Soil structure and plant growth: Impact of bulk density and biopores

Compacted soils are not uniformly hard; they usually contain structural cracks and biopores, the continuous large pores that are formed by soil fauna and by roots of previous crops. Roots growing in compacted soils can traverse otherwise impenetrable soil by using biopores and cracks and thus gain access to a larger reservoir of water and nutrients. Experiments were conducted in a growth chamber to determine the plant response to a range of uniform soil densities, and the effect of artificial and naturally-formed biopores. Barley plants grew best at an intermediate bulk density, which presumably represented a compromise between soil which was soft enough to allow good root development but sufficiently compact to give good root-soil contact. Artificial 3.2 mm diameter biopores made in hard soil gave roots access to the full depth of the pot and were occupied by roots more frequently than expected by chance alone. This resulted in increased plant growth in experiments where the soil was allowed to dry. Our experiments suggest that large biopores were not a favourable environment for roots in wet soil; barley plants grew better in pots containing a network of narrow biopores made by lucerne and ryegrass roots, responded positively to biopores being filled with peat, and some pea radicles died in biopores. A theoretical analysis of water uptake gave little support to the hypothesis that water supply to the leaves was limiting in either very hard or very soft soil. The net effect of biopores to the plant would be the benefits of securing extra water and nutrients from depth, offset by problems related to poor root-soil contact in the biopore and impeded laterals in the compacted biopore walls.     

Maize stomatal conductance in the field: its relationship with soil and plant water potentials, mechanical constraints and ABA concentration in the xylem sap

Abstract. Stomatal conductance, leaf water potential, soil water potential and concentration of abscisic acid (ABA) in the xylem sap were measured on maize plants growing in the field, in two treatments with contrasting soil structures. Soil compaction affected the stomatal conductance, but this effect was no longer observed if the soil water potential was increased by irrigation. Differences in leaf water potential did not account for the differences in conductance between treatments. Conversely, the relationship between stomatal conductance and concentration of ABA in the xylem sap was consistent during the experiment. The proposed interpretation is that stomatal conductance was controlled by the root water potential via an ABA message. Control of the stomatal conductance by the leaf water potential or by an effect of mechanical stress on the roots is unlikely.     

Soil solute concentration and water uptake by single lupin and radish plant roots

Application of computer assisted tomography to gamma and X-ray attenuation measurements and Na⁺-LIX microelectrodes were used to determine the spatial distributions of soil water content and Na⁺ concentrations respectively near single roots of eighteen day old lupin and radish plants. These quantities were monitored at root depths of 3, 6 and 9 cm and at zero, 2, 4, 6, and 8 hour intervals from the diurnal commencement of transpiration. The plants were subjected to two levels of transpirational demand and five Na⁺ soil solution concentration levels. Water extraction rates for the lupin and radish roots increased continuously with time but were substantially reduced with increasing Na⁺ concentration in the treatment. Water uptake was uniform along the length of the essentially constant diameter lupin roots but decreased along the tapering radish roots as the diameter and hence the surface area per unit length of the roots decreased. The accumulation of Na⁺ at the root surfaces of both plants increased gradually with time in a near linear fashion and was slightly higher under the higher transpiration demand. These increases were not exponential as would be expected with non-absorption by the roots and this is considered to be due to back diffusion at the relatively high water contents used. At these water contents matric potentials had a much smaller influence on transpiration than osmotic potentials. The relationships between leaf water potentials (Ψ₁) and osmotic potentials at the root surfaces were linear with the decreases in Ψ₁ almost exactly reflecting the decreases in Ψ_π indicating rapid plant adjustment. Leaf water potentials decreased progressively with time and the relationships between leaf water potential and the transpiration rate were also linear supporting the suggestion of constant plant resistances at any given concentration.     

