Root elongation, water stress, and mechanical impedance: a review of limiting stresses and beneficial root tip traits

Root elongation in drying soil is generally limited by a combination of mechanical impedance and water stress. Relationships between root elongation rate, water stress (matric potential), and mechanical impedance (penetration resistance) are reviewed, detailing the interactions between these closely related stresses. Root elongation is typically halved in repacked soils with penetrometer resistances >0.8-2?MPa, in the absence of water stress. Root elongation is halved by matric potentials drier than about -0.5?MPa in the absence of mechanical impedance. The likelihood of each stress limiting root elongation is discussed in relation to the soil strength characteristics of arable soils. A survey of 19 soils, with textures ranging from loamy sand to silty clay loam, found that ～10% of penetration resistances were >2?MPa at a matric potential of -10?kPa, rising to nearly 50% >2?MPa at - 200?kPa. This suggests that mechanical impedance is often a major limitation to root elongation in these soils even under moderately wet conditions, and is important to consider in breeding programmes for drought-resistant crops. Root tip traits that may improve root penetration are considered with respect to overcoming the external (soil) and internal (cell wall) pressures resisting elongation. The potential role of root hairs in mechanically anchoring root tips is considered theoretically, and is judged particularly relevant to roots growing in biopores or from a loose seed bed into a compacted layer of soil.     

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.     

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.     

Number,position, diameter and initial direction of growth of primary roots in Musa

To understand soil colonization by a root system, information is needed on the architecture of the root system. In monocotyledons, soil exploration is mainly due to the growth of adventitious primary roots. Primary root emergence in banana was quantified in relation to shoot and corm development. Root emergence kinetics were closely related to the development of aerial organs. Root position at emergence on the corm followed an asymptotic function of corm dry weight, so that the age of each root at a given time could be deduced from its position. Root diameter at emergence was related to the position of the roots on the corm, with younger roots being thicker than older ones. However, root diameters were not constant along a given root, but instead decreased with the distance to the base; roots appear to be conical in their basal and apical parts. Root growth directions at emergence were variable, but a high proportion of the primary roots emerged with a low angle to the horizontal. Further research is needed to evaluate whether these initial trajectories are conserved during root development. Results presented in this study are in good agreement with those reported for other monocotyledons such as maize and rice. They give quantitative information that will facilitate the development of models of root system architecture in banana.     

Effect of Osmotic Stress on Turgor Pressure in Mung Bean Root Cells

Turgor pressure in cells of the elongating region of intactmung bean roots was directly measured by using the pressure-probetechnique. After the external osmotic pressure had been increasedfrom 0 MPa to 0.5 MPa, turgor pressure rapidly decreased byabout 0.5 MPa from 0.65 MPa to 0.14 MPa and root elongationstopped. Subsequent turgor regulation was clearly confirmed,which followed the osmotic adjustment to maintain a constantdifference in the osmotic pressure between root-cell sap andthe external medium ( II). It took at least 6 h for turgor pressureto recover to an adjusted constant level of about 0.5 MPa dueto turgor regulation, but rootelongation resumed within onlyan hour after the osmotic treatment. Therefore, the resumptionof root elongation under osmotic stress could not have beendirectly connected with turgor regulation. Furthermore, sincethe amounts of decrease in turgor pressure just after applicationsof various degrees of osmotic stress could be interpreted inrelation to those in II, hydraulic conductivity between theinside and the outside of root cells must be large enough toattain water potential equilibrium rapidly in response to osmoticstress. We conclude that turgor pressure in the cells of theelongating region of mung bean roots is determined mainly by II because of water potential equilibrium. (Received January 27, 1987; Accepted May 21, 1987)     

Maize lateral root developmental plasticity induced by mild water stress. I: Genotypic variation across a high‐resolution series of water potentials

Lateral root developmental plasticity induced by mild water stress was examined across a high‐resolution series of growth media water potentials (Ψ_w) in two genotypes of maize. The suitability of several media for imposing near‐stable Ψ_w treatments on transpiring plants over prolonged growth periods was assessed. Genotypic differences specific to responses of lateral root growth from the primary root system occurred between cultivars FR697 and B73 over a narrow series of water stress treatments ranging in Ψ_w from ?0.25 to ?0.40 MPa. In FR697, both the average length and number of first‐order lateral roots were substantially enhanced at a Ψ_w of ?0.25 MPa compared with well‐watered controls. These effects were separated spatially, occurring primarily in the upper and lower regions of the axial root, respectively. Furthermore, first‐order lateral roots progressively increased in diameter with increasing water stress, resulting in a maximum 2.3‐fold increase in root volume at a Ψ_w of ?0.40 MPa. In B73, in contrast, the length, diameter, nor number of lateral roots was increased in any of the water stress treatments. The genotype‐specific responses observed over this narrow range of Ψ_w demonstrate the necessity of high‐resolution studies at mild stress levels for characterization of lateral root developmental plasticity.     

