Water-Transporting System in Higher Plants and Its Elements: 6. Relationship between Root Pressure and Transpiration in an Intact Plant

Novel experimental data were adapted to a previously devised concept of water relations in plant roots. The improved concept explains the formation in the root stele of a high water potential that exceeds the water potential of the root environment. The physical basis of this phenomenon relies on active metabolism of the peripheral root zone (cortex) and, more importantly, on the unloading of assimilates from the root central cylinder (stele) to the outer cylinder (cortex). The unloading not only raises the water potential of the root stele but also increases the hydraulic resistance; these two factors account for the elevation of root pressure. The process of unloading assimilates from the root stele to the cortex is apparently promoted by the transit of the ascending water flow through the cortex. This flow, enriched with the dissolved oxygen of the root environment, stimulates the unloading and metabolic conversion of assimilates in the root symplast.     

Effects of phosphate and pH stress on the growth and function of apple roots

Summary Effects of phosphate and pH stress on the growth and uptake functions of apple roots were studied over a period of fourteen days using split-root (2-way) seedlings in solution culture. The level of P fed to either or both halves of the root system was varied and a demineralized water control was also included. pH treatments consisted of using acidic nutrient solutions (pH 3 to 4) or nutrient solutions adjusted to pH 5.0 before use.Solution pH proved of paramount importance for the expression of P deficiency effects on root growth and water uptake. Where initial solution pH was favourable for root growth (pH 5), P deficiency stimulated root growth and water uptake per seedling even if the stress was localized. On the other hand, acidic solutions and the water control inhibited root growth and water uptake compared with +P controls. Where solution pH was favourable, P stress also led to an increase in the mean length per root versus the +P control suggesting that the plant adapted to stress by developing an exploratory type of root.Water use per seedling was predominantly a function of root size rather than leaf area since the treatments influenced root size to a much greater extent than leaf area. Uptake was positively related to root size in that adjusted solutions gave a higher water use than nonadjusted solutions. However, efficiency of water use per unit weight of root was consistently higher in the nonadjusted solutions and this appeared to be due to the presence of a larger number of root tips per unit weight of root in such solutions compared with root systems in pH adjusted solutions.Uptake of P per half root was higher from pH adjusted than from nonadjusted solutions and was also increased by increasing the P concentration. Further, for any one treatment P uptake per half root increased throughout the experiment indicating that uptake was influenced by root growth. However, in contrast to water uptake, uptake of P per unit weight or per unit surface area of root was not changed by pH adjustment nor was this parameter of uptake concentration dependent. That is, the above-mentioned pH and concentration effects on P uptake were mediated through effects on root growth.Comparing localized versus uniform placement of P, uptake of P was significantly higher from the uniform application. However, uptake from localized placement at pH 5 was markedly higher than uptake under pH stress and therefore if the pH of the medium remains favourable for root growth then the lower value for localized placement could probably be compensated for by further increasing the concentration of P applied.     

Plantlet morphology and the regulation of net water loss in tissue-cultured Douglas-fir

Tissue-culture plantlets of Douglas-fir [ Pseudotsuga menziesii (Mirb.) Franco] were highly susceptible to detrimental water loss upon removal from culture in vitro. Control of net water loss was related to shoot and root morphology. Relative water content after 3 h of atmospheric water stress was positively correlated to root number, root surface area, and the length of the longest root, and was inversely correlated to the ratio of needle surface area/root surface area. High relative water content apparently was a result of a higher rate of water uptake among plantlets with beneficial morphological features. It is recommended that, to improve the ability of a plantlet to withstand water stress during acclimatization, beneficial root system features be focused upon during plantlet production.     

Wheat root diversity and root functional characterization

Background and Aims

Under limited moisture conditions, roots can play an outstanding role with respect to yield stability by effective absorption of water from soil. A targeted integration of root traits into plant breeding programs requires knowledge on the existing root diversity and access to easy and cost-effective methods. This study aimed to assess wheat root diversity, root properties in relation to water regime, and the efficiency of root capacitance for in situ screening.

