Three-dimensional visualization and quantification of water content in the rhizosphere

? Despite the importance of rhizosphere properties for water flow from soil to roots, there is limited quantitative information on the distribution of water in the rhizosphere of plants. ? Here, we used neutron tomography to quantify and visualize the water content in the rhizosphere of the plant species chickpea (Cicer arietinum), white lupin (Lupinus albus), and maize (Zea mays) 12 d after planting. ? We clearly observed increasing soil water contents (θ) towards the root surface for all three plant species, as opposed to the usual assumption of decreasing water content. This was true for tap roots and lateral roots of both upper and lower parts of the root system. Furthermore, water gradients around the lower part of the roots were smaller and extended further into bulk soil compared with the upper part, where the gradients in water content were steeper. ? Incorporating the hydraulic conductivity and water retention parameters of the rhizosphere into our model, we could simulate the gradual changes of θ towards the root surface, in agreement with the observations. The modelling result suggests that roots in their rhizosphere may modify the hydraulic properties of soil in a way that improves uptake under dry conditions.     

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.     

Drying of mucilage causes water repellency in the rhizosphere of maize: measurements and modelling

Background and Aims

Although maize roots have been extensively studied, there is limited information on the effect of root exudates on the hydraulic properties of maize rhizosphere. Recent experiments suggested that the mucilaginous fraction of root exudates may cause water repellency of the rhizosphere. Our objectives were: 1) to investigate whether maize rhizosphere turns hydrophobic after drying and subsequent rewetting; 2) to test whether maize mucilage is hydrophobic; and 3) to find a quantitative relation between rhizosphere rewetting, particle size, soil matric potential and mucilage concentration.

Methods

Maize plants were grown in aluminum containers filled with a sandy soil. When the plants were 3-weeks-old, the soil was let dry and then it was irrigated. The soil water content during irrigation was imaged using neutron radiography. In a parallel experiment, ten maize plants were grown in sandy soil for 5 weeks. Mucilage was collected from young brace roots growing above the soil. Mucilage was placed on glass slides and let dry. The contact angle was measured with the sessile drop method for varying mucilage concentration. Additionally, capillary rise experiments were performed in soils of varying particle size mixed with maize mucilage. We then used a pore-network model in which mucilage was randomly distributed in a cubic lattice. The general idea was that rewetting of a pore is impeded when the concentration of mucilage on the pore surface (g cm^?2) is higher than a given threshold value. The threshold value depended on soil matric potential, pore radius and contract angle. Then, we randomly distributed mucilage in the pore network and we calculated the percolation of water across a cubic lattice for varying soil particle size, mucilage concentration and matric potential.

Results

Our results showed that: 1) the rhizosphere of maize stayed temporarily dry after irrigation; 2) mucilage became water repellent after drying. Mucilage contact angle increased with mucilage surface concentration (gram of dry mucilage per surface area); 3) Water could easily cross the rhizosphere when the mucilage concentration was below a given threshold. In contrast, above a critical mucilage concentration water could not flow through the rhizosphere. The critical mucilage concentration decreased with increasing particle size and decreasing matric potential.

Conclusions

These results show the importance of mucilage exudation for the water fluxes across the root-soil interface. Our percolation model predicts at what mucilage concentration the rhizosphere turns hydrophobic depending on soil texture and matric potential. Further studies are needed to extend these results to varying soil conditions and to upscale them to the entire root system.

     

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.     

Root effects on soil water and hydraulic properties

Plants can affect soil moisture and the soil hydraulic properties both directly by root water uptake and indirectly by modifying the soil structure. Furthermore, water in plant roots is mostly neglected when studying soil hydraulic properties. In this contribution, we analyze effects of the moisture content inside roots as compared to bulk soil moisture contents and speculate on implications of non-capillary-bound root water for determination of soil moisture and calibration of soil hydraulic properties. In a field crop of maize (Zea mays) of 75 cm row spacing, we sampled the total soil volumes of 0.7 m × 0.4 m and 0.3 m deep plots at the time of tasseling. For each of the 84 soil cubes of 10 cm edge length, root mass and length as well as moisture content and soil bulk density were determined. Roots were separated in 3 size classes for which a mean root porosity of 0.82 was obtained from the relation between root dry mass density and root bulk density using pycnometers. The spatially distributed fractions of root water contents were compared with those of the water in capillary pores of the soil matrix. Water inside roots was mostly below 2–5% of total soil water content; however, locally near the plant rows it was up to 20%. The results suggest that soil moisture in roots should be separately considered. Upon drying, the relation between the soil and root water may change towards water remaining in roots. Relations depend especially on soil water retention properties, growth stages, and root distributions. Gravimetric soil water content measurement could be misleading and TDR probes providing an integrated signal are difficult to interpret. Root effects should be more intensively studied for improved field soil water balance calculations. Presented at the International Conference on Bioclimatology and Natural Hazards, Pol’ana nad Detvou, Slovakia, 17–20 September 2007.     