Root elongation is restricted by axial but not by radial pressures: so what happens in field soil?

Root distribution determines largely the zone of soil that roots have access to for water and nutrient uptake, and is of great importance especially if water and fertilizer input is restricted. Mechanical impedance is the major limitation to root elongation in many field soils. Until now, experiments have focused largely on the axial resistance to root growth. In a fascinating study of the radial forces exerted by the roots of chickpea, root extension, diameter change, and the radial forces that axially unimpeded roots exert are reported: Kolb et al. (this volume) record radial stresses of about 0.3?MPa that are broadly consistent with cell turgor pressures, but, interestingly, find no restriction to axial elongation. This result is in marked contrast to large decreases in elongation of pea radicles resulting from much smaller axial pressures reported elsewhere in the literature (e.g., an 85?% decrease in root elongation in response to axial pressures of?<?0.1?MPa). The situation is different also from that in homogeneous soil, where root penetration resistance pressures of 0.4-1.0?MPa are typically required to halt root elongation. Soil structure and strength properties will determine the balance of axial and radial pressures on an individual root tip, and hence the root elongation response. It appears that a degree of radial confinement may help roots to extend axially into hard soil. This result also complements recent findings that in strong field soils the availability of soil macropores has a large influence on regulating the root-elongation rates of seedlings.     

Field evaluation of laboratory techniques for predicting the ability of roots to penetrate strong soil and of the influence of roots on water sorptivity

The ability of two laboratory screening techniques to predict the abilities of roots of eight crop species to penetrate a compacted soil were evaluated and compared in a field experiment. A soil tilled to remove the effects of mechanical resistance was planted with the same species to serve as a control. Depth of root penetration, root density and the influence of the roots on the sorptivity of water were measured.Roots of all species penetrated deeper in the deep tilled than compacted soils. There were differences in the ability of roots of the species to penetrate the compacted soil. Generally dicotyledonous species had more roots penetrating to depth in both the compact and deep tilled soils. Within the main species classifications, lupin and safflower (dicotyledons) and oats and barley (monocotyledons) had the highest penetration into the compacted soil.Water sorptivities in the deep tilled soils were higher than those of the compact soil. Soil from planted treatments had higher sorptivities than soil which had not been planted. This is attributed to biopores left by the roots. Sorptivities of soils which had dicotyledonous species were generally higher than those of monocotyledons. The soil planted with safflower produced the highest sorptivity in the compacted layer (0.1–0.3 m).A comparison of the accuracy of the two laboratory screening methods in predicting the field penetration of roots suggest that the method involving mechanical stress was better than that involving osmotic stress. Relative root diameter was found to be a better indicator of the penetration ability of roots than relative root elongation.     

Scale of Physiological Processes Sensitivity to PEG-Induced Water Stress in Scots Pine Seedlings

The idea that water deficit strengthening induces concerted changes of plant physiological parameters is rather widespread. However, such changes are often difficult to identify due to challenges in establishments and maintenance of required water stress intensities using solid substrates. Therefore, we exposed Scots pine (Pinus sylvestris L.) seedlings to the range of water potentials from–0.15 to–1.5 MPa in PEG-water culture to identify the series of physiological parameters differently sensitive to water stress. We observed that even mild water stress (–0.15 MPa) inhibited root elongation, which could be one of the main pine seedlings vulnerabilities under drought. Active accumulation of osmolytes was already induced by mild water deficit and further increased with water stress severity. Plant fresh biomass growth sensitivity was more related to changes of relative water content (RWC) than to changes in tissue water content or dry weight accumulation. Plants were able to grow and accumulate dry weight down to–0.5 MPa, but lower medium water potentials (–1.0 and–1.5 MPa) suppressed growth and heavily damaged root cells, as judged from many-fold increase of Ca²⁺ content in roots. Chlorophyll a content was surprisingly sensitive to water stress, while carotenoids level was increased under severe stress conditions. In conclusion, the experimental system with stepwise water potential values allowed us to analyze the sensitivity scale of a range of P. sylvestris physiological processes to water stress. It was largely similar to those described earlier for other plant species, but its peculiarities were high sensitivity of root elongation, marked resistance of biomass growth to water deficit and well-developed ability to osmotic adjustment.     

