Root elongation, water stress, and mechanical impedance: a review of limiting stresses and beneficial root tip traits

Root elongation in drying soil is generally limited by a combination of mechanical impedance and water stress. Relationships between root elongation rate, water stress (matric potential), and mechanical impedance (penetration resistance) are reviewed, detailing the interactions between these closely related stresses. Root elongation is typically halved in repacked soils with penetrometer resistances >0.8-2?MPa, in the absence of water stress. Root elongation is halved by matric potentials drier than about -0.5?MPa in the absence of mechanical impedance. The likelihood of each stress limiting root elongation is discussed in relation to the soil strength characteristics of arable soils. A survey of 19 soils, with textures ranging from loamy sand to silty clay loam, found that ～10% of penetration resistances were >2?MPa at a matric potential of -10?kPa, rising to nearly 50% >2?MPa at - 200?kPa. This suggests that mechanical impedance is often a major limitation to root elongation in these soils even under moderately wet conditions, and is important to consider in breeding programmes for drought-resistant crops. Root tip traits that may improve root penetration are considered with respect to overcoming the external (soil) and internal (cell wall) pressures resisting elongation. The potential role of root hairs in mechanically anchoring root tips is considered theoretically, and is judged particularly relevant to roots growing in biopores or from a loose seed bed into a compacted layer of soil.     

Root elongation of seedling peas through layered soil of different penetration resistances

Field soils contain localized zones of larger penetration resistance within peds and compacted layers, while cracks and biopores offer low resistance pathways to roots. Root responses to such localized conditions have not been investigated in detail. This study examined what happens to the root elongation rate when roots grew through a layer of hard soil into a layer of looser soil for a 4 day period. The experiment was performed twice; firstly with the shoot in continuous darkness, and secondly with it exposed to a day-night cycle to prevent etiolation of the shoot. Pea seedlings were grown in columns of a sandy loam soil which was packed to bulk densities of 0.85, 1.1, 1.3 or 1.4 Mg/m³ in the top layer and 0.85 Mg/m³ in the bottom layer. The root elongation rate in the top layer of 1.4 Mg/m³ soil (penetrometer resistance=1.8 MPa) was only 55% of the elongation rate in the top layer of 0.85 Mg/m³ soil (penetrometer resistance=0.06 MPa). The elongation rate of roots that had grown through the top layer of 1.4 Mg/m³ soil into the bottom layer of loose soil was reduced by some residual effect of the mechanical impedance. The root elongation rate in the bottom layer of loose soil decreased as the penetrometer resistance of the top layer of soil increased. The daily elongation rate of the roots in the bottom layer that had grown through the 1.4 Mg/m³ soil averaged only about 65% of the elongation rate of the roots that had grown through the 0.85 Mg/m³ soil. This residual effect of mechanical impedance on root elongation persisted for at least 2 days and was more severe in the day-night cycle experiment than in the dark experiment. These results have important implications for modelling root elongation in any soil in which the soil strength changes with distance or with time.     

Penetrometer resistance,root penetration resistance and root elongation rate in two sandy loam soils

Root penetration resistance and elongation of maize seedling roots were measured directly in undisturbed cores of two sandy loam soils. Root elongation rate was negatively correlated with root penetration resistance, and was reduced to about 50 to 60% of that of unimpeded controls by a resistance of between 0.26 and 0.47 MPa. Resistance to a 30° semiangle, 1 mm diameter penetrometer was between about 4.5 and 7.5 times greater than the measured root penetration resistance. However, resistance to a 5° semiangle, 1 mm diameter probe was approximately the same as the resistnace to root penetration after subtracting the frictional component of resistance. The diameter of roots grown in the undisturbed cores was greater than that of roots grown in loose soil, probably as a direct result of the larger mechanical impedance in the cores.     

