Water uptake and structural plasticity along roots of a desert succulent during prolonged drought

Desert succulents resume substantial water uptake within 1–2 d of the cessation of drought, but the changes in root structure and hydraulic conductivity underlying such recovery are largely unknown. In the monocotyledonous leaf succulent Agave deserti Engelm. substantial root mortality occurred only for lateral roots near the soil surface; nearly all main roots were alive at 180 d of drought. New main roots were initiated and grew up to 320 mm at soil water potentials lower than – 5·0 MPa, utilizing water from the shoot. The hydraulic conductivity of distal root regions decreased 62% by 45 d of drought and 70% thereafter. After 7 d of rewetting, root hydraulic conductivity was restored following 45 d of drought but not after 90 and 180 d. The production of new lateral roots and the renewed apical elongation of main roots occurred 7–11 d after rewetting following 180 d of drought. Hydraulic conductivity was higher in the distal region than at midroot and often increased again near the root base, where many endodermal cells lacked suberin lamellae. Suberization and xylem maturation were influenced by the availability of moisture, suggesting that developmental plasticity along a root allows A. deserti to capitalize on intermittent or heterogeneous supplies of water.     

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.     

Hydraulic Conductances of the Soil, the Root-Soil Air Gap, and the Root: Changes for Desert Succulents in Drying Soil

Water movement to and from a root depends on the soil hydraulicconductivity coefficient (L^soil), the distance across any root-soilair gap, and the hydraulic conductivity coefficient of the root(L^P). After analytical equations for the effective conductanceof each part of the pathway are developed, the influences ofsoil drying on the soil water potential and L^soil are describedduring a 30 d period for a loamy sand in the field. The influencesof soil drying on L^P for three desert succulents, Agave deserti,Ferocactus acanthodes, and Opuntia ficus-indica, are also describedfor a 30 d period. To quantify the effects of soil drying onthe development of a root-soil air gap, diameters of 6-week-oldroots of the three species were determined at constant watervapour potentials of –1.0 MPa and –10 MPa as wellas with the water vapour potential decreasing at the same rateas soil drying during a 30 d period. The shrinkage observedfor roots initially 2·0 mm in diameter averaged 19% duringthe 30d period. The predominant limiting factor for water movementwas L^P of the root for the first 7 d of soil drying, the root-soilair gap for the next 13 d, and L^soil thereafter. Compared withthe ease of water uptake from a wet soil, the decrease in conductancesduring soil drying, especially the decrease in L^soil causedthe overall conductance to decrease by 3 x 10³-fold during the30 d period for the three species considered, so relativelylittle water was lost to the dry soil. Such rectifier-like behaviourof water movement in the soil-root system resulted primarilyfrom changes in L^soil and, presumably, is a general phenomenonamong plants, preventing water loss during drought but facilitatingwater uptake after rainfall. Key words: Agave deserti, Ferocactus acanthodes, Opuntia ficus-indica, rectification, soil water potential, water movement     

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     

Thermal and Water Relations of Roots of Desert Succulents

Two succulent perennials from the Sonoran Desert, Agave desertiEngelm. and Ferocactus acanthodes (Lem.) Britton and Rose, loselittle water through their roots during drought, yet respondrapidly to light rainfall. Their roots tend to be shallow, althoughabsent from the upper 20 mm or so of the soil. During 12–15d after a rainfall, new root production increased total rootlength by 47 per cent to 740 m for A. deserti and by 27 percent to 230 m for F. acanthodes; root dry weight then averagedonly 15 per cent of shoot dry weight. The annual carbon allocatedto dry weight of new roots required 11 per cent of shoot carbondioxide uptake for A. deserti and 19 per cent for F. acanthodes.Elongation of new roots was greatest near a soil temperatureof 30°C, and lethal temperature extremes (causing a 50 percent decrease in root parenchyma cells taking up stain) were56°C and -7°C. Soil temperatures annually exceeded themeasured tolerance to high temperature at depths less than 20mm, probably explaining the lack of roots in this zone. Attached roots immersed in solutions with osmotic potentialsabove -2·6 MPa could produce new lateral roots, with50 per cent of maximum elongation occurring near -1·4MPa for both species. Non-droughted roots lost water when immersedin solutions with osmotic potentials below -0·8 MPa,and root hydraulic conductance decreased markedly below about-1·2 MPa. Pressure-volume curves indicated that, fora given change in water potential, non-droughted roots lostthree to five times more water than droughted roots, non-droughtedleaves, or non-droughted stems. Hence, such roots, which couldbe produced in response to a rainfall, will lose the most tissuewater with the onset of drought, the resulting shrinkage beingaccompanied by reduced root hydraulic conductance, less contactwith drying soil, and less water loss from the plant to thesoil. Agave deserti, Ferocactus acanthodes, roots, soil, temperature, water stress, drought, Crassulacean acid metabolism, succulents     