Germination and seedling growth of carrots under salinity and moisture stress

Poor crop stand is a common problem in saline areas. Germination and seedling emergence may be depressed as a result of impeded aeration, saline or dry conditions. In this study, we examined the effects of salinity and moisture stress and their interactions on seed germination and seedling growth of carrots. Variable soil matric and osmotic potentials were either obtained by equilibrating soil salinized to different degrees on a 0.5 MPa ceramic plate soil moisture extractor or by adding different amounts of salt solutions to the same mass of air-dried soil, based on a previously determined soil moisture release curve, and allowing to equilibrate for 1 week. Germination decreased significantly in the investigated silty soil (Aquic Ustifluvent) at soil moisture potentials higher than −0.01 MPa, whereas osmotic potentials as low as −0.5 MPa did not influence germination. Matric potentials of −0.3 and −0.4 MPa, respectively, resulted in a strong decrease (35–95%) of germination and delayed germination by 2 to 5 days in the silty soil to which different amounts (18 and 36%, respectively) and sizes (0.8–1.2 mm and 1.5–2.2 mm, respectively) of sand particles had been added. No effect of sand and grain diameter was detected. Germination was not affected by comparable osmotic potentials. Seedling growth showed a much higher sensitivity than germination to decreasing matric potentials, but was not affected by osmotic potentials ranging from −0.05 to −0.5 MPa. Optimum shoot growth occurred at matric potentials between −0.025 and −0.1 MPa. Shoot and root growth decreased markedly at matric potentials higher than −0.01 MPa. Fresh weight of shoots decreased gradually at matric potentials lower than −0.2 MPa. Root growth was significantly increased at matric potentials of −0.1 to −0.3 MPa, whereas comparable osmotic potentials did not have equivalent effects. It is concluded that germination and seedling growth are differently affected by comparable matric and osmotic stresses and that water stress exerts a more negative effect than salt stress.     

Root elongation rate is correlated with the length of the bare root apex of maize and lupin roots despite contrasting responses of root growth to compact and dry soils

Background

We investigated interacting effects of matric potential and soil strength on root elongation of maize and lupin, and relations between root elongation rates and the length of bare (hairless) root apex.

Methods

Root elongation rates and the length of bare root apex were determined for maize and lupin seedlings in sandy loam soil of various matric potentials (?0.01 to ?1.6 MPa) and bulk densities (0.9 to 1.5 Mg m^?3).

Results

Root elongation rates slowed with both decreasing matric potential and increasing penetrometer resistance. Root elongation of maize slowed to 10 % of the unimpeded rate when penetrometer resistance increased to 2 MPa, whereas lupin elongated at about 40 % of the unimpeded rate. Maize root elongation rate was more sensitive to changes in matric potential in loosely packed soil (penetrometer resistances <1 MPa) than lupin. Despite these differing responses, root elongation rate of both species was linearly correlated with length of the bare root apex (r² 0.69 to 0.97).

Conclusion

Maize root elongation was more sensitive to changes in matric potential and mechanical impedance than lupin. Robust linear relationships between elongation rate and length of bare apex suggest good potential for estimating root elongation rates for excavated roots.     

Influence of temperature and water potential on root growth of white oak

Root growth of white oak ( Quercus alba L.) was observed under field conditions using a rhizotron. The effects of temperature, soil water potential, and leaf water potential were evaluated on three measures of root growth and development: root elongation rate, number of growing roots, and root growth intensity (sum of projected root area compared to the total root viewing area). Root elongation rate was linearly related to changes in soil temperature and soil water potential. At soil temperatures less than 17deg;C, temperature was the dominant factor affecting rate of growth, bat at temperatures greater than 17°C soil water potential became the important factor. Unlike root elongation rate, the number of growing roots and root growth intensity increased at cold soil temperatures (8°C) and at soil water potentials of-0.3 to -0.8 MPa. At high soil water potentials (-0.1 MPa) root elongation rate reached a maximum while the number of growing roots and root growth intensity were low. These differences showed that root growth and development were not exclusively affected by the soil environment. In addition, the relationship between root growth and predawn leaf water potential suggested that root growth was a contributing factor to the drought resistance of white oak.     

