Formation and Stabilization of Rhizosheaths of Zea mays L. (Effect of Soil Water Content)

Field observations have shown that rhizosheaths of grasses formed under dry conditions are larger, more coherent, and more strongly bound to the roots than those formed in wet soils. We have quantified these effects in a model system in which corn (Zea mays L.) primary roots were grown through a 30-cm-deep prepared soil profile that consisted of a central, horizontal, "dry" (9% water content) or "wet" (20% water content) layer (4 cm thick) sandwiched between damp soil (15-17% water content). Rhizosheaths formed in dry layers were 5 times the volume of the subtending root. In wet layers, rhizosheaths were only 1.5 times the root volume. Fractions of the rhizosheath soil were removed from individual roots by three successive treatments; sonication, hot water, and abrasion. Sonication removed 50 and 90% of the soil from rhizosheaths formed in dry and wet soils, respectively. After the heat treatment, 35% of the soil still adhered to those root portions where rhizosheaths had developed in dry soil, compared with 2% where sheaths had formed in wet soil. Root hairs were 4.5 times more abundant and were more distorted on portions of roots from dry layers than from wet layers. Drier soil enhanced adhesiveness of rhizosheath mucilages and stimulated the formation of root hairs; both effects stabilize the rhizosheath. Extensive and stable rhizosheaths may function in nutrient acquisition in dry soils.     

The distribution and abundance of wheat roots in a dense,structured subsoil – implications for water uptake

We analysed the abundance, spatial distribution and soil contact of wheat roots in dense, structured subsoil to determine whether incomplete extraction of subsoil water was due to root system limitations. Intact soil cores were collected to 1.6 m below wheat crops at maturity on a red Kandosol in southern Australia. Wheat roots, remnant roots, soil pores and root–soil contact were quantified at fresh breaks in the soil cores. In surface soil layers (<0.6 m) 30–40% of roots were clumped within pores and cracks in the soil, increasing to 85–100% in the subsoil (>0.6 m), where 44% of roots were in pores with at least three other roots. Most pores contained no roots, with occupancy declining from 20% in surface layers to 5% in subsoil. Wheat roots clumped into pores contacted the surrounding soil via numerous root hairs, whereas roots in cracks were appressed to the soil surface and had very few root hairs. Calculations assuming good root–soil contact indicated that root density was sufficient to extract available subsoil water, suggesting that uptake is constrained at the root–soil interface. To increase extraction of subsoil water, genetic targets could include increasing root–soil contact with denser root hairs, and increasing root proliferation to utilize existing soil pores.     

Drought stress and plant ecotype drive microbiome recruitment in switchgrass rhizosheath

The rhizosheath, a layer of soil grains that adheres firmly to roots, is beneficial for plant growth and adaptation to drought environments. Switchgrass is a perennial C4 grass which can form contact rhizosheath under drought conditions. In this study, we characterized the microbiomes of four different rhizocompartments of two switchgrass ecotypes (Alamo and Kanlow) grown under drought or well-watered conditions via 16S ribosomal RNA amplicon sequencing. These four rhizocompartments, the bulk soil, rhizosheath soil, rhizoplane, and root endosphere, harbored both distinct and overlapping microbial communities. The root compartments (rhizoplane and root endosphere) displayed low-complexity communities dominated by Proteobacteria and Firmicutes. Compared to bulk soil, Cyanobacteria and Bacteroidetes were selectively enriched, while Proteobacteria and Firmicutes were selectively depleted, in rhizosheath soil. Taxa from Proteobacteria or Firmicutes were specifically selected in Alamo or Kanlow rhizosheath soil. Following drought stress, Citrobacter and Acinetobacter were further enriched in rhizosheath soil, suggesting that rhizosheath microbiome assembly is driven by drought stress. Additionally, the ecotype-specific recruitment of rhizosheath microbiome reveals their differences in drought stress responses. Collectively, these results shed light on rhizosheath microbiome recruitment in switchgrass and lay the foundation for the improvement of drought tolerance in switchgrass by regulating the rhizosheath microbiome.     