Hydraulic properties of rice and the response of gas exchange to water stress

We investigated the role of xylem cavitation, plant hydraulic conductance, and root pressure in the response of rice (Oryza sativa) gas exchange to water stress. In the field (Philippines), the percentage loss of xylem conductivity (PLC) from cavitation exceeded 60% in leaves even in watered controls. The PLC versus leaf water potential relationship indicated diurnal refilling of cavitated xylem. The leaf water potential causing 50 PLC (P(50)) was -1.6 MPa and did not differ between upland versus lowland rice varieties. Greenhouse-grown varieties (Utah) were more resistant to cavitation with a 50 PLC of -1.9 MPa but also showed no difference between varieties. Six-day droughts caused concomitant reductions in leaf-specific photosynthetic rate, leaf diffusive conductance, and soil-leaf hydraulic conductance that were associated with cavitation-inducing water potentials and the disappearance of nightly root pressure. The return of root pressure after drought was associated with the complete recovery of leaf diffusive conductance, leaf-specific photosynthetic rate, and soil-leaf hydraulic conductance. Root pressure after the 6-d drought (61.2 +/- 8.8 kPa) was stimulated 7-fold compared with well-watered plants before drought (8.5 +/- 3.8 kPa). The results indicate: (a) that xylem cavitation plays a major role in the reduction of plant hydraulic conductance during drought, and (b) that rice can readily reverse cavitation, possibly aided by nocturnal root pressure.     

Transport constraints on water use by the Great Basin shrub, Artemisia tridentata

As soil and plant water status decline, decreases in hydraulic conductance can limit a plant's ability to maintain gas exchange. We investigated hydraulic limitations for Artemisia tridentata during summer drought. Water use was quantified by measurements of soil and plant water potential ( Ψ ), transpiration and leaf area. Hydraulic transport capacity was quantified by vulnerability to water stress-induced cavitation for root and stem xylem, and moisture release characteristics for soil. These data were used to predict the maximum possible steady-state transpiration rate ( E _crit) and minimum leaf xylem pressure ( Ψ _crit). Transpiration and leaf area declined by ~ 80 and 50%, respectively, as soil Ψ decreased to –2·6 MPa during drought. Leaf-specific hydraulic conductance also decreased by 70%, with most of the decline predicted in the rhizosphere and root system. Root conductance was projected to be the most limiting, decreasing to zero to cause hydraulic failure if E _crit was exceeded. The basis for this prediction was that roots were more vulnerable to xylem cavitation than stems (99% cavitation at –4·0 versus –7·8 MPa, respectively). The decline in water use during drought was necessary to maintain E and Ψ within the limits defined by E _crit and Ψ _crit.     

Drought-Induced Xylem Dysfunction in Petioles,Branches, and Roots of Populus balsamifera L. and Alnus glutinosa (L.) Gaertn

Variation in vulnerability to xylem cavitation was measured within individual organs of Populus balsamifera L. and Alnus glutinosa (L.) Gaertn. Cavitation was quantified by three different techniques: (a) measuring acoustic emissions, (b) measuring loss of hydraulic conductance while air-dehydrating a branch, and (c) measuring loss of hydraulic conductance as a function of positive air pressure injected into the xylem. All of these techniques gave similar results. In Populus, petioles were more resistant than branches, and branches were more resistant than roots. This corresponded to the pattern of vessel width: maximum vessel diameter in 1- to 2-year-old roots was 140 [mu]m, compared to 65 and 45 [mu]m in rapidly growing 1-year-old shoots and petioles, respectively. Cavitation in Populus petioles started at a threshold water potential of -1.1 MPa. The lowest leaf water potential observed was -0.9 MPa. In Alnus, there was no relationship between vessel diameter and the cavitation response of a plant organ. Although conduits were narrower in petioles than in branches, petioles were more vulnerable to cavitation. Cavitation in petioles was detected when water potential fell below -1.2 MPa. This value equaled midday leaf water potential in late June. As in Populus, roots were the most vulnerable organ. The significance of different cavitation thresholds in individual plant organs is discussed.     

