After 48 h growth under controlled lighting, root elongation and root diameter were measured. Root elongation of all species was reduced by increasing levels of external osmotic stress. Dicotyledonous species were affected more than monocotyledons at potentials of ? 0.25 and ? 0.5 MPa but less at ? 1.0 MPa. Root diameters of all the species were thicker than those of the unstressed at potentials of ? 0.25 and ? 0.5 MPa. At a potential of ? 1.0 MPa the dicotyledons were still thicker, though not by as much as they were at ? 0.25 and ? 0.5 MPa. The monocotyledons, in contrast, were thinner at ? 1.0 MPa. There was a significant positive correlation (r=0.81, p <0.01) between root diameter and root elongation at ? 1.0 MPa potential. Species were ranked according to the relative root elongation (RRE) and relative root thickness (RRT) at the highest level of stress (? 1.0 MPa). In both rankings dicotyledonous species were in the top ranks and monocotyledous species were in lower positions. The results are compared with those for the elongation and thickening of roots growing against external mechanical stress obtained in a previous study. There were good correlations between the responses observed for the two types of external stress. The implications of these findings are discussed.     

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.     

Screening the ability of rice roots to overcome the mechanical impedance of wax layers: importance of test conditions and measurement criteria

The development of a wax layer method for screening the ability of rice (Oryza sativa L.) roots to overcome mechanical impedance is described. Wax layers (3 mm thick) made of mixtures of white soft paraffin and paraffin wax were installed 50 mm deep in tubes of sand. The sand was watered with nutrient solution and planted with 3-d old rice seedlings. The numbers of root axes per plant that had penetrated the wax layers 24 d after planting were counted. The ratio of penetrated to total root axes per plant gave a misleading measure of root penetration ability, as rice varieties differed in the ratio of penetrated to total axes in a low impedance (3% wax) control. In non-flooded conditions, a 60% wax layer decreased root penetration (number of roots penetrating the wax layer per plant) to a mean of 74% of the low impedance control, whereas an 80% wax layer decreased mean root penetration to 31% of the control. The best measure of root penetration in non-flooded conditions was the number of axes penetrating an 80% wax layer. Flooding decreased root penetration of a 60% wax layer to a mean of 26% of the low impedance control. The best measure of root penetration in flooded conditions was the number of axes penetrating a 60% wax layer.     

Changes in growth,porosity, and radial oxygen loss from adventitious roots of selected mono‐ and dicotyledonous wetland species with contrasting types of aerenchyma

Growth in stagnant, oxygen‐deficient nutrient solution increased porosity in adventitious roots of two monocotyledonous (Carex acuta and Juncus effusus) and three dicotyledonous species (Caltha palustris, Ranunculus sceleratus and Rumex palustris) wetland species from 10 to 30% under aerated conditions to 20–45%. The spatial patterns of radial oxygen loss (ROL), determined with root‐sleeving oxygen electrodes, indicated a strong constitutive ‘barrier’ to ROL in the basal root zones of the two monocotyledonous species. In contrast, roots of the dicotyledonous species showed no significant ‘barrier’ to ROL when grown in aerated solution, and only a partial ‘barrier’ when grown in stagnant conditions. This partial ‘barrier’ was strongest in C. palustris, so that ROL from basal zones of roots of R. sceleratus and R. palustris was substantial when compared to the monocotyledonous species. ROL from the basal zones would decrease longitudinal diffusion of oxygen to the root apex, and therefore limit the maximum penetration depth of these roots into anaerobic soil. Further studies of a larger number of dicotyledonous wetland species from a range of substrates are required to elucidate the ecophysiological consequences of developing a partial, rather than a strong, ‘barrier’ to ROL.     