Distribution of cutin and suberin biomarkers under forest trees with different root systems

Background and aims

The rhizosphere, the soil immediately surrounding roots, provides a critical bridge for water and nutrient uptake. The rhizosphere is influenced by various forms of root–soil interactions of which mechanical deformation due to root growth and its effects on the hydraulics of the rhizosphere are the least studied. In this work, we focus on developing new experimental and numerical tools to assess these changes.

Methods

This study combines X-ray micro-tomography (XMT) with coupled numerical simulation of fluid and soil deformation in the rhizosphere. The study provides a new set of tools to mechanistically investigate root-induced rhizosphere compaction and its effect on root water uptake. The numerical simulator was tested on highly deformable soil to document its ability to handle a large degree of strain.

Results

Our experimental results indicate that measured rhizosphere compaction by roots via localized soil compaction increased the simulated water flow to the roots by 27 % as compared to an uncompacted fine-textured soil of low bulk density characteristic of seed beds or forest topsoils. This increased water flow primarily occurred due to local deformation of the soil aggregates as seen in the XMT images, which increased hydraulic conductivity of the soil. Further simulated root growth and deformation beyond that observed in the XMT images led to water uptake enhancement of ~50 % beyond that due to root diameter increase alone and demonstrated the positive benefits of root compaction in low density soils.

Conclusions

The development of numerical models to quantify the coupling of root driven compaction and fluid flow provides new tools to improve the understanding of plant water uptake, nutrient availability and agricultural efficiency. This study demonstrated that plants, particularly during early growth in highly deformable low density soils, are involved in active mechanical management of their surroundings. These modeling approaches may now be used to quantify compaction and root growth impacts in a wide range of soils.     

Reverse flow in roots of Sesbania rostrata measured using the constant power heat balance method

This investigation was performed to examine qualitatively and quantitatively the reverse flow in partially dried roots of Sesbania rostrata using the constant power heat balance method. First, a semi-empirical technique for estimating sheath conductance of sap-flow sensors without assuming that sap flow is zero at night was proposed. Sap flow measured with the heat balance method was compared with water uptake as measured by a potometric method. Sap flow was overestimated by 56·1% for a 3·3-mm-diameter root, and by 40·0% for 6·1 mm and 33·3% for 8·8 mm roots. However, high correlation coefficients between the rates of water uptake and sap flow demonstrated that calibration would provide reliable values for root sap flow. To detect reverse flow, a split root experiment was conducted using a S. rostrata plant with its root system divided between dry and wet compartments. Daily sap flow of the drying compartment declined whereas that in ‘wet’ root increased, suggesting that the decrease in water uptake by ‘dry’ roots was offset by the ‘wet’ roots. Reverse flow was observed at night in the root on the dry side of the container when the soil water potential was less than –0·30 MPa. The total amount of water released into the soil during the night period was estimated to be 22·5 g.     

MAIN ROOT-LATERAL ROOT JUNCTIONS OF TWO DESERT SUCCULENTS: CHANGES IN AXIAL AND RADIAL COMPONENTS OF HYDRAULIC CONDUCTIVITY DURING DRYING

The influence of junctions between main roots and lateral roots on water flow was investigated for the desert succulents Agave deserti and Ferocactus acanthodes during 21 d of drying in soil. Under wet conditions, the junctions did not restrict xylem water flow from lateral roots to main roots, consistent with predictions of axial conductance based on vessel diameters. Embolism caused by drying reduced such axial conductance more at the junctions than in adjoining root regions. Connective tracheary elements at the junctions were abundantly pitted and had large areas of unlignified primary wall, apparently making them more susceptible to embolism than vessels or tracheids elsewhere in the roots. Unlike the decrease in axial conductance, the overall hydraulic conductivity of the junction increased during drying because of an increase in the conductivity of the radial pathway. Despite such increases, main roots may not lose substantial amounts of water to a dry soil during drought, initially because embolism at the junctions can limit xylem flow and later because soil hydraulic conductivity decreases. Moreover, the increased root hydraulic conductivity and a potentially rapid recovery from embolism by connective tracheary elements may favor water uptake near the junctions upon soil rewetting.     