Water stress induced cavitation and embolism in some woody plants

A comparison was made of the relative vulnerability of xylem conduits to cavitation and embolism in three species [ Thuja occidentalis L., Tsuga canadensis (L.) Carr. and Acer saccharum Marsh.]. Waterlogged samples of wood were air dehydrated while measuring relative water loss, loss of hydraulic conductance, cumulative acoustic emissions (= cavitations) and xylem water potential. Most cavitation events and loss of hydraulic conductance occurred while water potential declined from – 1 to –6 MPa. There were differences in vulnerability between species. Other people have hypothesized that large xylem conduits (e.g. vessels) should be more vulnerable to cavitations than small conduits (e.g. tracheids). Our findings are contrary to this hypothesis. Under water stress, the vessel bearing wood retained water better than tracheid bearing wood. However, within a species large conduits were more prone to cavitation than small conduits.     

Long-term acclimatization of hydraulic properties, xylem conduit size, wall strength and cavitation resistance in Phaseolus vulgaris in response to different environmental effects

Phaseolus vulgaris grown under various environmental conditions was used to assess long-term acclimatization of xylem structural characteristics and hydraulic properties. Conduit diameter tended to be reduced and 'wood' density (of 'woody' stems) increased under low moisture ('dry'), increased soil porosity ('porous soil') and low phosphorus ('low P') treatments. Dry and low P had the largest percentage of small vessels. Dry, low light ('shade') and porous soil treatments decreased P50 (50% loss in conductivity) by 0.15-0.25 MPa (greater cavitation resistance) compared with 'controls'. By contrast, low P increased P50 by 0.30 MPa (less cavitation resistance) compared with porous soil (the control for low P). Changes in cavitation resistance were independent of conduit diameter. By contrast, changes in cavitation resistance were correlated with wood density for the control, dry and porous soil treatments, but did not appear to be a function of wood density for the shade and low P treatments. In a separate experiment comparing control and porous soil plants, stem hydraulic conductivity (kh), specific conductivity (ks), leaf specific conductivity (LSC), total pot water loss, plant biomass and leaf area were all greater for control plants compared to porous soil plants. Porous soil plants, however, demonstrated higher midday stomatal conductance to water vapour (gs), apparently because they experienced proportionally less midday xylem cavitation.     

Intra-specific variation in xylem cavitation in interior live oak (Quercus wislizenii A. DC.).

Xylem cavitation induced by water stress reduces plant hydraulic conductance and can indicate the habitat a species evolved in and its phylogenetic background. Species differ widely in cavitation resistance, but less is known about intra-specific variation. Cavitation resistance was assessed for field-collected adult and sapling size classes from three populations of interior live oak (Quercus wislizenii A. DC.) in California, USA. Root and stem cavitation resistance of two-year old seedlings from a greenhouse experiment was also measured. Cavitation resistance curves were determined by injecting air into the vascular system to induce cavitation and measuring the subsequent decline in hydraulic conductance. Based on the air-seeding hypothesis, the absolute value of the air pressures should be equivalent to the tensions that cause cavitation under dehydrating conditions. Conductance declined exponentially with applied pressure for both roots and stems. Comparisons between populations did not reveal significant differences despite good statistical power. The 50% loss in conductance point occurred between 1.0-1.6 MPa; conductance declined more slowly thereafter. Conductance was 21-30% of maximum at 4.0 MPa and 7-14% at 8.0 MPa. Saplings exhibited a nearly identical pattern compared with adults except at 4.0 MPa, where saplings exhibited slightly less cavitation (7%). Greenhouse seedling stems were more resistant compared with both field-collected adults and with seedling roots. The 50% loss in conductance point occurred at 0.83 and 2.6 MPa for seedling roots and stems, respectively. Seedling stems maintained conductance of 20.9% at 8.0 MPa while most roots were fully cavitated between 5.0-8.0 MPa.     

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.     