Methods

Root morphological, anatomical properties and root capacitance of wheat species from different ploidy levels were studied under field conditions in 2 years contrasting in water regime. Soil water content was weekly measured.

Results

Significant genotypic differences were observed for most root traits. The investigated genotypes exploited different strategies to maximize soil water depletion, e.g. high topsoil root length density, low tissue mass density, high specific root length, deep rooting and looser xylem vessels. Multivariate statistics of root traits revealed an acceptable genotypic differentiation according to regional origin, genetics and capacity to extract soil water.

Conclusions

Under supply-driven environments, dehydration avoidance via water uptake maximization can be achieved through high topsoil rooting density. In this regard, root capacitance can be useful for in situ screening.     

Water use and sodium chloride uptake by apple trees

Summary Apple trees grown with their root systems split into halves were used to study the effects of non-uniform salinity stress within a root system upon salt and water uptake. Water uptake declined rapidly when sodium chloride solution (90 meq l⁻¹) was added to any root zone but uptake increased correspondingly in the non-saline root zone of each tree. This changed pattern of water uptake with partial salinization did not change the total water use by the trees compared with their water use when neither root zone was salt stressed. After a‘steady-state’ condition of water uptake had been reached 80 to 85% of the water was taken up in the non-saline root zone. Irrigation at three soil matric potential intervals of −6.6, −33 and −66 kPa allowed to develop in the non-saline root zone of each tree did not affect water use responses. Leaf concentrations of Ca, Mg and K were unaffected by treatments. Chloride and Na concentrations increased in leaves with exposure to salinity stress in half root zones and with increasing soil matric potential stress. Some evidence was obtained using tritium enriched water that water was transferred from a non-saline root zone into a saline root zone but the volume involved was unmeasurable.     

Root growth and water uptake in winter wheat under deficit irrigation

Root growth is critical for crops to use soil water under water-limited conditions. A field study was conducted to investigate the effect of available soil water on root and shoot growth, and root water uptake in winter wheat (Triticum aestivum L.) under deficit irrigation in a semi-arid environment. Treatments consisted of rainfed, deficit irrigation at different developmental stages, and adequate irrigation. The rainfed plots had the lowest shoot dry weight because available soil water decreased rapidly from booting to late grain filling. For the deficit-irrigation treatments, crops that received irrigation at jointing and booting had higher shoot dry weight than those that received irrigation at anthesis and middle grain filling. Rapid root growth occurred in both rainfed and irrigated crops from floral initiation to anthesis, and maximum rooting depth occurred by booting. Root length density and dry weight decreased after anthesis. From floral initiation to booting, root length density and growth rate were higher in rainfed than in irrigated crops. However, root length density and growth rate were lower in rainfed than in irrigated crops from booting to anthesis. As a result, the difference in root length density between rainfed and irrigated treatments was small during grain filling. The root growth and water use below 1.4 m were limited by a caliche (45% CaCO₃) layer at about 1.4 m profile. The mean water uptake rate decreased as available soil water decreased. During grain filling, root water uptake was higher from the irrigated crops than from the rainfed. Irrigation from jointing to anthesis increased seasonal evapotranspiration, grain yield, harvest index and water-use efficiency based on yield (WUE), but did not affect water-use efficiency based on aboveground biomass. There was no significant difference in WUE among irrigation treatments except one-irrigation at middle grain filling. Due to a relatively deep root system in rainfed crops, the higher grain yield and WUE in irrigated crops compared to rainfed crops was not a result of rooting depth or root length density, but increased harvest index, and higher water uptake rate during grain filling.     