Simultaneous measurement of root force and elongation for seedling pea roots

A new method was developed to measure simultaneously, continuously,and non-destructively the elongation rate and the force exertedby the roots of seedlings grown in moist air. A pea (Pisum sativumL. cv. Helka) seedling was suspended inside a modified sampletube on one side of a pulley, with the tip of the radicle pushingon to a force transducer through a hole in the tube. The forceon the root tip was monitored by the force transducer and couldbe adjusted by adding or removing mass from the counterweighton the other side of the pulley. As the root grew, the sampletube was raised and the elongation of the root was monitoredusing a linear variable differential transformer (LVDT) attachedto the thread connecting the sample tube and counterweight.The changes in elongation rate were recorded which occurredin response to increases and decreases in the applied force.Forces of up to 125 mN were exerted on the root, correspondingto forces per unit final cross-sectional area (i.e. root growthpressures) of up to 0.1 MPa. As soon as the force on the root was changed there was a rapidreversible compression or extension of the root. Superimposedon this elastic/viscoelastic deformation, the root elongationrate slowed by more than 50% within 30 min of increasing theforce applied to the root by 100 mN. A similarly fast but smallerincrease in growth rate occurred when the force was removed.Both of these ‘fast’ responses were followed bya longer period of more gradual change in the root elongationrate over a period of 20 h or longer. Both ‘fast’and ‘slow’ responses may be explained in terms ofa modified Lockhart model of growth. The initial ‘fast’response of the root is probably due to the immediate changein the effective pressure (i.e. the turgor pressure minus theyield stress and external resisting pressure) available to drivecell elongation. The reason for the second slower adjustmentof the elongation rate is not known, but is probably due tosome combination of a decrease in the rate of cell productionand/or a stiffening of the cell walls in the longitudinal directionwith increasing mechanical resistance. The increase in rootdiameter in response to mechanical impedance decreased the rootgrowth pressure that the root exerted, but was associated witha slower root elongation rate. Key words: Compaction, mechanical impedance, penetration resistance, root diameter, soil strength     

A model of water flow through plants incorporating shoot/root 'message' control of stomatal conductance

Abstract. A model of water flow from the soil into the plant, and from the plant to the atmosphere is described. There are three state variables in the model: the soil, root and shoot water contents. The flow rate of water from the soil to the root is calculated by dividing the gradient in water potential by a resistance, comprising the resistance from the bulk soil to the root surface, and that from the root surface to the root interior. The resistance in the soil depends on the soil hydraulic conductivity, which in turn depends on the soil water potential. The flow rate from the root to the shoot is given by the gradient in water potential divided by a resistance, which depends on the structural dry mass of the plant. Transpiration is described by the Penman-Monteith equation. The plant water characteristics can be modified to take account of osmotic and cell wall rigidity parameters. The model incorporates the concept of shoot/root ‘messages’ of water stress, which influence stomatal conductance. The message works through the generation of a hormone as the pressure potential in the shoot (mesophyll) or root falls. This hormone induces a shift of osmoticum from the guard cells to the surrounding mesophyll cells, which causes an increase (i.e. closer to zero) in the osmotic potential in these cells. This, in turn, causes a decrease in their pressure potential, and so reduces stomatal conductance. The model is used as a framework to address some of the issues that have recently been raised concerning the role of water potential in describing water flow through plants. We conclude that, with the hormone present, there is unlikely to be a unique relationship between stomatal conductance and shoot total water potential, since stomatal conductance depends on the pressure potential in the guard cells, which may differ from that in other cells. Nevertheless, this does not imply that water potential is not an important, and indeed fundamental, component for describing water flow through plants. Other aspects of water flow through plants are also considered, such as diurnal patterns of shoot, root and soil water potential components. It is seen that these may differ from the commonly held view that, as the soil dries down, they all attain the same values during the dark period, and which, as we show, is largely unsubstantiated either theoretically or experimentally.     

TRAP: a modelling approach to below-ground carbon allocation in temperate forests

Tree root systems, which play a major role in below-ground carbon (C) dynamics, are one of the key research areas for estimating long-term C cycling in forest ecosystems. In addition to regulating major C fluxes in the present conditions, tree root systems potentially hold numerous controls over forest responses to a changing environment. The predominant contribution of tree root systems to below-ground C dynamics has been given little emphasis in forest models. We developed the TRAP model, i.e. Tree Root Allocation of Photosynthates, to predict the partitioning of photosynthates between the fine and coarse root systems of trees among series of soil layers. TRAP simulates root system responses to soil stress factors affecting root growth. Validation data were obtained from two Belgian experimental forests, one mostly composed of beech (Fagus sylvatica L.) and the other of Scots pine (Pinus sylvestris L.). TRAP accurately predicted (R = 0.88) night-time CO₂ fluxes from the beech forest for a 3-year period. Total fine root biomass of beech was predicted within 6% of measured values, and simulation of fine root distribution among soil layers was accurate. Our simulations suggest that increased soil resistance to root penetration due to reduced soil water content during summer droughts is the major mechanism affecting the distribution of root growth among soil layers of temperate Belgian forests. The simulated annual rate of C input to soil litter due to the fine root turnover of the Scots pine was 207 g C m^–2 yr^–1. The TRAP model predicts that fine root turnover is the single most important source of C to the temperate forest soils of Belgium.     

