Soil penetration resistance, a suitable soil property to account for variations in root elongation and branching

Background and Aims

Root system development is affected by soil conditions. The effects of bulk density, water content and penetration resistance on root development processes were investigated in peach trees.

Methods

Peach tree rootstocks were grown in various soil conditions, combining two bulk densities (1.2 and 1.5?g soil.cm^-3) and three water contents (0.14, 0.17 and 0.20?g.g^-1soil). Root parameters (tip diameter, length of apical unbranched zone, branching density and diameters of main and lateral roots) and plant growth (leaves, branches, trunk, root dry mass) were measured. Root growth processes (elongation, branching) were studied using relationships between root parameters.

Results

The proportion of biomass allocated to each plant compartment was similar whatever the soil conditions. Variations in root development were best explained by the variation in penetration resistance, rather than other soil properties. Increased soil penetration resistance reduced the root elongation rate, especially for thick roots. In addition, the branching pattern was affected. In soil with a high penetration resistance, the root system shape differs from a typical herringbone pattern.

Conclusions

These results allow quantification of the root system plasticity, and improve our understanding of the interactions between root development and soil properties.     

Fine root morphological and functional traits in Fagus sylvatica and Fraxinus excelsior saplings as dependent on species, root order and competition

Aims

The mechanisms of belowground competition are not well understood. Addressing literature reports on competition-induced changes in tree fine root morphology, we conducted a growth experiment with tree saplings to investigate competition effects on important root morphological and functional traits in a root order-focused analysis.

Methods

European beech and European ash saplings were grown for 34 months in containers under greenhouse conditions in monoculture (2 conspecific plants), in mixture (1 beech and 1 ash) or as single plants. The root system was fractionated according to root orders and eight morphological and functional properties were determined.

Results

Root order was the most influential factor affecting the fine root traits (except for root diameter and δ¹³C); a significant species identity effect was found for root diameter, tissue density, N concentration and δ¹³C. Ash fine roots were thicker, but had lower tissue densities, contained more N and had systematically higher δ¹³C values than beech roots. The competition treatments had no significant effect on morphological root traits but altered δ¹³C in the 2nd root order.

Conclusion

Neither intra- nor interspecific root competition affected fine root morphology significantly suggesting that competition-induced root modification may not be a universal phenomenon in temperate trees.     

Variation of carbon age of fine roots in boreal forests determined from 14C measurements

Background and aims

The main objectives of this study were to determine how the carbon age of fine root cellulose varies between stands, tree species, root diameter and soil depth. In addition, we also compared the carbon age of fine roots from soil cores of this study with reported values from the roots of the same diameter classes of ingrowth cores on the same sites.

Methods

We used natural abundance of ¹⁴C to estimate root carbon age in four boreal Norway spruce and Scots pine stands in Finland and Estonia.

Results

Age of fine root carbon was older in 1.5–2 mm diameter fine roots than in fine roots with <0.5 mm diameter, and tended to be older in mineral soil than in organic soil. Fine root carbon was older in the less fertile Finnish spruce stands (11–12 years) than in the more fertile Estonian stand (3 and 8 years), implying that roots may live longer in less fertile soil. We further observed that on one of our sites carbon in live fine roots with the 1.5–2 mm diameter was of similar C age (7–12 years) than in the ingrowth core roots despite the reported root age in the ingrowth cores – being not older than 2 years.

Conclusions

From this result, we conclude that new live roots may in some cases use old carbon reserves for their cellulose formation. Future research should be oriented towards improving our understanding of possible internal redistribution and uptake of C in trees.     

Biomechanics of nodal, seminal and lateral roots of barley: effects of diameter, waterlogging and mechanical impedance

Background and aims

Biomechanical properties of cereal root systems largely control both resistance to root lodging and their ability to stabilise soil. Abiotic stresses can greatly modify root system growth and form. In this paper the effect of waterlogging and moderate mechanical impedance on root biomechanics is studied for both lateral roots and the main axes of barley.

Methods

Barley (Hordeum vulgare) plants were subjected to transient water-logging and moderate mechanical impedance in repacked soil columns. Roots were excavated, separated into types (nodal, seminal or lateral) and tested in tension to measure strength and elastic modulus.