Partition of photosynthates between shoot and root in spring wheat (Triticum aestivum L.) as a function of soil water potential and root temperature

Low soil water potential and low or high root temperatures are important stresses affecting carbon allocation in plants. This study examines the effects of these stresses on carbon allocation from the perspective of whole plant mass balance. Sixteen-day old spring wheat seedlings were placed in a growth room under precisely controlled root temperatures and soil water potentials. Five soil water potential treatments, from −0.03 MPa to −0.25 MPa, and six root temperature treatments, from 12 to 32°C were used. A mathematical model based on mass balance considerations was used, in combination with experimental measurements of rate of net photosynthesis, leaf area, and shoot/root dry masses to determine photosynthate allocation between shoot and root. Partitioning of photosynthates to roots was the lowest at 22–27°C root temperature regardless soil water potential, and increased at both lower and higher root temperatures. Partitioning of photosynthates to the roots increased with decreasing soil water potential. Under the most favourable conditions, i.e. at −0.03 MPa soil water potential and 27°C root temperature, the largest fraction, 57%, of photosynthates was allocated to the shoots. Under the most stressed conditions, i.e. at −0.25 MPa soil water potential and 32°C root temperature, the largest fraction, more than 80%, of photosynthates was allocated to roots.     

Effect of water stress on root meristems in woody and herbaceous plants during the first stage of development

We investigated the effect of water stress on the root system architecture of pine saplings and pea seedlings during the first stage of development. Attention was focused on meristematic tissue situated at the root tip because of the leading role played by the tissue in the planning of root system architecture. The data showed that both species are extremely sensitive and that plants arrest their growth immediately during water stress treatment. When stress treatment was not intense, both species recovered growth but presented modifications in the root system architecture. In pine saplings, the modification in root system architecture was the consequence of fine root meristems not recovering from water stress. The saplings survived by producing new lateral meristems from the cortical tannin zone above the fine root tip. In the case of pea seedlings, the meristematic tissues in the primary root arrested proliferation during water stress although they recovered when the event occurred during the first hours of germination. The response was different when water stress was enforced on older seedlings. In this case, root meristems never completely recovered their proliferation despite the increase in proline content observed in the cells. The modification of root system architecture in pea seedlings depended on the arrest of primary root elongation and the formation of new root laterals. As regards the primary roots, water stress treatment induced along the axis the formation of irregular ‘swellings’ in the cortical zone above the meristematic zone. Anatomical investigations suggested that such swellings may have derived from the changes in elongation direction of derivatives. The formation of new laterals was observed in hydroponic cultures when water stress treatment was enforced slowly and prolonged for a long time. The production of new lateral meristems may have been a similar response of woody and herbaceous plants to water stress conditions. It is not known whether these new meristems present characteristics of resistance to water stress. This revised version was published online in June 2006 with corrections to the Cover Date.     

Use of the root contact concept,an empirical leaf conductance model and pressure-volume curves in simulating crop water relations

A simulation model “DanStress” was developed for studying the integrated effects of soil, crop and climatic conditions on water relations and water use of field grown cereal crops. The root zone was separated into 0.1 m deep layers of topsoil and subsoil. For each layer the water potential at the root surface was calculated by a single root model, and the uptake of water across the root was calculated by a root contact model. Crop transpiration was calculated by Monteith's combination equation for vapour flow. Crop conductance to water vapour transfer for use in Monteith's combination equation was scaled up from an empirical stomatal conductance model used on sunlit and shaded crop surfaces of different crop layers. In the model, transpirational water loss originates from root water uptake and changes in crop water storage. Crop water capacitance, used for describing the water storage, was derived from the slope of pressure-volume (PV) curves of the leaves. PV curves were also used for deriving crop water potential, osmotic potential, and turgor pressure. The model could simulate detailed diurnal soil-crop water relations during a 23-day-drying cycle with time steps of one hour. During the grain filling period in spring barley (Hordeum distichum L.), grown in a sandy soil in the field, measured and predicted values of leaf water and osmotic potential, RWC, and leaf stomatal conductance were compared. Good agreement was obtained between measured and predicted values at different soil water deficits and climatic conditions. In the field, measured and predicted volumetric soil water contents (θ) of topsoil and subsoil layers were also compared during a drying cycle. Predicted and measured θ-values as a function of soil water deficits were similar suggesting that the root contact model approach was valid. From the investigation we concluded: (I) a model, which takes the degree of contact between root surface and soil water into account, can be used in sandy soil for calculation of root water uptake, so that the root conductance during soil water depletion only varies by the degree of contact; (II) crop conductance, used for calculation of crop transpiration, can be scaled up from an empirical single leaf stomatal conductance model controlled by the level of leaf water potential and micrometeorological conditions; (III) PV curves are usable for describing crop water status including crop water storage.     