Plasticity of rhizosphere hydraulic properties as a key for efficient utilization of scarce resources

Background

It is known that the soil near roots, the so-called rhizosphere, has physical and chemical properties different from those of the bulk soil. Rhizosphere properties are the result of several processes: root and soil shrinking/swelling during drying/wetting cycles, soil compaction by root growth, mucilage exuded by root caps, interaction of mucilage with soil particles, mucilage shrinking/swelling and mucilage biodegradation. These processes may lead to variable rhizosphere properties, i.e. the presence of air-filled gaps between soil and roots; water repellence in the rhizosphere caused by drying of mucilage around the soil particles; or water accumulation in the rhizosphere due to the high water-holding capacity of mucilage. The resulting properties are not constant in time but they change as a function of soil condition, root growth rate and mucilage age.

Scope

We consider such a variability as an expression of rhizosphere plasticity, which may be a strategy for plants to control which part of the root system will have a facilitated access to water and which roots will be disconnected from the soil, for instance by air-filled gaps or by rhizosphere hydrophobicity. To describe such a dualism, we suggest classifying rhizosphere into two categories: class A refers to a rhizosphere covered with hydrated mucilage that optimally connects roots to soil and facilitates water uptake from dry soils. Class B refers to the case of air-filled gaps and/or hydrophobic rhizosphere, which isolate roots from the soil and may limit water uptake from the soil as well water loss to the soil. The main function of roots covered by class B will be long-distance transport of water.

Outlook

This concept has implications for soil and plant water relations at the plant scale. Root water uptake in dry conditions is expected to shift to regions covered with rhizosphere class A. On the other hand, hydraulic lift may be limited in regions covered with rhizosphere class B. New experimental methods need to be developed and applied to different plant species and soil types, in order to understand whether such dualism in rhizosphere properties is an important mechanism for efficient utilization of scarce resources and drought tolerance.     

Hydraulic lift in drought-tolerant and -susceptible maize hybrids

Hydraulic lift was investigated in a greenhouse study involving two drought-tolerant maize (Zea mays L.) hybrids (TAES176 and P3223) and a drought-susceptible hybrid (P3225) during the flowering stage. Root systems were grown in two soil compartments – a drier upper soil and a wetter deep soil. The plants were shaded for 3 h during the daytime. Soil volumetric water content (Ø_v) in the upper pots was measured with time domain reflectometry (TDR) before and after shading. An increase in Ø_v in the upper pot was detected with TDR in the drought-tolerant hybrids following 3 h of shading, but not in the drought-susceptible hybrid. Furthermore, water exuded from roots in the top soil layers was greater in the more drought-tolerant TAES176 than in P3223 (489 vs. 288 g per pot in 3 h, P<0.005). The sizable amount of water from hydraulic lift allowed TAES176 to reach a peak transpiration rate 27–42% higher than the drought-susceptible hybrid P3225 on the days when the evaporative demand was high. To our knowledge, this is the first experiment that reveals a significant surge of transpiration due to hydraulic lift following midday shading. Hydraulic lift also prevented soil moisture depletion in the upper pots with TAES176, but not with P3223 or P3225. Root characteristics may be responsible for differences in hydraulic lift of the three maize hybrids. There were 2.3–3.3-fold more primary roots in the deep moist soil in P3223 and TAES176 than in P3225 that may enable these hybrids to absorb and transport water at faster rates. Therefore, more water can be exuded into the upper drier soil when transpiration is suppressed by shading. Larger primary roots (20–28% larger diameter) and a higher root volume in the upper soil in TAES176 and P3223 than in P3225 may contribute to higher root hydraulic conductance and greater water efflux from the roots. The negligible hydraulic lift in P3225 may also relate to higher night-time transpiration of the hybrid. This report has documented, for the first time, the existence of genetic variations in hydraulic lift among maize hybrids and links between hydraulic lift and drought tolerance within maize plants. It appears that one of drought tolerance mechanisms in maize may lie in the extent of hydraulic lift.     