Results

Water-logging and mechanical impedance substantially changed root system growth whilst root biomechanical properties were affected by waterlogging. Root strength was generally greater in thin roots and depended on root type. For example, seminal roots 0.4–0.6 mm in diameter were approximately seven times stronger and five times stiffer than lateral roots of the same diameter when mechanically impeded. Root sample populations typically exhibited negative power-law relationships between root strength and diameter for all root types. Mechanical impedance slowed seminal root elongation by approximately 50 % and resulted in a 15 % and 11 % increase in the diameter of in nodal and seminal roots respectively. Power-law relationships between root diameter and root biomechanical properties corresponded to the different root types. Coefficients for between root diameter, strength and elastic modulus improved when separated by root type, with R² values increasing in some roots from 0.05 to 0.71 for root strength and 0.08 to 0.74 for elastic modulus.

Conclusions

Moderate mechanical impedance did not influence the tensile strength of roots, but, waterlogging diminished the relationship between root strength and diameter. Separation of root type improved predictions of root strength and elastic modulus using power-law regressions.     

Drought changes the structure and elemental composition of very fine roots in seedlings of ten woody tree species. Implications for a drier climate

Background and aims

Water availability is often one of the most limiting factors for plants. Climate change predictions for many areas suggest an intensification of water limitation. The ability of a plant to modify its root characteristics can be an important mechanism for preventing drought stress.

Methods

We studied the drought response of seedlings of 10 woody species and compared the biomass allocation, vertical root distribution across different root diameters, and the key traits of very fine roots (root diameter <0.5 mm) under two water regimes (no water limitation and severe drought).

Results

Under drought conditions, the very fine roots had a higher specific root length (SRL, root length: biomass ratio), smaller root diameter and higher root tissue mass density, as well as a lower nitrogen concentration. A higher value of the mean root plasticity index was related to higher drought resistance. A quantitative literature review showed that there was a wide variation in the effect of the drought on SRL, thus there was not a clear effect of drought on SRL.

Conclusions

Certain species have the necessary root traits and plasticity to survive drought. We have identified plasticity in root characteristics as a whole-plant trait which plays a significant role in separating out species into those which are vulnerable and those which are resistant to drought.     

Comparative analysis of Cd and Zn impacts on root distribution and morphology of Lolium perenne and Trifolium repens: implications for phytostabilization

Backgrounds and aims

The phytostabilization potential of plants is a direct function of their root systems. An experimental design was developed to investigate the impact of Cd and Zn on the root distribution and morphology of Lolium perenne and Trifolium repens.

Methods

Seedlings were transplanted into columns filled with washed quartz and irrigated daily with Cd- or Zn-containing nutrient solutions during 1 month. Root biomass, root length density (RLD) and diameter were subsequently quantified as a function of depth. Pot experiments were also performed to quantify metal, lignin and structural polysaccharides concentrations as well as cell viability.

Results

Lolium perenne accumulated Cd and Zn in the roots whereas T. repens was unable to restrict heavy metal translocation. Cadmium and Zn reduced rooting depth and RLD but induced thick shoot-borne roots in L. perenne. Cd-induced root swelling was related to lignification occurring in the exodermis and parenchyma of central cylinder. Hemicelluloses and lignin did not play a key role in root metal retention. Cadmium slightly reduced mean root cell viability whereas Zn increased this parameter in comparison to Cd.

Conclusions

Even though plant species like Lolium perenne and Trifolium repens may appear suitable for a phytostabilization scheme based on their shoot metal tolerance, exposure to toxic heavy metals drastically impairs their root distribution. This could jeopardize the setting up of phytostabilization trials. The metal-induced alterations of root system properties are clearly metal- and species-specific. At sites polluted with multiple metals, it is therefore recommended to first test their impact on the root system of multiple plant species so as to select the most appropriate species for each site.     

Grassland root demography responses to multiple climate change drivers depend on root morphology

Aims

We examine how root system demography and morphology are affected by air warming and multiple, simultaneous climate change drivers.

Methods

Using minirhizotrons, we studied root growth, morphology, median longevity, risk of mortality and standing root pool in the upper soil horizon of a temperate grassland ecosystem for 3 years. Grassland monoliths were subjected to four climate treatments in a replicated additive design: control (C); elevated temperature (T); combined T and summer precipitation reduction (TD); combined TD and elevated atmospheric CO₂ (TDCO₂).