Structure,ecology and physiology of root clusters – a review

Hairy rootlets, aggregated in longitudinal rows to form distinct clusters, are a major part of the root system in some species. These root clusters are almost universal (1600 species) in the family Proteaceae (proteoid roots), with fewer species in another seven families. There may be 10–1000 rootlets per cm length of parent root in 2–7 rows. Proteoid roots may increase the surface area by over 140× and soil volume explored by 300× that per length of an equivalent non-proteoid root. This greatly enhances exudation of carboxylates, phenolics and water, solubilisation of mineral and organic nutrients and uptake of inorganic nutrients, amino acids and water per unit root mass. Root cluster production peaks at soil nutrient levels (P, N, Fe) suboptimal for growth of the rest of the root system, and may cease when shoot mass peaks. As with other root types, root cluster production is controlled by the interplay between external and internal nutrient levels, and mediated by auxin and other hormones to which the process is particularly sensitive. Proteoid roots are concentrated in the humus-rich surface soil horizons, by 800× in Banksia scrub-heath. Compared with an equal mass of the B horizon, the A₁ horizon has much higher levels of N, P, K and Ca in soils where species with proteoid root clusters are prominent, and the concentration of root clusters in that region ensures that uptake is optimal where supply is maximal. Both proteoid and non-proteoid root growth are promoted wherever the humus-rich layer is located in the soil profile, with 4× more proteoid roots per root length in Hakea laurina. Proteoid root production near the soil surface is favoured among hakeas, even in uniform soil, but to a lesser extent, while addition of dilute N or P solutions in split-root system studies promotes non-proteoid, but inhibits proteoid, root production. Local or seasonal applications of water to hakeas initiate non-proteoid, then proteoid, root production, while waterlogging inhibits non-proteoid, but promotes proteoid, root production near the soil surface. A chemical stimulus, probably of bacterial origin, may be associated with root cluster initiation, but most experiments have alternative interpretations. It is possible that the bacterial component of soil pockets rich in organic matter, rather than their nutrient component, could be responsible for the proliferation of proteoid roots there, but much more research on root cluster microbiology is needed.     

Soil water dynamics in an oak stand

A model describing water uptake by plants with particular attention to the soil-root interface under transient conditions is derived and discussed. Field data on a daily scale enable the unknown parameters of the model to be determined with the help of an identification technique. The model is then used to analyse the experimental results presented in part I of this paper. The loss of total conductivity of the soil-tree system under drought conditions whereas the metabolism of the trees seems to remain unaffected can be explained by the increase of the soil-root resistance. In fact this resistance becomes the limiting factor when the volumetric soil water content decreases (below =0.33 for the superficial layer and 0.36 for the deeper one in the studied case). Such values can be frequently encountered at the end of summer.     

Water uptake profile response of corn to soil moisture depletion

The effects of soil moisture distribution on water uptake of drip‐irrigated corn were investigated by simultaneously monitoring the diurnal evolution of sap flow rate in stems, of leaf water potential, and of soil moisture, during intervals between successive irrigations. The results invalidate the steady‐state resistive flow model for the continuum. High hydraulic capacitance of wet soil and low hydraulic conductivity of dry soil surrounding the roots damped significantly diurnal fluctuations of water flow from bulk soil to root surface. By contrast, sap flow responded directly to the large diurnal variation of leaf water potential. In wet soil, the relation between the diurnal courses of uptake rates and leaf water potential was linear. Water potential at the root surface remained nearly constant and uniformly distributed. The slope of the lines allowed calculating the resistance of the hydraulic path in the plant. Resistances increased in inverse relation with root length density. Soil desiccation induced a diurnal variation of water potential at the root surface, the minimum occurring in the late afternoon. The increase of root surface water potential with depth was directly linked to the soil desiccation profile. The development of a water potential gradient at the root surface implies the presence of a significant axial resistance in the root hydraulic path that explains why the desiccation of the soil upper layer induces an absolute increase of water uptake rates from the deeper wet layers.     