Formation of aggregates by plant roots in homogenised soils

The influence of root growth and water regime on the formation of aggregates was studied in modified minirhizotrons under controlled conditions. Two soils, a black earth (67% clay) and a red-brown earth (19% clay) were ground and forced through a 0.5 mm sieve. Ryegrass, pea and wheat were grown for fifteen wetting and drying (wd) cycles for 5 months. Another set of minirhizotrons was not planted and served as a control. Measurements of aggregate size distribution (ASD), aggregate tensile strength (ATS), aggregate stability (AS), aggregate bulk density (ABD) and organic carbon (OC) were made on single aggregates of the 2–4 mm fraction. The results showed that aggregates of the black earth which has a high clay content and shrink/swell properties had more smaller aggregates with higher ATS, AS and ABD than those from the red-brown earth. It was also found that for both soils: (1) w/d cycles and higher root length density (RLD) increased the proportions of smaller aggregates and aggregate strength; (2) differences in the ability of the plant species to influence aggregation was evident and seemed to be related to the RLD. The RLD was in the order ryegrass > wheat > pea. Mechanisms likely to be involved in processes of aggregate formation and stabilization are discussed. They include cracking of soil due to tensile stresses generated during drying of a shrinking soil; changes in pore water pressure within the soil mass caused by water uptake by plant roots generating effective stresses; and biological processes associated with plant roots and root exudates.     

Genotypic variation in the ability of wheat roots to penetrate wax layers

Background and aims

The role of the root system in mediating crop yields has recently been emphasised, resulting in several laboratory approaches for phenotyping root traits. We aimed to determine the existence of, and reasons for, genotypic variation in wheat (Triticum aestivum L.) root penetration of strong wax layers.

Methods

Three contrasting groups (UK elite lines, CIMMYT lines and near-isogenic lines of cv Mercia containing dwarfing and semi-dwarfing Rht alleles) comprising 18 different genotypes with contrasting phenologies were studied. We determined the ability of roots of these genotypes to penetrate strong wax layers and the angular spread of the root systems.

Results

There were no intrinsic differences in root system ability to penetrate strong wax layers (consistent with the similar root diameter of all lines) since greater root penetration was simply related to more root axes. Recording root penetration of concentric zones of the wax layer demonstrated that cv. Battalion had a root system with a smaller angular spread than cv. Robigus, which had the root system with the greatest angular spread.

Conclusions

There was limited genotypic variability in root penetration of strong layers within the wheat cultivars studied. A key challenge will be to determine the physiological and agronomic significance of the variation in root angular spread.     

Specialized 'dauciform' roots of Cyperaceae are structurally distinct, but functionally analogous with 'cluster' roots

When grown in nutrient solutions of extremely low [P] (相似文献   

Root penetration through a high bulk density soil layer: differential response of a crop and weed species

Weed control remains one of the greatest problems in agro-ecological systems. An important factor controlling crop and weed competition for below ground resources is the presence of compacted soil layers or ‘hard pans’. In a series of experiments, we investigated the ability of roots of soybean (Glycine max L.) and the weeds sicklepod (Senna obtusifolia L.) and Palmer amaranth (Amaranthus palmeri S. Wats) to penetrate through a compacted soil layer and acquire N from lower in the soil profile. Soil columns were constructed to simulate a compacted soil layer with different bulk densities ~8 cm beneath the soil surface. Results indicated that roots of the two weed species penetrated high bulk density soil layers more effectively than those of four soybean lines. Root penetration was not related with growth rates among the species or soybean genotypes. Overall root and shoot growth of the weeds was sustained when downward root growth was inhibited, while both declined with soybean, even under high fertility conditions. The weeds also acquired relatively high amounts of $^{15} {\text{N}} - {\text{NO}}_{\text{3}}^ - $ from buried patches beneath the high bulk density layers compared to soybean. The results indicate that the weed species would have a competitive advantage when plow pans are present, and an attempt is made to relate this advantage with competitive dynamics observed in the field.     

Effects of phosphate sorptivity on long-term plant recovery and effectiveness of fertilizer phosphate in soils

Summary A greenhouse experiment was carried out on 30 soils of eastern Australia in the first year and 15 soils in the following years to determine the effects of soil phosphate sorptivity and phosphate removal in harvested material on effectiveness and recovery of fertilizer phosphate by white clover during a four year period.Phosphate recovery by the clover and fertilizer effectiveness were primarily negative functions of phosphate sorptivity. After the first year there was a progressive decrease in the sorptivity effect such that the subsequent decreases in phosphate recovery and effectiveness were largest on weakly sorptive soils and smallest on strongly sorptive soils. In the long term the cumulative effects of the apparently slow immobilizing reactions on the weakly sorptive soils tended to equalize the effects of faster reactions on strongly sorptive soils so that the actual recovery and effectiveness in the final year were similar on all soils regardless of their sorptivities. The cumulative recovery and cumulative effectiveness in the long term, however, remained strongly negative functions of sorptivity. Phosphate removal in harvested material caused large decreases in phosphate effectiveness on all soils and particularly on weakly sorptive soils.     