Water uptake and structural plasticity along roots of a desert succulent during prolonged drought

Desert succulents resume substantial water uptake within 1–2 d of the cessation of drought, but the changes in root structure and hydraulic conductivity underlying such recovery are largely unknown. In the monocotyledonous leaf succulent Agave deserti Engelm. substantial root mortality occurred only for lateral roots near the soil surface; nearly all main roots were alive at 180 d of drought. New main roots were initiated and grew up to 320 mm at soil water potentials lower than – 5·0 MPa, utilizing water from the shoot. The hydraulic conductivity of distal root regions decreased 62% by 45 d of drought and 70% thereafter. After 7 d of rewetting, root hydraulic conductivity was restored following 45 d of drought but not after 90 and 180 d. The production of new lateral roots and the renewed apical elongation of main roots occurred 7–11 d after rewetting following 180 d of drought. Hydraulic conductivity was higher in the distal region than at midroot and often increased again near the root base, where many endodermal cells lacked suberin lamellae. Suberization and xylem maturation were influenced by the availability of moisture, suggesting that developmental plasticity along a root allows A. deserti to capitalize on intermittent or heterogeneous supplies of water.     

Influence of pastoral fallow on plant root growth and soil physical and chemical characteristics in a hill pasture

Pastoral fallowing over a growing season (October–May) has a profound effect on standing biomass and sward structure, and should have an impact on below ground plant growth and soil biological activities. Two field studies were conducted to compare the effects of pastoral fallow with rotational grazing on root growth and soil physical and chemical properties. Root growth and distribution was altered by pastoral fallowing and there was significantly (P < 0.01) less root biomass at 0–50 mm depth of soil in the fallowed sward than the grazed sward. Compared with the grazed treatment, pastoral fallow increased soil air permeability at 500 mm tension by 38%, saturated hydraulic conductivity by 26%, unsaturated hydraulic conductivity at 20 mm tension by 67% and soil moisture by 10–15%, and reduced soil bulk density by 11%. Fallowing had little effect on soil nutrients both at the end of fallowing, except for small reductions in K and Mineral N levels at 0–75 mm soil depth, and two to three years after fallowing.     

Microtensiometer technique for in situ measurement of soil matric potential and root water extraction from a sandy soil

The suitability of microtensiometers to measure the spatial variation of soil matric potential and its diurnal change was tested in a pot experiment with pearl millet (Pennisetum americanum [L.] Leeke) in a sandy soil as the soil dried out.The temporal and spatial resolution of this technique allowed precise measurement of soil matric potential and thus estimation of soil water extraction from different compartments as well as from the whole rooting zone. The technique also allowed the measurement of rehydration of plants at night and root water uptake rate per unit soil volume or per unit root length. The precision of determination of root water uptake depended greatly on the accuracy of the estimate of hydraulic conductivity, which was derived from a bare soil and might be different for a cropped soil owing to aggregation induced by the root system. A linear relationship between root length and water uptake was found (r²=0.82), irrespective of variation in soil water content between compartments and despite the variation in root age, xylem differentiation and suberin formation expected to exist between different compartments of the rooting zone. As the experiment was carried out in a range of soil matric potentials between –4 and –30 kPa, drought stress did not occur. Further information at lower soil matric potentials are required, to address questions such as the importance of soil resistance for water uptake, or which portion of the root system has to be stressed to induce hormonal signals to the shoot. The microtensiometer technique can be applied to soil matric potentials up to –80 kPa.     