Soil and plant water stress in an Appalachian oak forest in relation to topography and stand age

Summary A Forest Site Quality Index (FSQI) formulated to predict site quality in Ridge and Valley terrain based on the topographic parameters of aspect, slope inclination and slope position was used to verify moisture gradients along the southeast face of Potts Mountain in Craig County, Virginia. A gradient of site quality index values representing xeric to mesic sites was established in both recently clearcut and adjacent uncut second-growth forest stands. Soil moisture content was determined gravimetrically at ten day intervals from May to October, 1981. Plant moisture stress measurements were taken in conjunction with soil moisture sampling using the pressure chamber technique on three dominant hardwood tree species.For both clearcut and uncut forest stands, a general gradient of increasing soil moisture availability with increasing FSQI was evident, although differences were not large between index values of 8 and 11 in either stand type. Soil water potential and predawn plant water potential exhibited a strong seasonal trend, their direct relationship suggesting that available soil water is probably the critical factor controlling base P levels. Growth limiting stress levels began in late July and continued for the remainder of the growing season.Funding for this research was granted through Cooperative Research Agreement # 18-882, USDA SE Forest Experiment Station and the Forestry Department of Virginia Polytechnic Institute and State University, Blacksburg, VA 24061.     

Root morphology, hydraulic conductivity and plant water relations of high-yielding rice grown under aerobic conditions

Background and Aims

Increasing physical water scarcity is a major constraint for irrigated rice (Oryza sativa) production. ‘Aerobic rice culture’ aims to maximize yield per unit water input by growing plants in aerobic soil without flooding or puddling. The objective was to determine (a) the effect of water management on root morphology and hydraulic conductance, and (b) their roles in plant–water relationships and stomatal conductance in aerobic culture.

Methods

Root system development, stomatal conductance (g_s) and leaf water potential (Ψ_leaf) were monitored in a high-yielding rice cultivar (‘Takanari’) under flooded and aerobic conditions at two soil moisture levels [nearly saturated (> –10 kPa) and mildly dry (> –30 kPa)] over 2 years. In an ancillary pot experiment, whole-plant hydraulic conductivity (soil-leaf hydraulic conductance; K_pa) was measured under flooded and aerobic conditions.

Key Results

Adventitious root emergence and lateral root proliferation were restricted even under nearly saturated conditions, resulting in a 72–85 % reduction in total root length under aerobic culture conditions. Because of their reduced rooting size, plants grown under aerobic conditions tended to have lower K_pa than plants grown under flooded conditions. Ψ_leaf was always significantly lower in aerobic culture than in flooded culture, while g_s was unchanged when the soil moisture was at around field capacity. g_s was inevitably reduced when the soil water potential at 20-cm depth reached –20 kPa.

Conclusions

Unstable performance of rice in water-saving cultivations is often associated with reduction in Ψ_leaf. Ψ_leaf may reduce even if K_pa is not significantly changed, but the lower Ψ_leaf would certainly occur in case K_pa reduces as a result of lower water-uptake capacity under aerobic conditions. Rice performance in aerobic culture might be improved through genetic manipulation that promotes lateral root branching and rhizogenesis as well as deep rooting.     

X-ray absorption and phase contrast imaging to study the interplay between plant roots and soil structure

Plant performance is, at least partly, linked to the location of roots with respect to soil structure features and the micro-environment surrounding roots. Measurements of root distributions from intact samples, using optical microscopy and field tracings have been partially successful but are imprecise and labour-intensive. Theoretically, X-ray computed micro-tomography represents an ideal solution for non-invasive imaging of plant roots and soil structure. However, before it becomes fast enough and affordable or easily accessible, there is still a need for a diagnostic tool to investigate root/soil interplay. Here, a method for detection of undisturbed plant roots and their immediate physical environment is presented. X-ray absorption and phase contrast imaging are combined to produce projection images of soil sections from which root distributions and soil structure can be analyzed. The clarity of roots on the X-ray film is sufficient to allow manual tracing on an acetate sheet fixed over the film. In its current version, the method suffers limitations mainly related to (i) the degree of subjectivity associated with manual tracing and (ii) the difficulty of separating live and dead roots. The method represents a simple and relatively inexpensive way to detect and quantify roots from intact samples and has scope for further improvements. In this paper, the main steps of the method, sampling, image acquisition and image processing are documented. The potential use of the method in an agronomic perspective is illustrated using surface and sub-surface soil samples from a controlled wheat trial. Quantitative characterization of root attributes, e.g. radius, length density, branching intensity and the complex interplay between roots and soil structure, is presented and discussed. This revised version was published online in June 2006 with corrections to the Cover Date.     