Results

Air warming (C vs T) and the combined climate change treatment (C vs TDCO₂) had a positive effect on root growth rate and standing root pool. However, root responses to climate treatment varied depending on diameter size class. For fine roots (≤ 0.1 mm), new root length and mortality increased under warming but decreased in response to elevated CO₂ (TD vs TDCO₂); for coarse roots (> 0.2 mm), length and mortality increased under both elevated CO₂ and combined climate change drivers.

Conclusions

Our data suggest that the standing roots pool in our grassland system may increase under future climatic conditions. Contrasted behaviour of fine and coarse roots may correspond to differential root activity of these extreme diameter classes in future climate.     

Longitudinal variation in cadmium influx in intact first order lateral roots of sunflower (Helianthus annuus. L)

Aims

Contamination of sunflower (Helianthus annuus L.) by cadmium (Cd) is a concern for food and feed safety as this species accumulates Cd to a greater extent than other crops. We examined the relationships between root architecture and Cd²⁺ uptake by roots.

Methods

We determined and mathematically modelled the longitudinal variation of Cd²⁺ influx in first order roots of sunflower grown in hydroponics by using short-term exposure to ¹⁰⁹Cd-labelled solutions (0.8 to 500 nM). Thereafter, by taking into account the longitudinal variation of the influx, we simulated the uptake of Cd²⁺ for 24 h by cohorts of roots characterised by various architectural characteristics.

Results

Cd²⁺ influx at the root tip was on average 2.9 times that of the basal region close to the taproot. The simulations indicated that the total Cd²⁺ uptake by root cohorts mainly depends on 1/ the root diameter and the number of roots, 2/ the value of the Cd²⁺ influx at the basal region 3/ the stronger influx at the root tip.

Conclusion

Considering a higher Cd²⁺ influx at the root tip may be important to understand the relationship between root architecture and Cd²⁺ uptake by the root system.     

Improved scaling of minirhizotron data using an empirically-derived depth of field and correcting for the underestimation of root diameters

Background and aims

Accurate data on the standing crop, production, and turnover of fine roots is essential to our understanding of major terrestrial ecological processes. Minirhizotrons offer a unique opportunity to study the dynamic processes of root systems, but are susceptible to several measurement biases.

Methods

We use roots extracted from minirhizotron tube surfaces to calculate the depth of field of a minirhizotron image and present a model to correct for the underestimation of root diameters obscured by soil in minirhizotron images.

Results

Non-linear regression analysis resulted in an estimated depth of field of 0.78 mm for minirhizotron images. Unadjusted minirhizotron data underestimated root net primary production and fine root standing crop by 61 % when compared to adjusted data using our depth of field and root diameter corrections. Changes in depth of field accounted for >99 % of standing crop adjustments with root diameter corrections accounting for <1 %.

Conclusions

Our results represent the first effort to empirically derive depth of field for minirhizotron images. This work may explain the commonly reported underestimation of fine roots using minirhizotrons, and stands to improve the ability of researchers to accurately scale minirhizotron data to large soil volumes.     

Coarse woody debris modifies surface soils of degraded temperate eucalypt woodlands

Aims

Reintroductions of coarse woody debris (CWD) to Australia’s temperate eucalypt woodlands have been proposed to address the paucity of CWD in these landscapes. This study aimed to quantify the effects of CWD on surface soils.

Methods

Values of C, N, C:N, P, NO₃ ^?, NH₄ ⁺, pH and electrical conductivity (EC) were measured adjacent to, and at reference distances from CWD. Soils were measured at depths of 0–1 cm, 1–3 cm and 3–5 cm for 12 individual CWD samples of varying decay classes and diameters. A linear mixed model was used to test the effects of the presence of CWD, soil depth and CWD decay class and diameter.

Results

Significantly larger values for C, N, C:N, P, NO₃ ^?, EC, and significantly smaller values for pH were found adjacent to CWD. The greatest impact of CWD was on the upper most surface soil. CWD decay class and diameter had little influence on the measured soil characteristics.