Comparative resistance of the soil and the plant to water transport

The resistances to liquid water transport in the soil and plant were determined directly and simultaneously from measurements of soil, root, and leaf water potentials and the flux of water through the soil-plant system to the sites of evaporation in the leaf. For soybean (Merr.) transporting water at a steady rate, water potential differences between soil and root were smaller than between root and leaf over the range of soil water potentials from −0.2 to −11 bars. As soil water was depleted, water flow through the soil and plant decreased to one-tenth the maximum rate, but both the soil resistance and plant resistance increased. The plant resistance remained larger than the soil resistance over the entire range of soil water availability. Previous suggestions that the soil is the major resistance have ignored the increase in plant resistance and/or assumed root densities that were too low.     

Hydraulic and chemical signalling in the control of stomatal conductance and transpiration.

Abscisic acid (ABA) transported in the xylem from root to shoot and perceived at the guard cell is now widely studied as an essential regulating factor in stomatal closure under drought stress. This provides the plant with a stomatal response mechanism in which water potential is perceived in the root as an indication of soil water status and available water resources. There is also ample evidence that stomata respond directly to some component of leaf water status. This provides additional information about water potential gradients developing between root and shoot as the result of water transport, allowing for a more stable regulation of shoot water status and better protection of the transport system itself. The precise location at which leaf water status is sensed, however, and the molecular events transducing this signal into a guard cell response are not yet known. Major questions therefore remain unanswered on how water stress signals perceived at root and leaf locations are integrated at the guard cell to control stomatal behaviour.     

Influence of soil water deficits on root growth of cotton seedlings

Summary Cotton (Gossypium hirsutum L. cv. H14) seedlings were raised in soil of differing soil water content in specially designed pots in which the roots had access to freely available water and nutrients located 2.5 cm below the base of the soil core. The time for root emergence from the soil core and the rate of root growth were measured daily from sowing to harvest. The root and shoot dry weight and leaf water potential were measured at the final harvest 16 days after sowing. As soil water content decreased, the root emerged from the soil earlier and the initial rate of root elongation was faster. In spite of the availability of freely available water, the plants in the soil at low water contents had significantly lower leaf water potentials than those in soil at high water contents. The root: shoot ratio increased as the soil water content decreased. This arose from an absolute increase in root weight, with shoot weight not being significantly affected.     

根源信号参与调控气孔行为的机制及其农业节水意义

在土壤干旱情况下,根源信号一方面向植物地上部分的长距离传输,为地上部分提供了土壤水分获取能力的测度,另一方面调控气孔开度,抑制蒸腾作用并提高植物的水分利用效率．文中综述了根源信号参与调控植物水分利用的生理机制和理论模型,指出该模型与根系吸水模型、气孔导度模型耦合,能够更好地反映植物叶片对土壤干旱以及大气干旱的响应、评述了在根源信号参与调控植物水分关系的基础上发展的调亏灌溉(RDI)、部分根系干旱(PRD)和控制性交替灌溉(CAI)等有效灌溉手段,有助于合理配置根系层供水量,通过根土相互作用和信号物质的传输,降低蒸腾和提高水分利用效率、另外,根源信号在调控根系生长发育、延缓地上部分生长以调节根冠比例,优化资源分配以利于生殖生长等方面均有所为,为全面提高农田水分利用效率提供节水生理基础。     