Relationships between plant and soil water status in vine (Vitis vinifera L.)

Soil water status and its effect on plant water status are commonly evaluated for water stress diagnosis in annual crops. We investigated the application of this method to vineyards, using the fraction of transpirable soil water (FTSW) to characterise the soil water deficit experienced by the plant. The stability of the relationship between FTSW and predawn leaf water potential (Ψp) was analysed over two years (2000–2001), in two contrasted soils in vineyards in south eastern France, both planted with the cultivar Syrah, but grafted on different rootstocks (SO4 and 140Ru). FTSW was determined from soil moisture measurements performed with a neutron probe down to 2.5 m, under the rows and between the rows (3 replicates in each case). Vertical and horizontal variations in soil water content were analysed and the upper and lower limits of total vine’s transpirable soil water (TTSW) were calculated for each soil. The lower limit was also compared with the value of soil moisture content determined at −1.5 MPa in the laboratory. FTSW could be calculated for the soil depth analysed, without distinguishing horizontal position (row or inter-row). The lower limit of TTSW for vine was higher than the soil water content at −1.5 MPa, except in the upper horizons (0–0.2 m) which are prone to soil evaporation. A single relationship between Ψp and FTSW was obtained for the two vineyards and for the two years of measurement. This relationship was similar to that established by Lebon et al. (2003) on Gewürztraminer/SO4 in a vineyard in northern France. FTSW can therefore be used as an indicator of the water deficit experienced in vineyards, provided that TTSW is correctly estimated.     

Modelling non-uniform soil water uptake by a single plant root

The assumption of uniform water flow to the root or uniform water potential at the root surface was shown by Hainsworth and Aylmore (1986, 1989) to be erroneous. The present paper demonstrates how the non-uniform uptake of water by a single root can be modeled. Differential equations are numerically solved to describe simultaneous water movement in the plant and in the soil. In the plant, boundary conditions are the water potentials at the root surface (Ψ_s) and in the xylem at the root base (Ψ_b). A set of difference equations describe the flow of water radially through the cortex to the xylem and in the xylem axially upwards to the base. For calculating the water flow in the soil and the values of Ψ_s, i.e. the boundary conditions for flow in the root, the finite element method (FEM) is used, the boundary conditions being the flux of water into the plant root and the zero flow across the wall, bottom and surface of a hypothetical soil cylinder surrounding the root. ei]Section editor: B E Clothier     

Salt-tolerant and plant growth-promoting bacteria isolated from Zn/Cd contaminated soil: identification and effect on rice under saline conditions

The bacteria of PDMCd0501, PDMCd2007, and PDMZnCd2003 were isolated from a Zn/Cd contaminated soil. They were classified as salt-tolerant bacteria in this experiment. The bacteria had indole-3-acetic acids (IAA) production, nitrogen fixation, and phosphate solubilization, under 8% (w/v) NaCl condition. Biochemical test (API 20E) and 16S rDNA sequencing identified PDMCd2007 and PDMCd0501 as Serratia sp. and PDMZnCd2003 was Pseudomonas sp. The effect of Pseudomonas sp. PDMZnCd2003 on the germination and seedlings of Oryza sativa L.cv. RD6 was determined under a salinity of 0–16 dS/m. The salinity levels of 4–16 dS/m affected to decrease germination and seedlings of rice. Comparison between uninoculated and inoculated system, however, Pseudomonas sp. PDMZnCd2003 had a negative impact on the rice growth. This unexpected effect was a case that should be concerned and studied further before application as a plant growth-promoting bacteria (PGPB).     

Plant and soil resistance to water flow in faba bean (Vicia faba L. major Harz.)