Conclusion

This is the first quantitative determination of the effects of eucalypt CWD on woodland soils in Australia. The effect of added CWD is rapid, occurring after just 2 years. The results suggest that the effects are due to the structural properties of CWD.     

The use of PET/CT scanning technique for 3D visualization and quantification of real-time soil/plant interactions

Aims

Conventional methodology using destructive sampling, which is laborious and has poor spatial and temporal resolution, has limited our understanding of soil-plant interactions. New non-invasive tomographic techniques have the potential to significantly improve our knowledge. In this study we demonstrated the simultaneous use of PET (positron emission tomography) and CT (X-ray computed tomography) to (a) non-destructively image a whole plant growing in sand, and (b) to link the observed morphology with recently assimilated C. The PET scanner was used to detect and visualize the location of the short-lived radioisotope ¹¹C (with a half-life of 20.4?min) taken up by the plant through ¹¹C-labelled CO₂. This provided information on carbon translocation and the metabolism of photo-assimilates in the plant as well as root structure. The CT scanners yielded data on soil and root structure.

Methods

A medical PET/CT scanner was used to scan a fodder radish plant growing in a pot with test soil composed of homogenous sand. We constructed an air-plant-soil controller system (APS) to control the environmental conditions, such as CO₂, temperature and light during the experiment. The plant was allowed to assimilate ¹¹CO₂ for 90?min before PET scanning was initiated. We carried out PET scanning for 60?min. Subsequently, the aerial parts of the plant was cut off and the pot was rescanned using a micro-CT scanner to obtain more detailed information on structure of the root system and the growth medium structure.

Results

The acquired PET and CT images gave images clearly visualizing the architecture and morphology of root and soil. Using a CT scanner, we were able to detect the main taproot located at 0 to 30?mm depth. With the PET scanner, we were able to measure a signal down to 82?mm below the surface of the sand. We found the highest concentration of ¹¹C at the position of the main root. The PET images, at different time intervals, showed the translocation and metabolisation of photo-assimilates from top to root. Using the micro-CT scanner (voxel size of 90?μm), we were able to detect roots down to 100?mm depth. These findings correlated the PET signals measured down to 82?mm depth.

Conclusions

We conclude that the simultaneous use of PET and CT technologies was successfully applied for soil-plant studies. The combined PET/CT technology has potential to provide new fundamental insight into soil-plant interactions and especially into the effect of abiotic stresses in spite of the limitation due to spatial resolution.     

Comment on: “root orientation can affect detection accuracy of ground-penetrating radar”

Introduction

In a recent paper, Tanikawa et al. Plant Soil 373:317–327, (2013) reported a considerable impact of root orientation on the accuracy of root detection and root diameter estimation by ground-penetrating radar (GPR).

Methods

In Tanikawa et al. Plant Soil 373:317–327, (2013), buried root samples in a sand box were scanned from multiple cross angles between root orientation and GPR transecting line under controlled conditions. Changes in radar waveform parameter of roots to different cross angles were investigated.

Results

Tanikawa et al. Plant Soil 373:317–327, (2013) clarified that 1) the variation in amplitude area (a signal strength related waveform parameter) to different cross angles fitted a sinusoidal waveform; and 2) the impact of root orientation on root diameter estimation by GPR could be mathematically corrected by applying a grid transect survey. However, we found that the quantitative relationship established in Tanikawa et al. Plant Soil 373:317–327, (2013) between amplitude area and cross angle was incorrect, and the application of a grid transect survey still underestimated root diameter.

Conclusion

The change in amplitude area to cross angle between transecting line and root orientation fits a sinusoidal waveform but different to that reported in Tanikawa et al. Plant Soil 373:317–327, (2013). The polarization of GPR wave may explain such sinusoidal variation in amplitude area to cross angle. The effect of root orientation on GPR-based root diameter estimation remains to be calibrated.     

IJ_Rhizo: an open-source software to measure scanned images of root samples

Background and aims

This paper provides an overview of the measuring capabilities of IJ_Rhizo, an ImageJ macro that measures scanned images of washed root samples. IJ_Rhizo is open-source, platform-independent and offers a simple graphic user interface (GUI) for a main audience of non-programmer scientists. Being open-source based, it is also fully modifiable to accommodate the specific needs of the more computer-literate users. A comparison of IJ_Rhizo’s performance with that of the widely used commercial package WinRHIZO? is discussed.