Root cap removal increases root penetration resistance in maize (Zea mays L)

The root cap assists the passage of the root through soil by means of its slimy mucilage secretion and by the sloughing of its outer cells. The root penetration resistance of decapped primary roots of maize (Zea mays L. cv. Mephisto) was compared with that of intact roots in loose (dry bulk density 1.0 g cm-3; penetration resistance 0.06 MPa) and compact soil (1.4 g cm-3; penetration resistance 1.0 MPa), to evaluate the contribution of the cap to decreasing the impedance to root growth. Root elongation rate and diameter were the same for decapped and intact roots when the plants were grown in loose soil. In compacted soil, however, the elongation rate of decapped roots was only about half that of intact roots, whilst the diameter was 30% larger. Root penetration resistances of intact and decapped seminal axis were 0.31 and 0.52 MPa, respectively, when the roots were grown in compacted soil. These results indicated that the presence of a root cap alleviates much of the mechanical impedance to root penetration, and enables roots to grow faster in compacted soils.     

Root elongation rate is correlated with the length of the bare root apex of maize and lupin roots despite contrasting responses of root growth to compact and dry soils

Background

We investigated interacting effects of matric potential and soil strength on root elongation of maize and lupin, and relations between root elongation rates and the length of bare (hairless) root apex.

Methods

Root elongation rates and the length of bare root apex were determined for maize and lupin seedlings in sandy loam soil of various matric potentials (?0.01 to ?1.6 MPa) and bulk densities (0.9 to 1.5 Mg m^?3).

Results

Root elongation rates slowed with both decreasing matric potential and increasing penetrometer resistance. Root elongation of maize slowed to 10 % of the unimpeded rate when penetrometer resistance increased to 2 MPa, whereas lupin elongated at about 40 % of the unimpeded rate. Maize root elongation rate was more sensitive to changes in matric potential in loosely packed soil (penetrometer resistances <1 MPa) than lupin. Despite these differing responses, root elongation rate of both species was linearly correlated with length of the bare root apex (r² 0.69 to 0.97).

Conclusion

Maize root elongation was more sensitive to changes in matric potential and mechanical impedance than lupin. Robust linear relationships between elongation rate and length of bare apex suggest good potential for estimating root elongation rates for excavated roots.     

水分胁迫与光照条件对棉花干物质和产量形成影响的数学模型

从作物冠层净同化速率入手,通过引入对CO2浓度、空气湿度、光照强度和土壤含水量反映较敏感的光能利用系数（β）,建立了考虑水分胁迫和光照条件对作物干物质积累与产量形成影响的数学模型,模型考虑了水分胁迫与低光照下冠层阻力增加的设定,将反映作物冠层水分状况的功能叶水势（Ψl）作为参数纳入本模型,通过对土壤相对含水量（Aw)、气温（Ta）、水汽压差（VPD）的多元回归估算出Ψl,并将空气动力学阻力（Ra）简化为风速（u）的函数,盆载试验应用实例和敏感性分析表明,该模型在诊断环境因子特别是土壤水分与光照因子对作物生长和产量构成的影响具有一定的实用性。     