An experiment was conducted to determine soil and plant resistance to water flow in faba bean under field conditions during the growing season. During each sampling period transpiration flux and leaf water potential measured hourly were used with daily measurements of root and soil water potential to calculate total resistance using Ohm's law analogy. Plant growth, root density and soil water content distributions with depth were measured. Leaf area and root length per plant reached their maximum value during flowering and pod setting (0.31 m² and 2200 m, respectively), then decreasing until the end of the growing period. Root distribution decreased with depth ranging, on average, between 34.2% (in the 0–0.25 m soil layer) and 18.1% (in the 0.75–1.0 m soil layer). Mean root diameter was 0.6 mm but most of the roots were less than 0.7 mm in diameter. Changes in plant and soil water potentials reflected plant growth characteristics and climatic patterns. The overall relationship between the difference in water potential between soil and leaf and transpiration was linear, with the slope equal to average plant resistance (0.0165 MPa/(cm³ m^-1 h^-1 10^-3). Different regression parameters were obtained for the various measurement days. The water potential difference was inversely related to transpiration at high leaf stomatal resistance and at high values of VPD. Total resistance decreased with transpiration flux in a linear relationship (r=−0.68). Different slope values were obtained for the different measurement days. Estimated soil resistance was much lower than the observed total resistance to water flow. The change from vegetative growth to pod filling was accompanied by an increase in plant resistance. The experimental results support previous findings that resistance to water flow through plants is not constant but is influenced by plant age, growth stage and environmental conditions. A more complex model than Ohm's law analogy may be necessary for describing the dynamic flow system under field conditions. This revised version was published online in June 2006 with corrections to the Cover Date.     

Drought adaptation of field grown wheat in relation to soil physical conditions

In order to investigate the effects of soil texture on possible non-hydraulic signals under field conditions, spring wheat plants (Triticum aestivum L. cv. Cadensa) grown in sand and loam soils and with a well developed root system were exposed to slow soil drying in the late vegetative stage of growth. Soil water potential and content were measured daily at different depths and plant responses were measured in flag leaves. When the average soil water potential in the top soil layers (0–25 cm depth in sand and 0–45 cm depth in loam) dropped to –60 or –70 kPa and the lower soil layers were still at field capacity, morning xylem [ABA] (0.03–0.04 vs. 0.06–0.08 mmol m-3) and midday leaf ABA concentration increased (250–300 vs. 400–450 ng/g DW) and leaf conductance decreased relatively to well-watered (control) plants (0.75–0.88 vs. 0.64–0.70 mol m-2 s-1). These responses took place before any decrease in leaf water potential occurred as compared with control plants, indicating that they were triggered by root-borne signals due to reduced root water status in the top soil layers. At this stage the soil water content was as low as 6% by volume, the fraction of roots in ‘wet’ soil was 0.12 and relative available soil water was 45% in sand and still high 20%, 0.48 and 70%, respectively, in loam of the whole soil profile indicating that roots were responding to soil water availability and not soil water content at a certain evaporative demand. In addition, similar responses occurred at high and low evaporative demands (3.4–5.2 vs. 0.6–4.0 mm/day of potential evapotranspiration). This revised version was published online in July 2006 with corrections to the Cover Date.     

