Downward flux of water through roots (i.e. inverse hydraulic lift) in dry Kalahari sands

Downward transport of water in roots, in the following termed “inverse hydraulic lift,” has previously been shown with heat flux techniques. But water flow into deeper soil layers was demonstrated in this study for the first time when investigating several perennial grass species of the Kalahari Desert under field conditions. Deuterium labelling was used to show that water acquired by roots from moist sand in the upper profile was transported through the root system to roots deeper in the profile and released into the dry sand at these depths. Inverse hydraulic lift may serve as an important mechanism to facilitate root growth through the dry soil layers underlaying the upper profile where precipitation penetrates. This may allow roots to reach deep sources of moisture in water-limited ecosystems such as the Kalahari Desert. Received: 1 January 1998 / Accepted: 1 April 1998     

Nitrogen Controls on Fine Root Substrate Quality in Temperate Forest Ecosystems

Nitrogen controls on fine root substrate quality (that is, nitrogen and carbon-fraction concentrations) were assessed using nitrogen availability gradients in the Harvard Forest chronic nitrogen addition plots, University of Wisconsin Arboretum, Blackhawk Island, Wisconsin, and New England spruce-fir transect. The 27 study sites encompassed within these four areas collectively represented a wide range of nitrogen availability (both quantity and form), soil types, species composition, aboveground net primary production, and climatic regimes. Changes in fine root substrate quality among sites were most frequently and strongly correlated with nitrate availability. For the combined data set, fine root nitrogen concentration increased (adjusted R ² = 0.46, P < 0.0001) with increasing site nitrate availability. Fine root “extractive” carbon-fraction concentrations decreased (adjusted R ² = 0.32, P < 0.0002), “acid-soluble” compounds increased (adjusted R ² = 0.35, P < 0.0001), and the “acid-insoluble” carbon fraction remained relatively high and stable (combined mean of 48.7 ± 3.1% for all sites) with increasing nitrate availability. Consequently, the ratio of acid-insoluble C–total N decreased (adjusted R ² = 0.40, P < 0.0001) along gradients of increasing nitrate availability. The coefficients of determination for significant linear regressions between site nitrate availability and fine root nitrogen and carbon-fraction concentrations were generally higher for sites within each of the four study areas. Within individual study sites, tissue substrate quality varied between roots in different soil horizons and between roots of different size classes. However, the temporal variation of fine root substrate quality indices within specific horizons was relatively low. The results of this study indicate that fine root substrate quality increases with increasing nitrogen availability and thus supports the substrate quality component of a hypothesized conceptual model of nitrogen controls on fine root dynamics that maintains that fine root production, mortality, substrate quality, and decomposition increase with nitrogen availability in forest ecosystems in a manner that is analogous to foliage.     

Hydraulic lift: consequences of water efflux from the roots of plants

Hydraulic lift is the passive movement of water from roots into soil layers with lower water potential, while other parts of the root system in moister soil layers, usually at depth, are absorbing water. Here, we review the brief history of laboratory and field evidence supporting this phenomenon and discuss some of the consequences of this below-ground behavior for the ecology of plants. Hydraulic lift has been shown in a relatively small number of species (27 species of herbs, grasses, shrubs, and trees), but there is no fundamental reason why it should not be more common as long as active root systems are spanning a gradient in soil water potential (Ψ_s) and that the resistance to water loss from roots is low. While the majority of documented cases of hydraulic lift in the field are for semiarid and arid land species inhabiting desert and steppe environments, recent studies indicate that hydraulic lift is not restricted to these species or regions. Large quantities of water, amounting to an appreciable fraction of daily transpiration, are lifted at night. This temporary partial rehydration of upper soil layers provides a source of water, along with soil moisture deeper in the profile, for transpiration the following day and, under conditions of high atmospheric demand, can substantially facilitate water movement through the soil-plant-atmosphere system. Release of water into the upper soil layers has been shown to afford the opportunity for neighboring plants to utilize this source of water. Also, because soils tend to dry from the surface downward and nutrients are usually most plentiful in the upper soil layers, lifted water may provide moisture that facilitates favorable biogeochemical conditions for enhancing mineral nutrient availability, microbial processes, and the acquisition of nutrients by roots. Hydraulic lift may also prolong or enhance fine-root activity by keeping them hydrated. Such indirect benefits of hydraulic lift may have been the primary selective force in the evolution of this process. Alternatively, hydraulic lift may simply be the consequence of roots not possessing true rectifying properties (i.e., roots are leaky to water). Finally, the direction of water movement may also be downward or horizontal if the prevailing Ψ_s gradient so dictates, i.e., inverse, or lateral, hydraulic lift. Such downward movement through the root system may allow growth of roots in otherwise dry soil at depth, permitting the establishment of many phreatophytic species. Received: 2 June 1997 / Accepted: 24 September 1997     