水分胁迫下AM真菌对沙打旺生长和抗旱性的影响

利用盆栽试验研究了水分胁迫条件下接种AM真菌对优良牧草和固沙植物沙打旺(Astragalus adsurgens Pall.)生长和抗旱性的影响。在土壤相对含水量为70%、50%和30%条件下,分别接种摩西球囊霉(Glomus mosseae)和沙打旺根际土著菌,不接种处理作为对照。结果表明,水分胁迫显著降低了沙打旺植株(无论接种AM真菌与否)的株高、分枝数、地上部干重和地下部干重,并显著提高了土著AM真菌的侵染率,对摩西球囊霉的侵染率无显著影响。接种AM真菌可以促进沙打旺生长和提高植株抗旱性,但促进效应因土壤含水量和菌种不同而存在差异。不同水分条件下,接种AM真菌显著提高了植株菌根侵染率、根系活力、地下部全N含量和叶片CAT活性。土壤相对含水量为30%和50%时,接种株地上部全N、叶片叶绿素、可溶性蛋白、脯氨酸含量和POD活性显著高于未接种株;接种AM真菌显著降低了叶片MDA含量;接种土著AM真菌的植株株高、分枝数、地上部和地下部干重显著高于未接种株。土壤相对含水量为30%时,接种AM真菌显著增加了地上部全P含量和叶片相对含水量;接种摩西球囊霉的植株株高、分枝数、地上部和地下部干重显著高于未接种株。水分胁迫40d,接种AM真菌显著提高了叶片可溶性糖含量。水分胁迫80d,接种株叶片SOD活性显著增加。菌根依赖性随水分胁迫程度增加而提高。沙打旺根际土著菌接种效果优于摩西球囊霉。水分胁迫和AM真菌的交互作用对分枝数、菌根侵染率、叶片SOD、CAT和POD活性、叶绿素、脯氨酸、可溶性蛋白、地上部全N和全P、地下部全N和根系活力有极显著影响,对叶片丙二醛和地下部全P有显著影响。AM真菌促进根系对土壤水分和矿质营养的吸收,改善植物生理代谢活动,从而提高沙打旺抗旱性,促进其生长。试验结果为筛选优良抗旱菌种,充分利用AM真菌资源促进荒漠植物生长和植被恢复提供了依据。     

Root responses to soil physical conditions; growth dynamics from field to cell

Root growth in the field is often slowed by a combination of soil physical stresses, including mechanical impedance, water stress, and oxygen deficiency. The stresses operating may vary continually, depending on the location of the root in the soil profile, the prevailing soil water conditions, and the degree to which the soil has been compacted. The dynamics of root growth responses are considered in this paper, together with the cellular responses that underlie them. Certain root responses facilitate elongation in hard soil, for example, increased sloughing of border cells and exudation from the root cap decreases friction; and thickening of the root relieves stress in front of the root apex and decreases buckling. Whole root systems may also grow preferentially in loose versus dense soil, but this response depends on genotype and the spatial arrangement of loose and compact soil with respect to the main root axes. Decreased root elongation is often accompanied by a decrease in both cell flux and axial cell extension, and recent computer-based models are increasing our understanding of these processes. In the case of mechanical impedance, large changes in cell shape occur, giving rise to shorter fatter cells. There is still uncertainty about many aspects of this response, including the changes in cell walls that control axial versus radial extension, and the degree to which the epidermis, cortex, and stele control root elongation. Optical flow techniques enable tracking of root surfaces with time to yield estimates of two-dimensional velocity fields. It is demonstrated that these techniques can be applied successfully to time-lapse sequences of confocal microscope images of living roots, in order to determine velocity fields and strain rates of groups of cells. In combination with new molecular approaches this provides a promising way of investigating and modelling the mechanisms controlling growth perturbations in response to environmental stresses.     

Analysis of a continuum model for root growth

We give an analysis by asymptotic expansions of a model for root growth. The model assumes growth is due entirely to mechanical forces created by the flux of water from the surrounding soil. We attempt to suggest ways in which the model could account for experimentally observed patterns of root growth. The model predicts that the rate of growth of the root is independent of the extensibility of the root wall.     

Effect of soil mechanical impedance on root growth of two rice varieties under field drought stress

Two upland rice varieties, Azucena and Bala, were screened for root growth under droughted and irrigated treatments in two field sites at the West Africa Rice Development Association (WARDA) experimental farm, Côte d’Ivoire, during the dry season of 1999/2000. The sites were chosen to represent contrasting soil profile penetration resistance (PR) characteristics on upland sites, although both were relatively impeding. The number of nodal root axes per unit area passing through horizontal transects (root density) was counted at 35, 56, 77 and 98 days after sowing (DAS) at 10 cm depth intervals. Azucena consistently maintained a greater root density than Bala and a greater proportion of Azucena roots grew to 30 cm depth (22.7% vs. 8.4% at 77 DAS). There was little detectable effect of water regime on root distribution but evidence of lower root numbers at depths below 20 cm in the higher PR site was revealed. A site by variety by soil depth interaction suggests that Azucena roots are more strongly affected by very high PR than those of Bala. PR between 0–30 cm depth increased greatly with decreasing soil water content during the drought as the soil dried. This increase is likely to have prevented or greatly impaired further nodal root growth within this layer. At 40 cm depth, PR was high (3–4 MPa) but did not increase during the drought. At this depth root growth rate was likely to be greatly reduced despite the availability of water. These results demonstrate that varietal differences in root morphology characterised in the laboratory can be also detected in impeding field soils as differences in the density of roots at depth. Relatively poor root growth in these fields in the absence of drought was probably due to the high mechanical impedance and/or the physiological stress of the plants in the dry season. Our results indicate that high mechanical impedance was a more fundamental constraint on root growth than soil water availability during the drought. Thus, varietal differences in root penetration ability might be very important for drought avoidance in soils of this type.     