Native root xylem embolism and stomatal closure in stands of Douglas-fir and ponderosa pine: mitigation by hydraulic redistribution

Hydraulic redistribution (HR), the passive movement of water via roots from moist to drier portions of the soil, occurs in many ecosystems, influencing both plant and ecosystem-water use. We examined the effects of HR on root hydraulic functioning during drought in young and old-growth Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] and ponderosa pine (Pinus ponderosa Dougl. Ex Laws) trees growing in four sites. During the 2002 growing season, in situ xylem embolism, water deficit and xylem vulnerability to embolism were measured on medium roots (2–4-mm diameter) collected at 20–30 cm depth. Soil water content and water potentials were monitored concurrently to determine the extent of HR. Additionally, the water potential and stomatal conductance (g_s) of upper canopy leaves were measured throughout the growing season. In the site with young Douglas-fir trees, root embolism increased from 20 to 55 percent loss of conductivity (PLC) as the dry season progressed. In young ponderosa pine, root embolism increased from 45 to 75 PLC. In contrast, roots of old-growth Douglas-fir and ponderosa pine trees never experienced more than 30 and 40 PLC, respectively. HR kept soil water potential at 20–30 cm depth above –0.5 MPa in the old-growth Douglas-fir site and –1.8 MPa in the old-growth ponderosa pine site, which significantly reduced loss of shallow root function. In the young ponderosa pine stand, where little HR occurred, the water potential in the upper soil layers fell to about –2.8 MPa, which severely impaired root functioning and limited recovery when the fall rains returned. In both species, daily maximum g_s decreased linearly with increasing root PLC, suggesting that root xylem embolism acted in concert with stomata to limit water loss, thereby maintaining minimum leaf water potential above critical values. HR appears to be an important mechanism for maintaining shallow root function during drought and preventing total stomatal closure.     

The dependence of root system properties on root system biomass of 10 North American grassland species

Dependence of the properties of root systems on the size of the root system may alter conclusions about differences in plant growth in different environments and among species. To determine whether important root system properties changed as root systems aged and accumulated biomass, we measured three important properties of fine roots (tissue density, diameter, and C:N) and three biomass ratios (root:shoot, fine:coarse, and shallow:deep) of monocultures of 10 North American grassland species five times during their second and third years of growth. With increasing belowground biomass, root tissue density increased and diameter decreased. This may reflect cortical loss associated with the aging of roots. For non-legumes, fine root C:N decreased with increasing root biomass, associated with decreases in soil solution NO^{3 –} concentrations. No changes in fine root C:N were detected with increasing belowground biomass for the two legumes we studied. Among all 10 species, there were generally no changes in the relative amounts of biomass in coarse and fine roots, root:shoot, or the depth placement of fine roots in the soil profile as belowground biomass increased. Though further research is needed to separate the influence of root system size, age of the roots, and changes in nutrient availability, these factors will need to be considered when comparing root functional traits among species and treatments.     

Water uptake and storage by rhizosheaths of Oryzopsis hymenoides: a numerical simulation

Rhizosheaths (sheaths of sand grains that form around the roots of some grasses) are common in perennial grasses that colonise sandy substrates. It has been hypothesised that rhizosheaths increase water availability by increasing the efficiency of water absorption. Others have suggested that rhizosheaths act as storage reservoirs for water. In either case rhizosheaths undoubtedly play an important role in the water relations of these grasses.
In an attempt to evaluate the main function of rhizosheaths, we developed a finite element cylindrical water flow model which enabled us to simulate water uptake by Oryzopsis hymenoides (Roem, and Shult.) Ricker. This model allowed us to estimate total water uptake by root systems with and without rhizosheaths and to compare these values to the extra water stored within the rhizosheath. The results of this study suggest that the presence of rhizosheaths is more important in reducing the total resistance to water flow within the rhizosphere than in enhancing water storage.     

Nitrogen and phosphorus cycling in relation to stand age of Ecucalyptus regnans F. Muell

Water movement between a root and the soil depends on the hydraulic conductances of the soil, the root, and the intervening root-soil air gap (L_gap) created as roots shrink during soil drying. To measure L_gap, segments of young cylindrical roots of Agave deserti about 3 mm in diameter were placed concentrically or eccentrically within tubes of moistened filter paper at a known water potential. As the width of the air gap between the filter paper and a concentrically located root was made smaller, measured L_gap increased less than did predicted L_gap based on isothermal conditions. For gaps of the size expected in the soil during water loss from roots (e.g., 10% of the root radius), the underprediction was about 70% and was primarily caused by a lowering of the root surface temperature accompanying water evaporation. As a root segment was eccentrically moved toward the filter paper, the measured L_gap increased. For the most eccentric case of touching the filter paper, the measured L_gap was 2.4-fold greater than for the concentric case, compared with an infinite L_gap predicted if the water potential were constant around the root surface. When a root touched soil with a water potential of -1.0MPa, L_gap estimated using a graphical method increased about 2.3-fold and the overall conductance of the root-soil system increased by 31% compared with the concentric case. For markedly eccentric locations of roots in air gaps, L_gap, which can be the principal conductance initially limiting water loss from roots to a drying soil, can be about 60% of the value predicted for the concentric isothermal case.     