Allometric Effects of <Emphasis Type="Italic">Agriophyllum squarrosum</Emphasis> in Response to Soil Nutrients,Water, and Population Density in the Horqin Sandy Land of China

We monitored the allometric effects for greenhouse-grown Agriophyllum squarrosum plants in response to variations in population density and the availability of soil nutrients and water. Biomass allocations were size-dependent. The plasticity of roots, stems, leaves, and reproductive effort was “true” in response to changes in nutrient content. At a low level of soil minerals, plants allocated more resources to the development of roots and reproductive organs than to leaves, but data for stem allocations were consistent for tradeoffs between the effects of nutrients and plant size. The plasticities of leaf allocation and reproductive effort were “true” whereas those of root and stem allocations were “apparent” in response to fluctuations in soil water, being a function of plant size. Decreasing soil water content was associated with higher leaf allocation and lower reproductive effort. Except for this “apparent” plasticity of leaf allocation, none was detected with population density on biomass allocation. Architectural traits were determinants of the latter. For roots, the determining trait was the ratio of plant height to total biomass; for stems and reproduction, plant height; and for leaves, the ratio of branch numbers to plant height.     

The importance of root gravitropism for inter-root competition and phosphorus acquisition efficiency: results from a geometric simulation model

We have observed that low soil phosphorus availability alters the gravitropic response of basal roots in common bean (Phaseolus vulgaris L.), resulting in a shallower root system. In this study we use a geometric model to test the hypotheses that a shallower root system is a positive adaptive response to low soil P availability by (1) concentrating root foraging in surface soil horizons, which generally have the highest P availability, and (2) reducing spatial competition for P among roots of the same plant. The growth of nine root systems contrasting in gravitropic response over 320 h was simulated in SimRoot, a dynamic three-dimensional geometric model of root growth and architecture. Phosphorus acquisition and inter-root competition were estimated with Depzone, a program that dynamically models nutrient diffusion to roots. Shallower root systems had greater P acquisition per unit carbon cost than deeper root systems, especially in older root systems. This was due to greater inter-root competition in deeper root systems, as measured by the volume of overlapping P depletion zones. Inter-root competition for P was a significant fraction of total soil P depletion, and increased with increasing values of the P diffusion coefficient (De), with root age, and with increasing root gravitropism. In heterogenous soil having greater P availability in surface horizons, shallower root systems had greater P acquisition than deeper root systems, because of less inter-root competition as well as increased root foraging in the topsoil. Root P acquisition predicted by SimRoot was validated against values for bean P uptake in the field, with an r ² between observed and predicted values of 0.75. Our results support the hypothesis that altered gravitropic sensitivity in P-stressed roots, resulting in a shallower root system, is a positive adaptive response to low P availability by reducing inter-root competition within the same plant and by concentrating root activity in soil domains with the greatest P availability. This revised version was published online in August 2006 with corrections to the Cover Date.     