Diurnal cycles of embolism formation and repair in petioles of grapevine (Vitis vinifera cv. Chasselas)

The impact of water deficit on stomatal conductance (g(s)), petiole hydraulic conductance (K(petiole)), and vulnerability to cavitation (PLC, percentage loss of hydraulic conductivity) in leaf petioles has been observed on field-grown vines (Vitis vinifera L. cv. Chasselas). Petioles were highly vulnerable to cavitation, with a 50% loss of hydraulic conductivity at a stem xylem water potential (Ψ(x)) of -0.95?MPa, and up to 90% loss of conductivity at a Ψ(x) of -1.5?MPa. K(petiole) described a daily cycle, decreasing during the day as water stress and evapotranspiration increased, then rising again in the early evening up to the previous morning's K(petiole) levels. In water-stressed vines, PLC increased sharply during the daytime and reached maximum values (70-90%) in the middle of the afternoon. Embolism repair occurred in petioles from the end of the day through the night. Indeed, PLC decreased in darkness in water-stressed vines. PLC variation in irrigated plants showed the same tendency, but with a smaller amplitude. The Chasselas cultivar appears to develop hydraulic segmentation, in which petiole cavitation plays an important role as a 'hydraulic fuse', thereby limiting leaf transpiration and the propagation of embolism and preserving the integrity of other organs (shoots and roots) during water stress. In the present study, progressive stomatal closure responded to a decrease in K(petiole) and an increase in cavitation events. Almost total closure of stomata (90%) was measured when PLC in petioles reached >90%.     

Shoot dieback during prolonged drought in Ceanothus (Rhamnaceae) chaparral of California: a possible case of hydraulic failure

Progressive diebacks of outer canopy branchlets of Ceanothus crassifolius were repeatedly observed after rainless periods up to 9 mo in duration in the Santa Monica Mountains of southern California. Mean xylem pressures of branchlets near the end of drought were as low as -11.2 MPa (N = 22) with a mean of about 60 dead branchlets per shrub. Inoculation (N = 15) with three species of fungi previously isolated from the same population of C. crassifolius did not promote dieback, suggesting that the observed decline was not fungal induced, as had been proposed. Further, at least 50% of healthy-appearing twigs, without symptoms of dieback, contained isolatible endophytic fungi. We used a centrifugal force method to determine the range of xylem pressure causing cavitation (vulnerability curves) for branchlets (N = 12) and roots (N = 16). We combined vulnerability curves with soil texture data (N = 6) into a water transport model that estimated the critical values (P(Lcrit)) of leaf xylem pressure associated with the loss of water from soil to foliage. Maximum P(Lcrit) was between -10 and -11 MPa and within the range of minimum measured xylem pressures of branchlets during drought and dieback. Branchlet dieback correlated with seasonal declines in xylem pressure in concert with declining safety margins from hydraulic failure. Symptoms of dieback were duplicated in the field by partially severing stem xylem that normally supplied branchlets with water. Taken together, these results indicate that loss of hydraulic conductance to foliage was the probable cause of the observed dieback in C. crassifolius. Partial dieback of peripheral branchlets, and its attendant reduction in evaporative surface area, may be a last-resort mechanism for whole-plant water conservation and drought survival in this species.     

Diurnal and seasonal variation in root xylem embolism in neotropical savanna woody species: impact on stomatal control of plant water status

Vulnerability to water-stress-induced embolism and variation in the degree of native embolism were measured in lateral roots of four co-occurring neotropical savanna tree species. Root embolism varied diurnally and seasonally. Late in the dry season, loss of root xylem conductivity reached 80% in the afternoon when root water potential (psi root) was about -2.6 MPa, and recovered to 25-40% loss of conductivity in the morning when psi root was about -1.0 MPa. Daily variation in psi root decreased, and root xylem vulnerability and capacitance increased with rooting depth. However, all species experienced seasonal minimum psi root close to complete hydraulic failure independent of their rooting depth or resistance to embolism. Predawn psi root was lower than psi soil when psi soil was relatively high (> -0.7 MPa) but became less negative than psi soil, later in the dry season, consistent with a transition from a disequilibrium between plant and soil psi induced by nocturnal transpiration to one induced by hydraulic redistribution of water from deeper soil layers. Shallow longitudinal root incisions external to the xylem prevented reversal of embolism overnight, suggesting that root mechanical integrity was necessary for recovery, consistent with the hypothesis that if embolism is a function of tension, refilling may be a function of internal pressure imbalances. All species shared a common relationship in which maximum daily stomatal conductance declined linearly with increasing afternoon loss of root conductivity over the course of the dry season. Daily embolism and refilling in roots is a common occurrence and thus may be an inherent component of a hydraulic signaling mechanism enabling stomata to maintain the integrity of the hydraulic pipeline in long-lived structures such as stems.     