Rectifier-like Activities of Roots of Two Desert Succulents

Axial and radial water flows for roots in response to appliedhydrostatic pressure drops, water loss from roots after variousperiods of drying, and development of new roots after rewettingdroughted plants were examined for two sympatric desert succulents.Agave deserti Engelm. and Ferocactus acanthodes (Lemaire) Brittonand Rose. For a 40 kPa hydrostatic pressure drop applied to20 mm long root pieces, radial water flows from the epidermisto the root xylem were 2- to 5-fold greater at the tip thanat midlength and were much less than axial flows along the xylem.Upon drying detached roots in air at 20 °C and a water vapoursaturation deficit of 1.2 kPa (50% relative humidity), radialwater flow decreased more than 10-fold in 3–6 h, and couldrecover to the original level 6 h after rewetting. The rateof water loss from attached roots of plants dried in air at20 °C and a 1.2 kPa saturation deficit decreased about 200-foldin 72 h, which would greatly limit water loss from the plantto a drying soil. At 96 h after rewetting roots of A. desertithat had been exposed to air at 20 °C and a 1.2 kPa saturationdeficit for 120 h, rehydration of existing roots and developmentof new roots contributed about equally to water uptake by thewhole plant. In summary, roots of these desert succulents canreadily take up water from a wet soil but do not lose much waterto a dry soil, thus effectively acting like rectifiers withrespect to plant-soil water movement. Key words: Agave, Cactus, Drought, Root, Water flow, Xylem     

Upscaling from Rhizosphere to Whole Root System: Modelling the Effects of Phospholipid Surfactants on Water and Nutrient Uptake

While the rhizosphere presents a different chemical, physical and biological environment to bulk soil, most experimental and modelling investigations of plant growth and productivity are based on bulk soil parameters. In this study, water and nutrient acquisition by wheat (Triticum aestivum L.) roots was investigated using rhizosphere- and root-system-scale modelling. The physical and chemical properties of rhizosphere soil could be influenced by phospholipid surfactants in the root mucilage. Two models were compared: a 2-dimensional (2D) Finite Element Method rhizosphere model, and a 3-dimensional (3D) root architecture model, ROOTMAP. ROOTMAP was parameterised to reproduce the results of the detailed 2D model, and was modified to include a rhizosphere soil volume. Lecithin (a phospholipid surfactant) could be exuded into the rhizosphere soil volume, decreasing soil water content and hydraulic conductivity at any given soil water potential, and decreasing phosphate adsorption to soil particles. The rhizosphere-scale modelling (5 × 5 mm² soil area, 10 mm root length, uptake over 12 h) predicted a reduction in water uptake (up to 16% at 30 kPa) and an increase in phosphate uptake (up to 4%) with lecithin exudation into the rhizosphere, but little effect on nitrate uptake, with only a small reduction in dry soil (1.6% at 200 kPa). The 3D root model reproduced the water (y = 1.013x, R² = 0.996), nitrate (y = 1x, R² = 1) and phosphate (y = 0.978x, R² = 0.998) uptake predictions of the rhizosphere model, providing confidence that a whole root system model could reproduce the dynamics simulated by a Finite Element Method rhizosphere model. The 3D root architecture model was then used to scale-up the rhizosphere dynamics, simulating the effect of lecithin exudation on water, nitrate and phosphate acquisition by a wheat root system, growing over 41 d. When applied to growing and responsive roots, lecithin exudation increased P acquisition by up to 13% in nutrient-rich, and 49% in relatively nutrient-poor soil. A comparison of wheat (Triticum aestivum L.) and lupin (Lupinus angustifolius L.) root architectures, suggested an interaction between the P acquisition benefit of rhizosphere lecithin and root architecture, with the more highly-branched wheat root structure acquiring relatively more P in the presence of lecithin than the sparsely-branched lupin root system.     