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.     

Influence of temperature and water potential on root growth of white oak

Root growth of white oak ( Quercus alba L.) was observed under field conditions using a rhizotron. The effects of temperature, soil water potential, and leaf water potential were evaluated on three measures of root growth and development: root elongation rate, number of growing roots, and root growth intensity (sum of projected root area compared to the total root viewing area). Root elongation rate was linearly related to changes in soil temperature and soil water potential. At soil temperatures less than 17deg;C, temperature was the dominant factor affecting rate of growth, bat at temperatures greater than 17°C soil water potential became the important factor. Unlike root elongation rate, the number of growing roots and root growth intensity increased at cold soil temperatures (8°C) and at soil water potentials of-0.3 to -0.8 MPa. At high soil water potentials (-0.1 MPa) root elongation rate reached a maximum while the number of growing roots and root growth intensity were low. These differences showed that root growth and development were not exclusively affected by the soil environment. In addition, the relationship between root growth and predawn leaf water potential suggested that root growth was a contributing factor to the drought resistance of white oak.     

The effect of mechanical impedance on root growth in pea (Pisum sativum). I. Rates of cell flux, mitosis, and strain during recovery

We studied the effect of mechanical impedance on cell flux and meristematic activity in pea roots. Pea seedlings ( Pisum sativum L. cv. Helka) were grown in cores of sand packed to dry bulk densities of either; 1.4 Mg m⁻³ with an additional 2.4 kg uniaxial load applied to the surface to increase the mechanical resistance to growth (penetration resistance of 1.5 MPa); or 1.0 Mg m⁻³ (penetration resistance of 0.05 MPa). A water content of 0.06 g g⁻¹ was chosen for optimum root growth. After 3 days, the seedlings were transferred to hydroponics, colchicine was added and the rate of cell doubling, mitotic index and length of the cell cycle was assessed. Cell flux in the third cortical layer was calculated for roots immediately removed from sand.Mechanical impedance slowed root extension to about 20% of the unimpeded rate, and final cell length was reduced to 50% of the unimpeded length. The rate of cell doubling was 3.4 times slower for roots recovering from mechanical impedance mostly as a result of a longer period spent in interphase. Cell flux in impeded roots was approximately half that of unimpeded roots (5 cells h⁻¹), and contributed to a shorter cell file and elongation zone, and a slower rate of root elongation.     

Field evaluation of laboratory techniques for predicting the ability of roots to penetrate strong soil and of the influence of roots on water sorptivity

The ability of two laboratory screening techniques to predict the abilities of roots of eight crop species to penetrate a compacted soil were evaluated and compared in a field experiment. A soil tilled to remove the effects of mechanical resistance was planted with the same species to serve as a control. Depth of root penetration, root density and the influence of the roots on the sorptivity of water were measured.Roots of all species penetrated deeper in the deep tilled than compacted soils. There were differences in the ability of roots of the species to penetrate the compacted soil. Generally dicotyledonous species had more roots penetrating to depth in both the compact and deep tilled soils. Within the main species classifications, lupin and safflower (dicotyledons) and oats and barley (monocotyledons) had the highest penetration into the compacted soil.Water sorptivities in the deep tilled soils were higher than those of the compact soil. Soil from planted treatments had higher sorptivities than soil which had not been planted. This is attributed to biopores left by the roots. Sorptivities of soils which had dicotyledonous species were generally higher than those of monocotyledons. The soil planted with safflower produced the highest sorptivity in the compacted layer (0.1–0.3 m).A comparison of the accuracy of the two laboratory screening methods in predicting the field penetration of roots suggest that the method involving mechanical stress was better than that involving osmotic stress. Relative root diameter was found to be a better indicator of the penetration ability of roots than relative root elongation.     