Methods

We compared IJ_Rhizo’s performance with that of the commercial package WinRHIZO? using two sets of images, one comprising test-line images, the second consisting of images of root samples collected in the field. IJ_Rhizo and WinRHIZO? estimates were compared by means of correlation and regression analysis.

Results

IJ_Rhizo “Kimura” and WinRHIZO? “Tennant” were the length estimates that were best linearly correlated with each other. Correlation between average root diameter estimates was weaker, due to the sensitivity of this parameter to thresholding and filtering of image background noise.

Conclusions

Overall, IJ_Rhizo offers new opportunities for researchers who cannot afford the cost of commercial software packages to carry out automated measurement of scanned images of root samples, without sacrificing accuracy.     

Root orientation can affect detection accuracy of ground-penetrating radar

Aim

Ground-penetrating radar (GPR) has been applied to detect coarse tree roots. The horizontal angle of a root crossing a scanning line is a factor that affects both root detection and waveform parameter values. The purpose of this study was to quantitatively evaluate the influence of root orientation (x, degree) on two major waveform parameters, amplitude area (A, dB × ns) and time interval between zero crossings (T, ns).

Methods

We scanned four diameter classes of dowels in a sandy bed as simulated roots using a 900 MHz antenna from multiple angles to clarify the relationships between the parameters and x.

Results

Angle x strongly affected reflection images and A values. The variation in A(x) fitted a sinusoidal waveform, whereas T was independent of x. The value of A scanning at 90° was estimated by A values of arbitrary x in two orthogonal transects. The sum of T in all reflected waveforms showed a significant linear correlation with dowel diameter.

Conclusions

We clarified that root orientation dramatically affected root detection and A values. The sum of T of all reflected waveforms was a suitable parameter for estimating root diameter. Applying grid transects can overcome the effects of root orientation.     

Spatial distribution and phenotypic variation in root cortical aerenchyma of maize (Zea mays L.)

Background and aims

Previous research has suggested that root cortical aerenchyma (RCA) can enhance soil exploration and crop performance by decreasing root respiration. This trait is a potential breeding target for adaptation to abiotic stresses such as drought and low nutrient availability. However, little is known of phenotypic variation in aerenchyma or its distribution among root classes.

Methods

The spatial distribution of RCA was evaluated in freehand sections from 13 sites in the root systems of 11 recombinant inbred and commercial lines of maize (Zea mays). RCA variation was evaluated in 583 recombinant inbred lines of maize at one sampling position.

Results

RCA varied significantly among root classes and axial positions. Genotypic differences were observed for the amount of RCA at corresponding sampling locations and for the mean amount of RCA across all sampling locations, but genotypes did not differ in the proportional distribution of RCA within the whole root system. The amount of RCA in a cross-section was independent of several other anatomical traits.

Conclusions

There is substantial genetic variation for RCA, and this variation is independent of other anatomical traits. RCA can be phenotyped in greenhouse-grown plants by sampling the middle parts of second- or third-whorl crown roots.     

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

Background and aims

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

Methods

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

Results

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

Conclusions

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

Longitudinal variation in cadmium influx in sunflower (Helianthus annuus L.) roots as depending on the growth substrate,root age and root order

Aims

In a previous work, we observed a longitudinal decrease in Cd²⁺ influx starting from the root tip in first order lateral roots of sunflower (Helianthus annuus L.) grown in hydroponics. This variable influx was expected to impact the total Cd²⁺ uptake depending on the root system architecture and on how steep was the decrease of the influx. Here, we examined the influence of the culture substrate, of the age and order of lateral roots on the longitudinal variation of Cd²⁺ influx.

Methods

By using short-term exposures to ¹⁰⁹Cd-labelled solution (5 to 200 nM), we compared the longitudinal variations in Cd²⁺ roots influx depending on the growth substrate (hydroponics or sand), on the root age and order.

Results

In second order laterals, Cd²⁺ influx decreased from the apex to the root base, as for first order laterals. For sand cultures compared to hydroponics, the mean Cd²⁺ influx was lower and decreased more steeply with the distance from the apex. The influx also decreased with increasing root age and order, markedly in hydroponics but less for sand cultures.