Partitioning water source and sinking process of a groundwater-dependent desert plant community

Aims

Root-specific responses to stress are not well-known, and have been largely based on indirect measurements of bulk soil water extraction, which limits mechanistic modeling of root function.

Methods

Here, we used neutron radiography to examine in situ root-soil water dynamics of a previously droughted black cottonwood (Populus trichocarpa) seedling, contrasting water uptake by the two major components of the root system that differed in initial recovery rate as apparent by ‘new’ (whiter, thinner), or ‘old’ (darker, thicker) parts of the fine root system.

Results

The smaller diameter ‘new’ roots had greater water uptake per unit surface area than the larger diameter ‘old’ roots, but they had less total surface area leading to less total water extraction; rates ranged from 0.0027–0.0116 g cm^?2 h^?1. The finest most-active roots were not visible in the radiographs, indicating the need to include destructive sampling. Analysis based on root-free bulk soil hydraulic properties indicated substantial redistribution of water via saturated/unsaturated flow and capillary wicking across the layers - suggesting water uptake dynamics following an infiltration event may be more complex than approximated by common soil hydraulic or root surface area modeling approaches.

Conclusions

Our results highlight the need for continued exploration of root-trait specific water uptake rates in situ, and impacts of roots on soil hydraulic properties – both critical components for mechanistic modeling of root function.

     

Abscisic acid signalling when soil moisture is heterogeneous: decreased photoperiod sap flow from drying roots limits abscisic acid export to the shoots

To investigate the contribution of different parts of the root system to total sap flow and leaf xylem abscisic acid (ABA) concentration ([X-ABA]_leaf), individual sunflower ( Helianthus annuus L.) shoots were grafted onto the root systems of two plants grown in separate pots and sap flow through each hypocotyl measured below the graft union. During deficit irrigation (DI), both pots received the same irrigation volumes, while during partial root zone drying (PRD) one pot ('wet') was watered and another ('dry') was not. During PRD, once soil water content ( θ ) decreased below a threshold, the fraction of sap flow from drying roots declined. As θ declined, root xylem ABA concentration increased in both irrigation treatments, and [X-ABA]_leaf increased in DI plants, but [X-ABA]_leaf of PRD plants actually decreased within a certain θ range. A simple model that weighted ABA contributions of wet and dry root systems to [X-ABA]_leaf according to the sap flow from each, better predicted [X-ABA]_leaf of PRD plants than either [X-ABA]_dry, [X-ABA]_wet or their mean. Model simulations revealed that [X-ABA]_leaf during PRD exceeded that of DI with moderate soil drying, but continued soil drying (such that sap flow from roots in drying soil ceased) resulted in the opposite effect.     

Measured and modelled differences in nutrient concentrations between rhizosphere and bulk soil in a Norway spruce stand

The gradient in soil characteristics from the bulk soil to the root surface is important to roots and to the organisms that live in the rhizosphere. Our ability to measure ion concentrations at the root surface is extremely limited, and models are largely untested. We used data from a well studied Norway spruce stand in SW Sweden to compare the measured difference in nutrient concentrations between rhizosphere and bulk soil with the difference predicted by a steady-state simulation model based on ecosystem budgets of nutrient uptake. The simulation model predicted depletion of NH₄, Ca, Mg, K in the rhizosphere, which shows that budgeted uptake rates were greater than the mass flow of bulk solution towards the root. In plots treated with ammonium sulphate, the model predicted an accumulation of S in the rhizosphere. In contrast, the observed rhizosphere concentrations were generally enriched in nutrients, relative to bulk soil. Collecting rhizosphere soil adhering to root surfaces may not be an appropriate method for describing the concentration gradient around the root. In addition, the simulation model omits some processes affecting conditions in the rhizosphere that are important to explaining nutrient uptake.     