Modelling Water in Crops and Plant Ecosystems

A water submodel is described that is specifically designedfor use with plant growth simulators that represent internalplant substrates and variable shoot:root partitioning. The modelcalculates water flow from soil to root, root to shoot, andshoot to the atmosphere, for a closed-canopy situation. As presentedhere, the model has three state variables: the masses of waterin the soil, root and shoot, and represents the processes ofevapotranspiration, rainfall interception and evaporation fromthe canopy, and drainage. The Penman –Monteith equationis used for crop transpiration. The fluxes of water from soilto root, and root to shoot, are driven by water potential difference.Tissue water potential and its components are calculated fromtissue water content and other plant variables and parameters.The model is able to simulate osmoregulation and describes avariable relationship between tissue water potential, its componentsand relative water content, depending on growth conditions.The model has elsewhere been integrated with two plant ecosystemmodels: for grassland and forest. The specific implementationand simulations given are for the Hurley pasture model (Thornleyand Verberne, 1989), a temperate grass vegetative growth model.The model gives reasonable predictions for diurnal changes inwater potential, drying-down behaviour and other quantitieswithin the scope of the model. Simulation; model; water relations; crop growth; grass     

盐胁迫对大豆根系木质部压力和Na+吸收的影响

取栽培大豆的水培幼苗为材料,用木质部压力探针和原子吸收分光光度计测定了盐胁迫条件下其根木质部压力和伤流液中Na~+含量的变化,以分析大豆抗盐吸水的机制.结果表明:在25～150 mmol/L NaCl的浓度范围内,随着盐胁迫强度的增加,大豆根木质部负压力的绝对值逐渐增大,但相对负压力和根的径向反射系数则逐渐减小;木质部伤流液中Na~+含量逐渐增加,但Na~+的相对含量则逐渐降低.同时,虽然根系吸水所需的木质部负压力(压力势)及根木质部伤流液的渗透势随着盐胁迫强度的增加都有所下降,但两者共同作用使木质部水势下降的幅度远远小于根外溶液水势(渗透势)下降的幅度,即随着根外溶液盐浓度的升高,根木质部溶液的总水势逐渐高出根外溶液的水势.上述结果说明,在盐胁迫下大豆可以利用相对小的木质部负压力逆水势梯度吸水,且通过避免对Na~+的过量吸收来适应盐胁迫环境.     

Soil and plant resistances to water uptake byVicia faba L.

Summary The hydraulic resistivity ofVicia faba L. roots grown in soil was estimated from steady state measurements of transpiration rate and leaf and soil water potentials. Root and stem axial resistivities, estimated from xylem vessel radii, were negligible. Root radial resistivity was estimated to be 1.3×10¹² sm⁻¹. This root radial resistivity value was used to estimate, root resistance to water uptake for a field crop ofVicia faba. Previously published results were used for root distribution and soil water contents at the drained upper limit (DUL) and the lower limit (LL) of extractable soil water. Soil resistance to water uptake was estimated from single root theory using the steady rate solution. At the DUL, root resistance was about 10⁵ times greater than soil resistance. At the LL, soil resistance exceeded root resistance for depths less than 0.3 m, but for depths greater than this soil resistance was smaller than root resistance. Estimates of possible uptake rates at given leaf water potentials indicated that overall soil resistance had a negligible influence upon uptake, even at the LL. The reliability of this result is examined in detail. It is concluded that over the complete range of extractable soil water contents soil resistanceper se would not have limited water use by this crop. This conclusion may also be valid for a wide range of soil and crop combinations.     