渗透胁迫对小麦幼苗根系呼吸的影响

用PEG—6000调节培养液的渗透势,研究了渗透胁迫对小麦幼苗根系呼吸作用的影响。在-0.5 MPa的溶液中根总呼吸强度显著降低,不同苗龄根的反应差异明显;随胁迫加强呼吸强度随之降低;根系ATP含量减少。在胁迫初期呼吸废物对呼吸强度的降低无补偿作用,而在后期(72 h后)则可提高呼吸强度。 中度水分胁迫下,HMP支路活性上升,EMP-TCAO途径活性降低;抗氰呼吸活性增大,而对氰敏感的系统活性减低;细胞色素氧化酶活性显著低于对照。     

Penetrometer resistance,root penetration resistance and root elongation rate in two sandy loam soils

Root penetration resistance and elongation of maize seedling roots were measured directly in undisturbed cores of two sandy loam soils. Root elongation rate was negatively correlated with root penetration resistance, and was reduced to about 50 to 60% of that of unimpeded controls by a resistance of between 0.26 and 0.47 MPa. Resistance to a 30° semiangle, 1 mm diameter penetrometer was between about 4.5 and 7.5 times greater than the measured root penetration resistance. However, resistance to a 5° semiangle, 1 mm diameter probe was approximately the same as the resistnace to root penetration after subtracting the frictional component of resistance. The diameter of roots grown in the undisturbed cores was greater than that of roots grown in loose soil, probably as a direct result of the larger mechanical impedance in the cores.     

Water stress induced by PEG decreases the maximum growth pressure of the roots of pea seedlings

Roots of plants growing in dry soil often experience large mechanical impedance because the decreased soil water content is associated with increased in soil strength. The combined effect of mechanical impedance and water stress hinders the establishment of seedlings in many soils, but little is known about the interaction between these two stresses. A method has been designed that, for the first time, measured the maximum axial force exerted by a root growing under controlled water stress. Using this technique the axial force exerted by a pea radicle was measured using a shear beam, while the seedling was suspended in an aerate solution of polyethylene glycol 20 000 at osmotic potentials between 0 and -0.45 MPa. The maximum growth force was then divided by the cross-sectional area of the root to give the maximum axial growth pressure. The value of maximum axial growth pressure decreased linearly from 0.66 and 0.35 MPa as the osmotic potentials of the solution of PEG decreased from 0 to -0.45 MPa. In dry soil, therefore, the maximum strength of soil that a root can penetrate is decreased because of the decrease in maximum growth pressure. The elongation rates of unimpeded roots were similar whether the roots were subject to either a matric potential in soil or to an osmotic potential in a solution of PEG.Key words: Pisum sativum L, pea, mechanical impedance, axial growth pressure, water stress, PEG 20 000.      

The Influence of Growing Temperature on the Growth and Morphology of Cereal Seedling Root Systems

The root systems of cereal seedlings (wheat, rye, barley, oats),grown in a series of liquid and solid media at 5, 15 and 25?C, were measured at intervals between sowing and emergence(shoot length 4.0 cm). In all cases, the length of seminal axesincreased linearly with time, and the rate of root extensionwas increased significantly by each 10 ?C increment in growingtemperature. In general, extension rates were lower in the solidmedia than in water culture; however, there was a strong interactionbetween medium and growing temperature, indicating that highertemperatures can compensate partly for the detrimental effectsof mechanical impedance. Examination of the influence of growing temperature upon rootaxis diameter was hampered by the observation that, for allfour species, apical root diameter was inversely related toroot length. It was concluded that comparison of the diametersof roots grown at different temperatures was valid only if theroots were of the same length (but different ages). Followingthis principle, it was found that, although cereal roots grownat 5 ?C tended to be slightly thicker than those at 15 ?C or25 ?C, this effect was small and rarely exceeded 10%. Publishedreports of very large differences in root diameter at differentgrowing temperatures were examined and found to be generallyunreliable. Lateral roots did not represent a significant fractionof the root system in any of the treatments studied. Root: shoot relationships at different growing temperatureswere examined by plotting root system length against shoot lengthfor each sample. It was found that, for the same shoot length,the root systems at 5 ?C were much shorter than at 25 ?C, theresponse at 15 ?C being rather more variable. This finding,which indicates that plants grown at low temperatures have alower capacity for water and solute uptake than those grownat laboratory temperatures, has important implications for thedesign of laboratory experiments. The influence of root zonetemperature upon root growth and physiology can be determinedunequivocably only if the roots have been exposed to the experimentaltemperature throughout their growth. Key words: Barley, Oats, Rye, Wheat, Root length, Root diameter, Root, shoot ratio     