Conclusion

Results suggested that for a given root surface area, the Cd²⁺ uptake by a root system should increase with increasing number of root tips and decreasing individual root length.     

Observations of below-ground characteristics of young redwood trees (Sequoia sempervirens) from two sites in New Zealand – implications for erosion control

Aims

Radiata pine (Pinus radiata D. Don) plantations are widely used to control erosion in New Zealand. However, other species with similar growth but longer rotation lengths and ability to coppice may offer future alternatives to radiata pine. Comparing performance of alternative species to radiata thus becomes important if policy is to be developed to promote them.

Methods

The below-ground characteristics (roots) of young redwood (Sequoia sempervirens (D. Don) Endl.) trees from two established plantations in New Zealand were examined and compared with those of radiata pine, and selected poplar and New Zealand native species.

Results

Roots with diameters less than 10 mm comprised over 99 % of total root length in 3-yr-old trees and 98 % of total root length in 4-yr-old trees. For roots greater than 2 mm in diameter, total root length of young redwood trees was greater than that of young radiata pine, poplar and the best performing New Zealand native plant. Total root length at a given root collar diameter for young (1–4 year old) redwood trees was significantly greater than for radiata pine trees. Roots of redwoods were finer and more numerous than for radiata but the below-ground biomass for a given root collar diameter showed no statistical difference between the two species.

Conclusions

Redwood, because of its comparable growth rate and the production of many fine lateral roots, has the potential to become a keystone erosion-control species in New Zealand, especially on steep lands where there is an increased risk of post-harvest landsliding associated with moderate to severe rainstorm events.     

Root lodging tolerance in Helianthus annuus (L.): associations with morphological and mechanical attributes of roots

Background and aims

The objectives of this study were to quantify the morphological and mechanical properties of the root-plate within two sunflower hybrids of contrasting susceptibility to root lodging; and to evaluate the effects of crop population density on these properties at two different development stages.

Methods

Two hybrids (CF29: tolerant, Zenit: sensitive) were grown at three densities: 5.6, 10 and 16 plants m^?2. At R2 (early reproductive) and R6 (end anthesis) development stages, plants were artificially lodged and stem biomass, total root biomass in the whole root-plate and in the 0–5 and >5 cm layers of the plate, root number (three diametrical categories: 0–1, 1.1–2, >2 mm), total root length, and root axial breakage force were assessed.

Results

CF29 root mass was twice that of Zenit with differences mainly in the top 5 cm of soil. This higher root-plate biomass of CF29 was associated with a greater root number and root length compared to Zenit within all root diameter categories. Roots of CF29 exhibited higher axial tension failure thresholds than those of Zenit, and these thresholds increased more sharply with root diameter in CF29 than in Zenit.

Conclusions

The better anchorage and tolerance to lodging of CF29 with respect to Zenit arose from additive actions of traits at both whole root-plate and individual root levels. These included total root-plate root length, root number, root biomass and root axial breakage force.     

Neutron imaging reveals internal plant water dynamics

Background and aims

Knowledge of plant water fluxes is critical for assessing mechanistic processes linked to biogeochemical cycles, yet resolving root water transport dynamics has been a particularly daunting task. Our objectives were to demonstrate the ability to non-invasively monitor individual root functionality and water fluxes within Zea mays L. (maize) and Panicum virgatum L. (switchgrass) seedlings using neutron imaging.

Methods

Seedlings were propagated for 1–3 weeks in aluminum chambers containing sand. Pulses of water or deuterium oxide were then tracked through the root systems by collecting consecutive radiographs during exposure to a cold-neutron source. Water flux was manipulated by cycling on a growth lamp to alter foliar demand for water.

Results

Neutron radiography readily illuminated root structure, root growth, and relative plant and soil water content. After irrigation there was rapid root water uptake from the newly wetted soil, followed by hydraulic redistribution of water through the root system to roots terminating in dry soil. Water flux within individual roots responded differentially to foliar illumination based on supply and demand of water within the root system.

Conclusions

Sub-millimeter scale image resolution revealed timing and magnitudes of root water uptake, redistribution within the roots, and root-shoot hydraulic linkages—relationships not well characterized by other techniques.