Leaf and root control of stomatal closure during drying in soybean

The stomatal conductance of an illuminated 2.5 cm² area of an intact soybean leaflet was the same whether the rest of the shoot was in light or darkness. This was true throughout soil drying cycles. Water potential of tissue immediately outside the illuminated area consistently decreased about 0.3 MPa upon illumination of the shoot. This erroneously suggested that stomatal conductance during soil drying did not respond to diurnal reductions in leaf water potential, but was controlled by root or soil water status. Tests showed that the water potential of tissue in the illuminated area did not change in the steady-state upon illumination of the rest of the shoot. Water potentials of shaded sections of leaves were not different from predawn water potentials, and were higher than leaf xylem pressure potentials as determined with a pressure chamber. These steep local gradients of leaf water potential suggest that there is minimal interchange of water among xylem elements leading from roots to different sections of leaves. The relationship between stomatal conductance and leaf water potential was the same whether leaf water potential was reduced by soil drying, application of polyethylene glycol (PEG) to the root system, lowering root temperature, or leaf excision. In the root cooling experiment, there was no soil drying, and with leaf excision, there was no root drying. The similarity of stomatal responses to leaf water potential in all cases strongly suggests control of conductance by a signal produced by local leaf water potential rather than root or soil water status in these experiments.     

Measurement of water fluxes and potentials in a single root-soil system

Summary The usefulness of a tensiometer-potometer system in investigations of water flow in the, vicinity of a plant root has been demonstrated. Measurements were made of the root-soil interface water potential, xylem potential and the distribution of water fluxes and root resistance along the length of a maize root. For a root growing in sand, the rhizosphere resistance was 3.5 to 8 times the radial resistance of the root at average rhizosphere, potentials of –250 m bars. For a root growing in sandy loam such rhizosphere resistance was not achieved until the average rhizosphere potential is approximately –2 bars.Contribution from the Department of Soil Science and Plant Nutrition, University of Western Australia, Nedlands, Western Australia 6009.     

Phenotypic plasticity and water flux rates of Citrus root orders under salinity

Knowledge about the root system structure and the uptake efficiency of root orders is critical to understand the adaptive plasticity of plants towards salt stress. Thus, this study describes the phenological and physiological plasticity of Citrus volkameriana rootstocks under severe NaCl stress on the level of root orders. Phenotypic root traits known to influence uptake processes, for example frequency of root orders, specific root area, cortical thickness, and xylem traits, did not change homogeneously throughout the root system, but changes after 6 months under 90 mM NaCl stress were root order specific. Chloride accumulation significantly increased with decreasing root order, and the Cl(-) concentration in lower root orders exceeded those in leaves. Water flux densities of first-order roots decreased to <20% under salinity and did not recover after stress release. The water flux densities of higher root orders changed marginally under salinity and increased 2- to 6-fold in second and third root orders after short-term stress release. Changes in root order frequency, morphology, and anatomy indicate rapid and major modification of C. volkameriana root systems under salt stress. Reduced water uptake under salinity was related to changes of water flux densities among root orders and to reduced root surface areas. The importance of root orders for water uptake changed under salinity from root tips towards higher root orders. The root order-specific changes reflect differences in vulnerability (indicated by the salt accumulation) and ontogenetic status, and point to functional differences among root orders under high salinity.     

Effect of water stress on leaf and root growth,and water uptake ofGmelina arborea ROXB. seedlings

Young seedlings ofGmelina arborea Roxb. were subjected to 2 weeks of drought. Despite the gradual reduction in stomatal conductance, leaf and root growth was not affected until the later part of the stress period. This was attributed to solute adjustment in the roots of the plants. As the severity of water stress increased, root growth was prolific in all the soil segments. As a result, water in the lowest soil segment was used to maintain plant turgor, which in turn sustains the leaf and root growth during the water-stress treatment. The influence of soil water content and soil water potential upon soil water uptake rate was also evaluated on soil profile basis. Rates of extraction began to decline in all soil segments as soon as soil water potential fell below -0.06 MPa, presumably as a result of vapour gaps between the root and soil (root: soil interface resistance). It is suggested that the growth of roots ofGmelina plants away from drying soil will minimize the resistance to water uptake.