Fine-root morphological and growth traits in a Turkey-oak stand in relation to seasonal changes in soil moisture in the Southern Apennines, Italy

We investigated the effects of seasonal changes in soil moisture on the morphological and growth traits of fine roots (<2?mm in diameter) in a mature Turkey-oak stand (Quercus cerris L.) in the Southern Apennines of Italy. Root samples (diameter:?<0.5, 0.5?C1.0, 1.0?C1.5, and 1.5?C2.0?mm) were collected with the Auger method. Mean annual fine-root mass and length on site was 443?g?m^?2 (oak fine roots 321?g?m^?2; other species 122?g?m^?2) and 3.18?km?m^?2 (oak fine roots 1.14?km?m^?2; other species 2.04?km?m^?2), respectively. Mean specific root length was 8.3?m?g^?1. All fine-root traits displayed a complex pattern that was significantly related to season. In the four diameter classes, both fine-root biomass and length peaked in summer when soil water content was the lowest and air temperature the highest of the season. Moreover, both fine-root biomass and length were inversely related with soil moisture (p?<?0.001). The finest roots (<0.5?mm in diameter) constituted an important fraction of total fine-root length (79?%), but only 21?% of biomass. Only in this root class, consequent to change in mean diameter, specific root length peaked when soil water content was lowest showing an inverse relationship (p?<?0.001). Furthermore, fine-root production and turnover decreased with increasing root diameter. These results suggest that changes in root length per unit mass, and pulses in root growth to exploit transient periods of low soil water content may enable trees to increase nutrient and water uptake under seasonal drought conditions.     

Root responses of flood-tolerant and flood-sensitive tree species to soil redox conditions

Summary Under controlled rhizotron conditions, roots of Taxodium distichum L., Quercus lyrata Walt, and Q. falcata var. pagodaefolia Ell. were subjected to low soil redox potentials. Root elongation was inhibited at low soil redox potentials. In T. distichum, redox potentials below +200 mV resulted in a significant inhibition of root elongation. In Q. falcata var. pagodaefolia and Q. lyrata, redox potentials below +350 mV resulted in complete cessation of root growth. Studies on root anatomy indicated that low soil redox potenials resulted in a changed cellular structure in the cortex of T. distichum. However, little change was noted in stress roots of oak species. Alcohol dehydrogenase activity in T. distichum roots was approximately doubled compared to control plants, indicating stimulated alcoholic fermentation. In T. distichum, alcoholic fermentation and anatomical changes contribute to flood tolerance but oak species lack these characteristics.     

A biophysical analysis of root growth under mechanical stress

The factors controlling root growth in hard soils are reviewed alongside summarised results from our recent studies. The turgor in cells in the elongation zone of roots pushes the apex forward, resisted by the external pressure of the soil and the tension in the cell walls. The external pressure of the soil consists of the pressure required to deform the soil, plus a component of frictional resistance between the root and soil. This frictional component is probably small due to the continuous sloughing of root cap cells forming a low-friction lining surrounding the root. Mechanically impeded roots are not only thicker, but are differently shaped, continuing to increase in diameter for a greater distance behind the root tip than in unimpeded roots. The osmotic potential decreases in mechanically impeded roots, possibly due to accumulation of solutes as a result of the slower root extension rate. This more negative osmotic potential is not always translated into increased turgor pressure, and the reasons for this require further investigation. The persistent effect of mechanical impedance on root growth is associated with both a stiffening of cell walls in the axial direction, and with a slowing of the rate of cell production.     

When the going gets tough: Responses of Sorghum bicolor L. (cv. Tegemeo) seedlings to soil penetration resistance

Summary

Sorghum bicolor L. (cv. Tegemeo) seedlings were grown for nine days in soil at field capacity packed to give a uniform penetration resistance (PR) of either 0.25, 1.00 or 1.75 MPa. Root biomass was not significantly affected by soil PR treatment. However, as PR increased to 1.75 MPa, the diameter of the seminal root axis increased by 52% whilst its length decreased by 30%. Shoot growth, in terms of oven dry (OD) weight and photo-synthetic area, was reduced in both the 0.25 MPa and 1.75 MPa treatments compared to the 1.00 MPa treatment. A reduced nutrient, water or oxygen supply to the roots were discounted as possible causes of the root and shoot responses to soil PR. It is suggested that the changes in root morphology between treatments were a direct result of the changes in soil PR. For shoot growth, in the 0.25 MPa treatment it is suggested that shoot growth was reduced as a result of an increase in the carbon sink strength of the roots.