Transverse hydraulic redistribution by a grapevine

Root hydraulic redistribution has been shown to occur in numerous plant species under both field and laboratory conditions. To date, such water redistribution has been demonstrated in two fundamental ways, either lifting water from deep edaphic sources to dry surface soils or redistributing water downward (reverse flow) when inverted soil Ψ_s gradients exist. The importance of hydraulic redistribution is not well documented in agricultural ecosystems under field conditions, and would be important because water availability can be temporally and spatially constrained. Herein we report that a North American grapevine hybrid (Vitis riparia × V. berlandieri cv 420 A) growing in an agricultural ecosystem can redistribute water from a restricted zone of available water under a drip irrigation emitter, laterally across the high resistance pathways of the trunk and into roots and soils on the non-irrigated side. Deuterium-labelled water was used to demonstrate lateral movement across the vine's trunk and reverse flow into roots. Water redistribution from the zone of available water and into roots distant from the source occurred within a relatively short time frame of 36 h, although overnight deposition into rhizosphere soils around the roots was not detected. Deuterium was eventually detected in rhizosphere soils adjacent to roots on the non-irrigated side after 7 d. Application of identical amounts of water with the same deuterium enrichment level (2%) to soils without grapevine roots showed that physical transport of water through the vapour phase could not account for either downward or transverse movement of the label. These results confirmed that root presence facilitated the transport of label into soils distant from the wetted zone. When deuterium-labelled water was allowed to flow directly into the trunk above the root–trunk interface, reverse flow occurred and lateral movement across the trunk and into roots originating around the collar region did not encounter large disproportionate resistances. Rapid redistribution of water into the entire root system may have important implications for woody perennial cultivars growing where water availability is spatially heterogeneous. Under the predominantly dry soil conditions studied in this investigation, water redistributed into roots may extend root longevity and increase the vines water capacitance during periods of high transpiration demand. These benefits would be enhanced by diminished water loss from roots, and could be equally important to other cited benefits of hydraulic redistribution into soils such as enhancement of nutrient acquisition.     

Effects of soil structural discontinuity on root and shoot growth and water use of maize

The role of roots penetrating various undisturbed soil horizons beneath loose layer in water use and shoot growth of maize was evaluated in greenhouse experiment. 18 undisturbed soil columns 20 cm in diameter and 20 cm in height were taken from the depths 30–50 cm and 50–70 cm from a Brown Lowland soil, a Pseudogley and a Brown Andosol (3 columns from each depth and soil). Initial resistance to penetration in undisturbed soil horizons varied from 2.5 to 8.9 MPa while that in the loose layer was 0.01 MPa. The undisturbed horizons had a major effect on vertical arrangement of roots. Root length density in loose layer varied from 96 to 126 km m^-3 while in adjacent stronger top layers of undisturbed horizons from 1.6 to 20.0 km m^-3 with higher values in upper horizons of each soil. For specific root length, the corresponding ranges were 79.4–107.7 m g^-1 (on dry basis) and 38.2–63.7 m g^-1, respectively. Ratios of root dry weight per unit volume of soil between loose and adjacent undisturbed layers were much lower than those of root length density indicating that roots in undisturbed horizons were produced with considerably higher partition of assimilates. Root size in undisturbed horizons relative to total roots was from 1.1 to 38.1% while water use from the horizons was from 54.1 to 74.0%. Total water use and shoot growth were positively correlated with root length in undisturbed soil horizons. There was no correlation between shoot growth and water use from the loose layers.     

Modelling the effect of varying soil water on root growth dynamics of annual crops

We present a simple framework for modelling root growth and distribution with depth under varying soil water conditions. The framework considers the lateral growth of roots (proliferation) and the vertical extension of roots (root front velocity). The root front velocity is assumed to be constant when the roots descend into an initially wet soil profile. The lateral growth of roots is governed by two factors: (1) the current root mass or root length density at a given depth, and (2) soil water availability at that depth.Under non-limiting soil water conditions, the increase in root mass at any depth is governed by a logistic equation so that the root length density (R _v) cannot exceed the maximum value. The maximumR _v, is assumed to be the same for all depths. Additional dry matter partitioned to roots is initially distributed according to the current root mass at each depth. As the root mass approaches the maximum value, less dry matter is partitioned to that depth.When soil water is limiting, a water deficit factor is introduced to further modify the distribution of root dry matter. It is assumed that the plant is an energy minimiser so that more root mass is partitioned to the wetter regions of the soil where least energy will be expended for root growth. Hence, the model allows for enhanced root growth in areas where soil water is more easily available.Simulation results show that a variety of root distribution patterns can be reproduced due to varying soil water conditions. It has been demonstrated that broad patterns of root distribution reported in the literature can also be simulated by the model.     