Root:soil adhesion in the maize rhizosphere: the rheological approach

This study was designed to investigate the strength of attachment of plant seedling roots to the soil in which they were grown. The study also assessed the effects of differing soil textures and differing soil matric potentials upon the strength of the root:soil attachment. A device for growing roots upon a soil surface was designed, and was used to produce roots which were attached to the soil. In order to quantify root:soil adhesion, roots of maize seedlings, grown on the soil surface, were subsequently peeled off using a universal test machine, in conjunction with simultaneous time-lapse video observation. To clarify the partitioning of energy in the root:soil peeling test, separate mechanical tests on roots, and on two adherent remoulded topsoil balls were also carried out. The seedling root was characterised by a low bending stiffness. The energy stored in bending was negligible, compared to the root:soil adhesion energy. The mechanical properties of two adherent remoulded topsoil balls were a decrease of the soil:soil adhesion energy as the soil:soil plastic energy increased. These two parameters were therefore interdependent. Using a video-camera system, it was possible to separate the different processes occurring during the root:soil peeling test, in particular, the seed:soil adhesion and the root:soil soil adhesion. An interpretation of the complex and variable force:displacement curves was thus possible, enabling calculation of the root:soil interfacial rupture energy. At a given suction (10 kPa), the results of the peeling test showed a clear soil texture effect on the value of the root:soil interfacial rupture energy. In contrast, for the same silty topsoil, the effect of the soil water suction on the value of the interfacial rupture energy was very moderate. The root:soil interfacial rupture energy was controlled mainly by a product of microscopic soil specific surface area and the macroscopic contact surface area between the root and the soil. Biological and physical interactions contributing to root:soil adhesion such as root:soil interlocking mechanics were also analysed and discussed. This revised version was published online in June 2006 with corrections to the Cover Date.     

Rhizoplane colonization of pea seedlings by Rhizobium leguminosarum and a deleterious root colonizing Pseudomonas sp. and effects on plant growth

Some pseudomonads produce a toxin that specifically inhibits winter wheat (Triticum aestivum L.) root growth and the growth of several microorganisms. The toxin does not inhibit pea (Pisum sativum) root growth, but the organisms are aggressive root colonizers and their effect on Rhizobium leguminosarum growth, colonization, and nodulation of peas was not known. Peas were grown in Leonard jars in the greenhouse. Pea roots were inoculated with R. leguminosarum, a toxin-producing Pseudomonas sp., both, or neither (control). The Pseudomonas sp. colonized pea roots more rapidly and in greater number than R. leguminosarum after ten days. In the presence of the Pseudomonas sp., the R. leguminosarum population on the rhizoplane was less at ten days. When the roots were inoculated with both R. leguminosarum and Pseudomonas sp., the number of nodules were greater than when R. leguminosarum was inoculated alone, but nodule dry weight and pea shoot biomass were similar to plants inoculated with only R. leguminosarum. Although these results need confirmation with non-sterile soil and field studies, these preliminary results indicate that peas will not be affected by wheat root-inhibitory rhizobacteria.     

The gradient between pre-dawn rhizoplane and bulk soil matric potentials, and its relation to the pre-dawn root and leaf water potentials of four species

Uptake of soil water by plants may result in significant gradients between bulk soil and soil in the vicinity of roots. Few experimental studies of water potential gradients in close proximity to roots, and no studies on the relationship of water potential gradients to the root and leaf water potentials, have been conducted. The occurrence and importance of pre-dawn gradients in the soil and their relation to the pre-dawn root and leaf water potentials were investigated with seedlings of four species. Pre-germinated seeds were grown without watering for 7 and lid in a silt loam soil with initial soil matric potentials of -0.02, -0.1 and -0.22 MPa. Significant gradients, independent of the species, were observed only at pre-dawn soil matric potentials lower than -0.25 MPa; the initial soil matric potentials were -0.1 MPa. At an initial bulk soil matric potential of -0.22 MPa, a steep gradient between bulk and rhizoplane soil was observed after 7 d for maize (Zea mays L. cv. Issa) and sunflower (Helianthus annuus L. cv. Nanus), in contrast to barley (Hordeum vulgare L. cv. Athos) and wheat (Triticum aestivum L. cv. Kolibri). Pre-dawn root water potentials were usually about the same as the bulk soil matric potential and were higher than the rhizoplane soil matric potential. Pre-dawn root and leaf water potentials tended to be much higher than rhizoplane soil matric potentials when the latter were lower than -0.5 MPa. It is concluded that plants tend to become equilibrated overnight with the wetter bulk soil or with wetter zones in the bulk soil. Plants can thus circumvent negative effects of localized steep pre-dawn soil matric potential gradients. This may be of considerable importance for water uptake and growth in drying soil.