Blair M. McKenzie

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.

The dynamics of root meristem distribution in the soil

Plants must develop efficient root architectures to secure access to nutrients and water in soil. This is achieved during plant development through a series of expansion and branching processes, mostly in the proximity of root apical meristems, where the plant senses the environment and explores immediate regions of the soil. We have developed a new approach to study the dynamics of root meristem distribution in soil, using the relationship between the increase in root length density and the root meristem density. Initiated at the seed, the location of root meristems in barley seedlings was shown to propagate, wave-like, through the soil, leaving behind a permanent network of roots for the plant to acquire water and nutrients. Data from observations on barley roots were used to construct mathematical models to describe the density of root meristems in space. These models suggested that the morphology of the waves of meristems was a function of specific root developmental processes. The waves of meristems observed in root systems of barley seedlings exploring the soil might represent a more general and fundamental aspect of plant rooting strategies for securing soil resources.

Influence of drying and ageing on the stabilization of earthworm (Lumbricidae) casts

Abstract The influence of drying and ageing on the stabilization of casts produced by the endoge´ic earthworm, Aporrectodea rosea, from a soil, which was hard-setting and low in organic matter, were investigated in the laboratory. Casts and uningested soil were aged-most for up to 32 days, dried for up to 21 days, or subjected to different wetting and drying cycles over 30 days. The dispersion index of aged-moist casts decreased from 0.40 to 0.25 over 32 days, while dispersion index of dried casts decreased from 0.40 to 0.01 over 21 days. The dispersion index of air-dried casts was not significantly increased by five cycles of wetting and drying. The dispersion index of dried casts was not significantly less than that of dried soil. In soils wetter than a matric potential of approximately –35kPa, stabilization of casts was probably due to a combination of cohesion of soil particles, age-hardening and growth of microorganisms. However, in soils drier than –35kPa, cementation was probably the major mechanism of stabilization. The addition of wheat straw to the soil prior to ingestion by earthworms increased dispersion from aged-moist casts, but did not influence dispersion from dried casts. The addition of wheat straw decreased the number of air-dried casts which slaked severely. The concentration of soluble carbohydrate decreased with dispersion index as casts and uningested soil were each dried. This suggested that soluble carbohydrate may have been denatured with or without being bonded to soil particles during drying.

Quantitative image analysis of earthworm-mediated soil displacement

The way soil is disrupted and deformed by earthworm movement is hard to quantify non-destructively. Two anecic earthworm species, Aporrectodea longa (Ude) and Lumbricus terrestris L., were compared in their effect on the soil around them as they burrow. Image analysis (particle image velocimetry, PIV) was used successfully to quantify the distance and direction of soil particle displacement by earthworm locomotion giving a unique insight into their effect on the soil structure development. The data collected from both species using PIV show a decline in soil displacement at increasing distance from the earthworm’s body. The quantity of soil being displaced was more to the sides of the earthworms (radially) than in front of the prostomium (axially). Also, L. terrestris displaced more soil than A. longa both axially and radially. The findings from this study suggest that PIV image analysis is a viable tool for examining soil displacement by earthworms and the method used has the potential to be applied to other soil organisms.

Do the roots of perennial ryegrass elongate to biopores filled with the casts of endogeic earthworms

Abstract Earthworm casts deposited in burrows may stimulate plant roots to elongate preferentially to those burrows. A model experiment was conducted with an artificial, impenetrable subsoil in which vertical holes were left empty, filled with earthworm casts, or filled with fertile soil. The roots of perennial ryegrass had elongated down through an aggregate seedbed and were deflected horizontally when they reached the artificial subsoil. After six days, approximately 6% of the roots that were deflected sideways, met with holes in the subsoil. However, the number of roots that met with holes filled with either casts, or soil, was not significantly different from the number of roots that met with empty holes. Under the conditions of our model system, there was no evidence that ryegrass roots elongated preferentially to and entered holes filled with earthworm casts.

Relationships between casts of geophagous earthworms (Lumbricidae, Oligochaeta) and matric potential

Laboratory experiments were conducted to investigate the effects of matric potential on the rate of cast production, and water content and bulk density of casts. Fresh, subsurface, and surface casts were collected from Aporrectodea caliginosa, A rosea and A. trapezoides kept in reconstituted soil cores at matric potentials between-25 and -2 kPa. Rates of cast production and water contents of fresh casts, subsurface casts, and surface casts increased as the matric potential of soil cores increased from -25 kPa to -2 kPa. The bulk density of fresh casts decreased as the matric potential of soil cores increased from -8.6 to -2 kPa. Aestivating earthworms were found in soil cores at all matric potentials except-2 kPa. The rate of cast production at the drier matric potentials may have been related to the water balance of the earthworms. As the matric potential of the soil cores decreased, the amount of water egested per cast remained constant, but overall the water egested per day decreased due to a reduction in the number of casts produced. Differences between the three species were identified in rates of cast production, with A. rosea generally producing more casts than either A. caliginosa or A. trapezoides, and in water contents and bulk densities of fresh casts.