Effects of liming and mycorrhizal colonization on soil phosphate depletion and phosphate uptake by maize (Zea mays L.) and soybean (Glycine max L.) grown in two tropical acid soils

The effects of liming and inoculation with the arbuscular mycorrhizal fungus, Glomus intraradices Schenck and Smith on the uptake of phosphate (P) by maize (Zea mays L.) and soybean (Glycine max [L.] Merr.) and on depletion of inorganic phosphate fractions in rhizosphere soil (Al-P, Fe-P, and Ca-P) were studied in flat plastic containers using two acid soils, an Oxisol and an Ultisol, from Indonesia. The bulk soil pH was adjusted in both soils to 4.7, 5.6, and 6.4 by liming with different amounts of CaCO₃.In both soils, liming increased shoot dry weight, total root length, and mycorrhizal colonization of roots in the two plant species. Mycorrhizal inoculation significantly increased root dry weight in some cases, but much more markedly increased shoot dry weight and P concentration in shoot and roots, and also the calculated P uptake per unit root length. In the rhizosphere soil of mycorrhizal and non-mycorrhizal plants, the depletion of Al-P, Fe-P, and Ca-P depended in some cases on the soil pH. At all pH levels, the extent of P depletion in the rhizosphere soil was greater in mycorrhizal than in non-mycorrhizal plants. Despite these quantitative differences in exploitation of soil P, mycorrhizal roots used the same inorganic P sources as non-mycorrhizal roots. These results do not suggest that mycorrhizal roots have specific properties for P solubilization. Rather, the efficient P uptake from soil solution by the roots determines the effectiveness of the use of the different soil P sources. The results indicate also that both liming and mycorrhizal colonization are important for enhancing P uptake and plant growth in tropical acid soils.     

Influence of temperature and soil drying on respiration of individual roots in citrus: integrating greenhouse observations into a predictive model for the field

In citrus, the majority of fine roots are distributed near the soil surface – a region where conditions are frequently dry and temperatures fluctuate considerably. To develop a better understanding of the relationship between changes in soil conditions and a plant’s below‐ground respiratory costs, the effects of temperature and soil drying on citrus root respiration were quantified in controlled greenhouse experiments. Chambers designed for measuring the respiration of individual roots were used. Under moist soil conditions, root respiration in citrus increased exponentially with changes in soil temperature (Q₁₀ = 1·8–2·0), provided that the changes in temperature were short‐term. However, when temperatures were held constant, root respiration did not increase exponentially with increasing temperatures. Instead, the roots acclimated to controlled temperatures above 23 °C, thereby reducing their metabolism in warmer soils. Under drying soil conditions, root respiration decreased gradually beginning at 6% soil water content and reached a minimum at <2% soil water content in sandy soil. A model was constructed from greenhouse data to predict diurnal patterns of fine root respiration based on temperature and soil water content. The model was then validated in the field using data obtained by CO₂ trapping on root systems of mature citrus trees. The trees were grown at a site where the soil temperature and water content were manipulated. Respiration predicted by the model was in general agreement with observed rates, which indicates the model may be used to estimate entire root system respiration for citrus.     

Root growth, developmental changes in the apex, and hydraulic conductivity for Opuntia ficus-indica during drought

Developmental changes in the root apex and accompanying changes in lateral root growth and root hydraulic conductivity were examined for Opuntia ficus-indica (L.) Miller during rapid drying, as occurs for roots near the soil surface, and more gradual drying, as occurs in deeper soil layers. During 7 d of rapid drying (in containers with a 3-cm depth of vermiculite), the rate of root growth decreased sharply and most root apices died; such a determinate pattern of root growth was not due to meristem exhaustion but rather to meristem mortality after 3 d of drying. The length of the meristem, the duration of the cell division cycle, and the length of the elongation zone were unchanged during rapid drying. During 14 d of gradual drying (in containers with a 6-cm depth of vermiculite), root mortality was relatively low; the length of the elongation zone decreased by 70%, the number of meristematic cells decreased 30%, and the duration of the cell cycle increased by 36%. Root hydraulic conductivity ( L _P) decreased to one half during both drying treatments; L _P was restored by 2 d of rewetting owing to the emergence of lateral roots following rapid drying and to renewed apical elongation following gradual drying. Thus, in response to drought, the apical meristems of roots of O. ficus-indica near the surface die, whereas deeper in the substrate cell division and elongation in root apices continue. Water uptake in response to rainfall in the field can be enhanced by lateral root proliferation near the soil surface and additionally by resumption of apical growth for deeper roots.     