CHANGES IN HYDRAULIC CONDUCTIVITY AND ANATOMY CAUSED BY DRYING AND REWETTING ROOTS OF AGAVE DESERTI (AGAVACEAE)

Concurrent determinations of changes in hydraulic conductivity and tissue anatomy were made for roots of Agave deserti excised during drying and following rewetting in soil. At 30 d of drought, hydraulic conductivity had declined less than twofold for older nodal roots, tenfold for young nodal roots, and more than 20-fold for lateral roots (“rain roots” occurring as branches on the nodal roots). These decreases were consistent with increases in cortical lacunae caused by cell shrinkage and collapse. Similarly, reduction of lacunae in response to rewetting after 7 d of drought corresponded to levels of recovery in hydraulic conductivity, with young nodal roots showing full recovery, lateral roots returning to only 21 % of initial conductivity, and older nodal roots changing only slightly. Increases in suberization in the exodermis, endodermis, and cortex adjacent to the endodermis in response to drying coincided with decreases in hydraulic conductivity. Measurements of axial hydraulic conductance per unit length before and after pressurization indicated that embolism caused reductions in axial conductance of 98% for lateral roots, 35% for young nodal roots, and 20% for older nodal roots at 7 d of drought. Embolism, cortical lacunae, and increasing suberization caused hydraulic conductivity to decline during drought in the three root types, thereby helping limit water loss to dry soil; the recovery in hydraulic conductivity for young nodal roots after rewetting would allow them to take up water readily once soil moisture is replenished.     

Changes in root hydraulic conductivity for two tropical epiphytic cacti as soil moisture varies

The tropical epiphytic cacti Epiphyllum phyllanthus and Rhipsalis baccifera experience extreme variations in soil moisture due to limited soil volumes and episodic rainfalls. To examine possible root rectification, whereby water uptake from a wet soil occurs readily but water loss to a dry soil is minimal, responses of root hydraulic conductivity (L_p) to soil drying and rewetting were investigated along with the underlying anatomical changes. After 30 d of soil drying, L_p decreased 50%–70% for roots of both species, primarily because increased suberization of the periderm reduced radial conductivity. Sheaths composed of soil particles, root hairs, and mucilage covered young roots and helped reduce root desiccation. Axial (xylem) conductance increased during drying due to vessel differentiation and maturation, and drought-induced embolism was relatively low. Within 4 d of rewetting, L_p for roots of both species attained predrought values; radial conductivity increased for young roots due to the growth of new branch roots initiated during drying and for older roots due to the development of radial breaks in the periderm. The decreases in L_p during drought reduced plant water loss to a dry soil, and yet maximal water uptake and transpiration occurred within a few days of rewetting, helping these epiphytes to take advantage of episodic rainfalls in a moist tropical forest.     

Rhizosheath formation and involvement in foxtail millet(Setaria italica) root growth under drought stress

The rhizosheath, a layer of soil particles that adheres firmly to the root surface by a combination of root hairs and mucilage, may improve tolerance to drought stress. Setaria italica(L.) P. Beauv.(foxtail millet), a member of the Poaceae family, is an important food and fodder crop in arid regions and forms a larger rhizosheath under drought conditions. Rhizosheath formation under drought conditions has been studied, but the regulation of root hair growth and rhizosheath size in response to soil moisture remains unclear. To address this question, in this study we monitored root hair growth and rhizosheath development in response to a gradual decline in soil moisture. Here, we determined that a soil moisture level of 10%–14%(w/w)stimulated greater rhizosheath production compared to other soil moisture levels. Root hair density and length also increased at this soil moisture level, which was validated by measurement of the expression of root hair-related genes.These findings contribute to our understanding of rhizosheath formation in response to soil water stress.     