A kinematic model of stretch-induced stress fiber turnover and reorientation

A kinetic model based on constrained mixture theory was developed to describe the reorganization of actin stress fibers in adherent cells in response to diverse patterns of mechanical stretch. The model was based on reports that stress fibers are pre-extended at a “homeostatic” level under normal, non-perturbed conditions, and that perturbations in stress fiber length destabilize stress fibers. In response to a step change in matrix stretch, the model predicts that stress fibers are initially stretched in registry with the matrix, but that these overly stretched fibers are gradually replaced by new fibers assembled with the homeostatic level of stretch in the new configuration of the matrix. In contrast, average fiber stretch is chronically perturbed from the homeostatic level when the cells are subjected to cyclic equibiaxial stretch. The model was able to describe experimentally measured time courses of stress fiber reorientation perpendicular to the direction of cyclic uniaxial stretch, as well as the lack of alignment in response to equibiaxial stretch. The model also accurately described the relationship between stretch magnitude and the extent of stress fiber alignment in endothelial cells subjected to cyclic uniaxial stretch. Further, in the case of cyclic simple elongation with transverse matrix contraction, stress fibers orient in the direction of least perturbation in stretch. In summary, the model predicts that the rate of stretch-induced stress fiber disassembly determines the rate of alignment, and that stress fibers tend to orient toward the direction of minimum matrix stretch where the rate of stress fiber turnover is a minimum.     

冬小麦近轴和远轴叶面气孔对土壤水分胁迫反应的敏感性

当根层土壤水分含量不足,作物体内出现水分胁迫时,小麦叶片两面气孔的反应有明显差异。远轴叶面气孔对水分胁迫的反应比近轴叶面气孔敏感。当出现水分胁迫时,远轴叶面气孔首先收缩,且收缩的程度比近轴叶面气孔大。远轴与近轴叶面气孔阻力的比值(r_(ab)/r_(ab))与根层平均土壤水势(Ψ_s)有关,当Ψ_s大于-50 kPa时,r_(ab)/r_(ad)基本稳定在1.5左右,而当Ψ_s小于-50 kPa时,r_(ab )/r_(ab)随Ψ_s降低而明显增加。     

Growth response of barley and tomato to nitrogen stress and its control by abscisic acid,water relations and photosynthesis

Barley (Hordeum vulgare L.) and tomato Lycopersicon esculentum Mill.) were grown hydroponically and examined 2, 5, and 10 d after being deprived of nitrogen (N) supply. Leaf elongation rate declined in both species in response to N stress before there was any reduction in rate of dryweight accumulation. Changes in water transport to the shoot could not explain reduced leaf elongation in tomato because leaf water content and water potential were unaffected by N stress at the time leaf elongation began to decline. Tomato maintained its shoot water status in N-stressed plants, despite reduced water absorption per gram root, because the decline in root hydraulic conductance with N stress was matched by a decline in stomatal conductance. In barley the decline in leaf elongation coincided with a small (8%) decline in water content per unit area of young leaves; this decline occurred because root hydraulic conductance was reduced more strongly by N stress than was stomatal conductance. Nitrogen stress caused a rapid decline in tissue NO ₃ ^- pools and in NO ₃ ^- flux to the xylem, particularly in tomato which had smaller tissue NO ₃ ^- reserves. Even in barley, tissue NO ₃ ^- reserves were too small and were mobilized too slowly (60% in 2 d) to support maximal growth for more than a few hours. Organic N mobilized from old leaves provided an additional N source to support continued growth of N-stressed plants. Abscisic acid (ABA) levels increased in leaves of both species within 2 d in response to N stress. Addition of ABA to roots caused an increase in volume of xylem exudate but had no effect upon NO ₃ ^- flux to the xylem. After leaf-elongation rate had been reduced by N stress, photosynthesis declined in both barley and tomato. This decline was associated with increased leaf ABA content, reduced stomatal conductance and a decrease in organic N content. We suggest that N stress reduces growth by several mechanisms operating on different time scales: (1) increased leaf ABA content causing reduced cell-wall extensibility and leaf elongation and (2) a more gradual decline in photosynthesis caused by ABA-induced stomatal closure and by a decrease in leaf organic N.Abbreviation and symbols ABA abscisic acid - c_i leaf internal CO₂ concentration - L_p root hydraulic conductance