Recovery of soil respiration after drying

Soils are frequently exposed to drying and wetting events and previous studies have shown that rewetting results in a strong but short-lived flush of microbial activity. The aim of this study was to determine the effect of the water content during the dry period on the size and duration of the flush and on the rate of recovery. Two soils (a sand and a sandy loam) were maintained at different water contents (WC) 30, 28 and 25 g water kg^?1 soil (sand) and 130, 105 and 95 g water kg^?1 soil (sandy loam) for 14 days, then rewet to the water content at which respiration was optimal [WC 35 (sand), WC200 (sandy loam)] and maintained at this level until day 68. Ground pea straw (C/N 26) was added and incorporated on day 1. The controls were maintained at the optimal water content throughout the 68 days. Respiration rates during the dry phase (days 1?C14) decreased with decreasing water content. The flush of respiration after rewetting peaked on day 15 in the sandy loam and on day 16 in the sand; it was greatest in the soils that had been maintained at the lowest water content [WC25 (sand) and WC95 (sandy loam)]. Cumulative respiration during the remainder of the incubation period in which all soils were maintained at optimal water content increased more strongly in the soils that had been dry compared to the constantly moist control. On the final day of the dry period (day 14), cumulative respiration in the dry soils was 29?C65% (sand) and 67?C94% (sandy loam) of the constantly moist control whereas on day 68 it was 80?C84% (sand) and 86?C96% (sandy loam). The greater increase in cumulative respiration in the previously dry soils can be explained by the reduced decomposition rates during the dry period which resulted in higher substrate availability on day 14 compared to the constantly moist control. Microbial community structure assessed by phospholipid fatty acid analyses changed over time in all treatments but was less affected by water content than respiration; it differed only between the highest and the lowest water content. These differences were maintained throughout the incubation period in the sandy loam and transiently in the sand. It can be concluded that the soil water content during the dry phase affects the size of the flush in microbial activity upon rewetting and that microbial activity in previously dried soils may not be fully restored even after 54 days of moist incubation, suggesting that drying of soil can have a significant and long-lasting impact on microbial functioning.     

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.     

Water uptake resumption following soil drought: a comparison among four barley genotypes

Soil water resulting from episodic growing season rainfall evaporatesrapidly in semi-arid regions. Plants may mnot benefit from suchwater additions if near-surface roots are unable to resume wateruptake rapidly following periods of soil water deficit. Ourobjectives were to develop a means of quantifying root uptakeresponses in the upper soil layer following rewetting aftersoil water deficit, and to evaluate the existence of genotypicdifferences among four diverse barley (Hordeum vulgare L.) genotypesin this regard. Plants were grown in replicate soil columnshaving hydraulically isolated surface and subsoil layers, andinstrumented with time-domain reflectometry (TDR) waveguides.The upper 0.05 m soil layer was allowed to dry to —1.8to —3.0 MPa for 10-14 d, during which time subsoil wetnesswas maintained at about —0.6 to —0.7 MPa. The time-courseof soil water uptake was monitored at 0.5 h intervals followingrewetting of the surface layer. Substantial water uptake began1 d after rewetting following 10 d, and 2-3 d after rewettingfollowing 14 d of water deficit. Rate of water uptake was morerapid in response to a second rewetting 5-7 d later. Consistentgenotypic responses in terms of cumulative water uptake on awhole plant and leaf area-specific basis were observed duringeach trial. These results have application to evaluating droughthardiness and interspecific competitive ability under semi-aridconditions, and to investigations of root physiological andmorphological changes that contribute to recovery from waterdeficit Key words: Hordeum vulgare, root water uptake, soil water deficit, time-domain reflectometry     

Temperature Effects on Recovery Time of Bacterial Growth After Rewetting Dry Soil

The effect of temperature on the recovery of bacterial growth after rewetting dry soil was measured in a soil that responded with bacterial growth increasing immediately upon rewetting in a linear fashion (type (i) response sensu Meisner et al. (Soil Biol Biochem 66: 188-192, 2013)). The soil was air-dried for 4 days and then rewetted at different temperatures. Bacterial growth over time was then estimated using the leucine incorporation method. At 25 °C, the recovery of bacterial growth to levels of a wet control soil was rapid, within 6 h, while at 15 °C, recovery time increased to around 60 h, becoming more than a week at 5 °C. The temperature dependency of the recovery time was well modeled by a square root function. Thus, temperature will not only directly affect growth rates but also affect length of transition periods, like resuscitation after a drying event. The temperature during the rewetting event thus has to be taken into consideration when analyzing the microbial response dynamics.