Shotgun proteomic analysis of long-distance drought signaling in rice roots

Rice (Oryza sativa L. cv. IR64) was grown in split-root systems to analyze long-distance drought signaling within root systems. This in turn underpins how root systems in heterogeneous soils adapt to drought. The approach was to compare four root tissues: (1) fully watered; (2) fully droughted and split-root systems where (3) one-half was watered and (4) the other half was droughted. This was specifically aimed at identifying how droughted root tissues altered the proteome of adjacent wet roots by hormone signals and how wet roots reciprocally affected dry roots hydraulically. Quantitative label-free shotgun proteomic analysis of four different root tissues resulted in identification of 1487 nonredundant proteins, with nearly 900 proteins present in triplicate in each treatment. Drought caused surprising changes in expression, most notably in partially droughted roots where 38% of proteins were altered in level compared to adjacent watered roots. Specific functional groups changed consistently in drought. Pathogenesis-related proteins were generally up-regulated in response to drought and heat-shock proteins were totally absent in roots of fully watered plants. Proteins involved in transport and oxidation-reduction reactions were also highly dependent upon drought signals, with the former largely absent in roots receiving a drought signal while oxidation-reduction proteins were strongly present during drought. Finally, two functionally contrasting protein families were compared to validate our approach, showing that nine tubulins were strongly reduced in droughted roots while six chitinases were up-regulated, even when the signal arrived remotely from adjacent droughted roots.     

Root growth,aerenchyma development,and oxygen transport in rice genotypes subjected to drought and waterlogging

Soils under field conditions may experience fluctuating soil water regimes ranging from drought to waterlogging. The inability of roots to acclimate to such changes in soil water regimes may result in reduced growth and function thereby, dry matter production. This study compared the root and shoot growth, root aerenchyma development, and associated root oxygen transport of aerobic and irrigated lowland rice genotypes grown under well-watered (control), waterlogged, and droughted soil conditions for 30 days. The aerobic genotypes were as tolerant as the irrigated lowland genotypes under waterlogging because of their comparable abilities to enhance aerenchyma that effectively facilitated O₂ diffusion to the roots for maintaining root growth and dry matter production. Under drought, aerobic genotypes were more tolerant than the irrigated lowland genotypes due to their higher ability to maintain nodal root production, elongation, and branching, thus, less reduction in dry matter production. Aerenchyma was also formed in droughted roots regardless of genotypes, but was resistant to internal O₂ transport under O₂ deficiency. The ability of roots to resist temporal variations in drought and waterlogging stresses might have strong implications for the adaptation of rice growing in environments with fluctuating soil water regimes.     

Fungal ectomycorrhizal community and drought affect root hydraulic properties and soil adherence to roots ofPinus pinaster seedlings

Pinus pinaster seedlings were grown in a sandy dune soil either inoculated withHebeloma cylindrosporum or let to natural colonisation. Six months later, half of the seedlings of both treatments were subjected to a 3-week moderate drought. Root colonisation analysis showed that root tips were colonised to almost 100% independent of the inoculation. DNA determination of the ectomycorrhizal morphotypes showed that inoculated seedlings were extensively mycorrhized byH. cylindrosporum (more than 75%) whereas non-inoculated seedlings were mycorrhized by the exotic speciesThelephora terrestris (50%) andLaccaria bicolor (30%) and to a lesser extent byH. cylindrosporum (20%). Drought did not affect these frequencies. Total plant biomass was not affected by the mycorrhizal status or by drought but the root/shoot biomass ratio as well as the root/leaf surface area ratio were much lower in seedlings extensively colonised byH. cylindrosporum. Root hydraulic conductivity was higher in plants mainly mycorrhized byH. cylindrosporum, showing that this fungus improved the water uptake capacity of the root system as compared toT. terrestris and/orL. bicolor. This positive effect was also found under drought but to a lesser extent.H. cylindrosporum also increased the amount of root-adhering soil as compared to the other fungal symbionts, illustrating the performance of this association in aggregating sandy soil particles and developing the rhizosheath. The origin of the reduced root hydraulic resistance byH. cylindrosporum mycorrhization is discussed for the whole path including soil, soil-root interface and root cortex.     