Soil physical strength rather than excess ethylene reduces root elongation of Eucalyptus seedlings in mechanically impeded sandy soils

Seedling establishment in heavily compact soils is hampered by poor root growth caused by soil chemical or physical factors. This study aims to determine the role of ethylene in regulating root elongation through mechanically impeded sandy soils using Eucalyptus todtiana F. Muell seedlings. Concentrations of ethephon (1, 10, and 100???M) were added to non-compact soils, and endogenous ethylene production from seedling roots was compared to ethylene production of roots grown in physically compacted field soils (98.6?% sand). The ethylene-inhibitor 3,5-diiodo-4-hydroxybenzoic acid (DIHB) (0.1???M) was included for each treatment to counteract the negative effects of excess ethylene or compact soils on root elongation. Root elongation was reduced in high ethylene soils by 49?% and high bulk density soils by 44?%. Root ethylene production increased ninefold in roots grown in the high ethylene environment (100???M), but decreased 80?% in compact soils. The use of DIHB did not alter root length and produced varying results with respect to ethylene production, suggesting an interaction effect involving high amounts of soil ethylene. While ethylene regulates root growth, the physical strength of sandy soils is the major factor limiting root elongation in mechanically impeded soils.     

The effect of soil strength on the growth of pigeonpea radicles and seedlings

The effect of soil strength on the growth of pigeonpea radicles and seedlings was investigated in cores of three clay soils prepared at different water contents and bulk densities in the laboratory.Radicle elongation directly into soil cores was reduced from 50–70 mm d^-1 at strengths less than 0.5 MPa to 0 mm d^-1 at 3.5–3.7 MPa. The response to soil strength was affected by the water content of the soil, presumably as a result of reduced oxygen availability in wetter soil. This effect was apparent in soils wet to air-filled porosities less than 0.15 m³ m^-3.Radicles were more sensitive to high soil strength (>1.5 MPa) than were seedling roots which encountered the same conditions at 60 mm in the profile. Radicle growth ceased at 3.5 MPa which reduced seedling root growth by only 60%.Despite a 60% reduction in root length in the high strength zone, seedling roots compensated in zones of loose soil above and below the compacted layer, and total root length and shoot growth were unaffected. There was no evidence of a root signal response which results in reduced shoot growth in some species in response to high soil strength.The proliferation of roots in surface layers and the delayed penetration of the root system to depth in compacted soil are likely to expose seedlings to a greater risk of water-deficit in the field, particularly under dryland conditions where plants rely on stored subsoil water for growth.     

Theoretical evidence that root penetration ability interacts with soil compaction regimes to affect nitrate capture

Background and AimsAlthough root penetration of strong soils has been intensively studied at the scale of individual root axes, interactions between soil physical properties and soil foraging by whole plants are less clear. Here we investigate how variation in the penetration ability of distinct root classes and bulk density profiles common to real-world soils interact to affect soil foraging strategies.MethodsWe utilize the functional–structural plant model ‘OpenSimRoot’ to simulate the growth of maize (Zea mays) root systems with variable penetration ability of axial and lateral roots in soils with (1) uniform bulk density, (2) plow pans and (3) increasing bulk density with depth. We also modify the availability and leaching of nitrate to uncover reciprocal interactions between these factors and the capture of mobile resources.Key ResultsSoils with plow pans and bulk density gradients affected overall size, distribution and carbon costs of the root system. Soils with high bulk density at depth impeded rooting depth and reduced leaching of nitrate, thereby improving the coincidence of nitrogen and root length. While increasing penetration ability of either axial or lateral root classes produced root systems of comparable net length, improved penetration of axial roots increased allocation of root length in deeper soil, thereby amplifying N acquisition and shoot biomass. Although enhanced penetration ability of both root classes was associated with greater root system carbon costs, the benefit to plant fitness from improved soil exploration and resource capture offset these.ConclusionsWhile lateral roots comprise the bulk of root length, axial roots function as a scaffold determining the distribution of these laterals. In soils with high soil strength and leaching, root systems with enhanced penetration ability of axial roots have greater distribution of root length at depth, thereby improving capture of mobile resources.