Root dynamics of Melaleuca halmaturorum in response to fluctuating saline groundwater

Melaleuca halmaturorum is a salt and waterlogging tolerant tree and thus often occurs in saline areas fringing permanent wetlands and in ephemeral swamps. The dominance of this tree in natural groundwater discharge areas may result in M. halmaturorum transpiration making a major contribution to groundwater discharge. To quantify this the seasonal changes in tree water sources in response to fluctuating soil salinity and waterlogging were examined. This study was conducted in a natural system where seasonally fluctuating saline groundwater (64 dS m^–1; 0.3–1.2 m deep) allowed the patterns of M. halmaturorum root water uptake to be followed over a 15 month period. Tree water sources were examined using the naturally occurring stable isotopes of water, while new root growth was examined using a field root observation window and from soil cores. The presence of isotopic fractionation of ²H under conditions of soil salinity and waterlogging was tested in a glasshouse experiment. Measurements of soil and leaf water potential were also made to examine the possible water sources and limits to water uptake. No isotopic fractionation was found by tree roots under conditions of salinity and waterlogging. M. halmaturorum trees were active in taking up groundwater at most times and combined this with a shallower soil water source replenished by rainfall in winter. Water uptake was concentrated in the deeper parts of the soil profile when the groundwater was at its deepest and salt had accumulated in the surface soils, at the end of summer. When groundwater rose, at the end of winter, roots responded by extracting water from near the soil surface (0–0.1 m), at the new watertable. This pattern of water uptake in response to groundwater fluctuations and salt accumulation in the surface soil was also reflected in new root tip appearance at the root observation window. Fluctuations in leaf water potential fallowed fluctuations in surface soil (0.1 m depth) water potential at all times. In winter leaf water potential reflected the absolute values of the surface soil water potential but in summer it was between surface soil and groundwater water potentials. We conclude that M. halmaturorum used groundwater in summer and a combination of rainfall and groundwater from the surface soils in winter. The ability to take up water from saline substrates through the maintenance of low leaf water potential, combined with this ability to rapidly alter root water uptake in response to changes in soil water availability contributed to the survival of M. halmaturorum in this saline swamp.     

Water relations and root activities of Buchloe dactyloides and Zoysia japonica in response to localized soil drying

Effects of localized soil drought stress on water relations, root growth, and nutrient uptake were examined in drought tolerant ‘Prairie’ buffalograss [Buchloe dactyloides (Nutt.) Engelm.] and sensitive ‘Meyer’ zoysiagrass (Zoysia japonica Steud.). Grasses were grown in small rhizotrons in a greenhouse and subjected to three soil moisture regimes: (1) watering the entire 80-cm soil profile (well-watered control); (2) drying 0–40 cm soil and watering the lower 40 cm (partially dried); (3) and drying the entire soil profile (fully dried). Drying the 0–40 cm soil for 28 days had no effect on leaf water potential (Ψ _leaf ) in Prairie buffalograss compared to the well-watered control but reduced that in Meyer zoysiagrass. Root elongation rate was greater for Prairie buffalograss than Meyer zoysiagrass under well-watered or fully dried conditions. Rooting depth increased with surface soil drying; with Prairie buffalograss having a larger proportion of roots in the lower 40 cm than Meyer zoysiagrass. The higher rates of water uptake in the deeper soil profile in the partially dried compared to the well-watered treatment and by Prairie buffalograss compared to Meyer zoysiagrass could be due to differences in root distribution. Root ¹⁵N uptake for Prairie buffalograss was higher in 0–20 cm drying soil in the partially dried treatment than in the fully dried treatment. Diurnal fluctuations in soil water content in the upper 20 cm of soil when the lower 40 cm were well-watered indicated water efflux from the deeper roots to the drying surface soil. This could help sustain root growth, maintain nutrient uptake in the upper drying soil layer, and prolong turfgrass growth under localized drying conditions, especially for the deep-rooted Prairie buffalograss. This revised version was published online in June 2006 with corrections to the Cover Date.