Soil textures rather than root hairs dominate water uptake and soil–plant hydraulics under drought

Although the role of root hairs (RHs) in nutrient uptake is well documented, their role in water uptake and drought tolerance remains controversial. Maize (Zea mays) wild-type and its hair-defective mutant (Mut; roothairless 3) were grown in two contrasting soil textures (sand and loam). We used a root pressure chamber to measure the relation between transpiration rate (E) and leaf xylem water potential (ψ_{leaf_x}) during soil drying. Our hypotheses were: (1) RHs extend root–soil contact and reduce the ψ_{leaf_x} decline at high E in dry soils; (2) the impact of RHs is more pronounced in sand; and (3) Muts partly compensate for lacking RHs by producing longer and/or thicker roots. The ψ_{leaf_x}(E) relation was linear in wet conditions and became nonlinear as the soils dried. This nonlinearity occurred more abruptly and at less negative matric potentials in sand (ca. −10 kPa) than in loam (ca. −100 kPa). At more negative soil matric potentials, soil hydraulic conductance became smaller than root hydraulic conductance in both soils. Both genotypes exhibited 1.7 times longer roots in loam, but 1.6 times thicker roots in sand. No differences were observed in the ψ_{leaf_x}(E) relation and active root length between the two genotypes. In maize, RHs had a minor contribution to soil–plant hydraulics in both soils and their putative role in water uptake was smaller than that reported for barley (Hordeum vulgare). These results suggest that the role of RHs cannot be easily generalized across species and soil textures affect the response of root hydraulics to soil drying.

Root hairs of maize do not show evident contribution to root growth, water uptake, and soil–plant hydraulics, whereas soil textures affect the response of root hydraulics to soil drying.     

The role of soil hydraulic conductivity on the spatial and temporal variation of root water uptake in drip-irrigated corn

A multiplexed TDR system and a heat-pulse system for stem sap flow measurements were used to determine the spatial and temporal pattern of root water uptake in field-grown corn. The TDR probes, 0.15 and 0.30 m in length, were buried vertically in the soil profile to a depth of 0.95 m below the soil surface and heat-pulse sensors were installed on the plant base. Nocturnal readings from TDR probes were used successfully to differentiate the two components of moisture change: root uptake and net drainage. The instantaneous rate of water extraction by the plant measured by the heat-pulse system agreed well with the integrated rate of root water uptake measured frequently (at half-hour or hourly intervals) by the TDR probes. This agreement enabled further exploration into the cause of the evolution of the spatial and temporal patterns of root water uptake during a drying cycle. The results indicated that right after irrigation in the well-watered soil profile, it is the spatial distribution of the roots that mainly determines the typical pattern of root extraction, in addition to the fact that the roots near the plant base are more effective than those farther away. The higher density and effectiveness of the roots near the plant base dry the soil rapidly so that soil hydraulic conductivity soon becomes a limiting factor for water uptake. Further analysis revealed that a decrease in root uptake occurs near the plant base under a given atmospheric demand when the relative bulk soil hydraulic conductivity decreases to 0.002K _r. This suggests that low conductivity (high resistance) in the soil near the plant base is the initial cause for downward and lateral shifting of the root uptake pattern. Note that this critical value of hydraulic conductivity is not universal since it depends on the soil type and atmospheric water demand during the period under observation. Therefore, prior to the application of moisture content or suction head as measures of water availability or to control irrigation scheduling, it is suggested that these parameters be calibrated by the soil K() or K() curves, respectively, for the expected atmospheric water demand for the specific crop and growing period.