Root hairs aid soil penetration by anchoring the root surface to pore walls

Highlight Root hairs anchored maize roots to the sides of pore walls, helping root tips to penetrate soil at intermediate soil bulk densities, shown by comparing wild-type seedlings with hairless mutants.

Destabilization of soil during the production of earthworm (Lumbricidae) and artificial casts

Abstract The mechanisms by which soil is destabilized in the digestive tract of endogéic earthworms were investigated with artificial casts, which were moulded with a syringe from slurries of a silty loam with or without gypsum and organic matter treatments, and compared to casts produced by Aporrectodea rosea (Lumbricidae). Both types of casts generally had the same levels of mechanical dispersion, observations of slaking, and particle size distribution when the casts were fresh, aged-moist for 30 days, or air-dried. Fresh casts were significantly more dispersive than the uningested soil despite the addition of gypsum or organic matter to the soil. However, the dispersion from aged-moist or air-dried casts was not substantially greater than that of uningested soil. Air-drying was more effective than moist-ageing in increasing the stability of casts and uningested soil. The concentration of soluble carbohydrate was greater in artificial casts produced from soil treated with sheep dung or xanthan gum, and in earthworm casts produced from soil treated with xanthan, than in the uningested soil of the same treatments. An increase in the concentration of soluble carbohydrate was related to an increase in dispersion. An attempt was made to simulate the addition of mucus to soil in the digestive tract of earthworms, by the addition of sucrose or xanthan gum to the slurry during the production of artificial casts. The addition of xanthan, but not sucrose, to the slurry increased mechanical dispersion relative to that of the uningested soil in the fresh treatment. Although the production of artificial casts destabilized soil to the same degree as earthworm casts, the artificial casts did not simulate all chemical, biochemical, and microbiological aspects of digestion.

Volume density of earthworm burrows in compacted cores of soil as estimated by direct and indirect methods

After earthworms of the species Aporrectodea caliginosa and A. rosea had burrowed in compacted cores of soil for 68 days the cores were sectioned horizontally. The upper surface of each sectioned layer of soil was photographed before it was dissected and the dimensions of all burrows within the layer measured. Volume densities calculated from the direct measurement of burrows were compared with the values calculated by stereology from data obtained from two image analysis methods, computerised image analysis and point counting with a systematic lattice. The assumption implicit in all stereology calculations was satisfied for this experiment in that the burrows of both species showed no preferred orientation in the compacted soil. Computerised image analysis could not measure the density of all burrows in the photographs because of the lack of contrast between cast-filled burrows and the soil and the complex shapes of the burrows. Although the volume densities of A. caliginosa burrows calculated from point counts were correlated with the values calculated from direct measurement, point counting over-estimated volume densities by two to three times. In the experiment, A. rosea produced too few airfilled burrows for the lattice to detect. The relative ratios of air-filled to cast-filled burrows calculated from the point counts suggested that approximately two-thirds and eight-ninths of the burrows of A. caliginosa and A. rosea, respectively, were filled with casts.

Liming impacts on soils, crops and biodiversity in the UK: A review.

Fertile soil is fundamental to our ability to achieve food security, but problems with soil degradation (such as acidification) are exacerbated by poor management. Consequently, there is a need to better understand management approaches that deliver multiple ecosystem services from agricultural land. There is global interest in sustainable soil management including the re-evaluation of existing management practices. Liming is a long established practice to ameliorate acidic soils and many liming-induced changes are well understood. For instance, short-term liming impacts are detected on soil biota and in soil biological processes (such as in N cycling where liming can increase N availability for plant uptake). The impacts of liming on soil carbon storage are variable and strongly relate to soil type, land use, climate and multiple management factors. Liming influences all elements in soils and as such there are numerous simultaneous changes to soil processes which in turn affect the plant nutrient uptake; two examples of positive impact for crops are increased P availability and decreased uptake of toxic heavy metals. Soil physical conditions are at least maintained or improved by liming, but the time taken to detect change varies significantly. Arable crops differ in their sensitivity to soil pH and for most crops there is a positive yield response. Liming also introduces implications for the development of different crop diseases and liming management is adjusted according to crop type within a given rotation. Repeated lime applications tend to improve grassland biomass production, although grassland response is variable and indirect as it relates to changes in nutrient availability. Other indicators of liming response in grassland are detected in mineral content and herbage quality which have implications for livestock-based production systems. Ecological studies have shown positive impacts of liming on biodiversity; such as increased earthworm abundance that provides habitat for wading birds in upland grasslands. Finally, understanding of liming impacts on soil and crop processes are explored together with functional aspects (in terms of ecosystems services) in a new qualitative framework that includes consideration of how liming impacts change with time. This holistic approach provides insights into the far-reaching impacts that liming has on ecosystems and the potential for liming to enhance the multiple benefits from agriculturally managed land. Recommendations are given for future research on the impact of liming and the implications for ecosystem services.