Vein recovery from embolism occurs under negative pressure in leaves of sunflower (Helianthus annuus)

Leaf veins undergo cavitation at water potentials (Psi(leaf)) commonly experienced by field-growing plants. Theoretically, embolism reversal should not be possible until xylem pressures rise by several kilopascals of atmospheric pressure, but recent evidence suggests that embolized conduits can be refilled even when surrounded by others at substantial tension (novel refilling). The present study reports 'novel refilling' occurring in leaf veins of sunflower (Helianthus annuus L.) while at Psi(leaf) = -0.33 MPa. Sixty per cent loss of vein hydraulic conductance (K(vein)) was recorded at Psi(leaf) < -0.65 MPa, while stem hydraulic conductance (K(stem)) was unaffected even at Psi(leaf) = -1.1 MPa. Loss of K(vein) was accompanied by stomatal closure. Water-stressed plants (Psi(leaf) = -1.1 MPa) were rehydrated overnight to different target water potentials achieved by using PEG at different concentrations as irrigation medium. K(vein) recovered by 50% at Psi(leaf) = -0.47 MPa and vein refilling was complete at Psi(leaf) = -0.33 MPa, i.e. well below the theoretical limit for conduit refilling (-0.05 MPa as calculated for sunflower minor veins). Mercurials supplied to detached leaves had no effect on the refilling process. Upon rehydration, recovery of K(vein) was not paralleled by recovery of whole-plant hydraulic conductance or leaf conductance to water vapour (g(L)), as a likely consequence of hydraulic failure of other components of the water pathway (root system or extravascular leaf compartments) and/or root-to-leaf chemical signalling. This is the first study providing experimental evidence for 'novel refilling' in a herbaceous dicot and highlighting the importance of this process in the leaf.     

Hydraulic vulnerability, vessel refilling, and seasonal courses of stem water potential of Sorbus aucuparia L. and Sambucus nigra L.

Differences in the seasonal variation in stem water potential between the two shrub species Sorbus aucuparia and Sambucus nigra were related with their vulnerability to xylem cavitation. It was also demonstrated indirectly that the two species differ in the extent to which they reverse cavitation. Seasonal variation in stem water potential was investigated during three growing seasons with in situ stem psychrometers. Sorbus experienced wide water potential variations and reached a minimum of -4.2 MPa during drought. Under the same microclimatic conditions, Sambucus experienced consistent stem water potentials with a minimum of -1.7 MPa. The relationship between percentage loss in hydraulic conductivity (PLC) and water potential (hydraulic vulnerability curve) of the two species differed in shape: a flat curve with nearly total loss of conductivity at -6 MPa was found for SORBUS: Sambucus showed a steep vulnerability curve with 90% loss conductivity at -2.2 MPa. Thus, Sambucus is extremely vulnerable to cavitation, but Sorbus is an almost invulnerable species. This different cavitation resistance adjusted the ranges of field stem water potential that the species experienced. Finally, seasonal courses of naturally occurring (native) embolism were compared with calculated PLC courses. This comparison indicates that Sorbus did not refill embolized xylem vessels whereas Sambucus reversed embolism. It was concluded that species which are highly vulnerable to cavitation and drought-induced embolism need refilling of embolized vessels as well as isohydric water potential patterns as two strategies of survival.     

Acclimation to Drought in Acer pseudoplatanus L. (Sycamore) Seedlings

A glasshouse experiment was conducted with well-watered andwater-stressed seedlings of sycamore (Acer pseudoplatanus L.)grown in soil columns. Water was withheld when the seedlingswere 82-d-old. Effects of soil drying on stomatal behaviour,water relations, xylem cavitation, and growth of leaves androots were evaluated. Stomatal conductance declined well before any observable changein bulk leaf water potentials, and was correlated with soilwater status. At seven weeks, osmotic potential had declinedby 0·51 MPa and 0·44 MPa at full and zero turgor,respectively. Drought significantly increased both bulk elasticmodulus and leaf dry weight to turgid weight ratio of water-stressedplants. Drought had no effect on relative water content at zeroturgor. Water cavitation in the xylem was detected as ultrasonic acousticemissions (AE). Water-stressed plants displayed significantlyhigher rates of AE than well-watered plants. Maximum rate ofAE coincided with the minimum level of stomatal conductanceand apparent rehydration of the leaves. Drought caused changes in the root distribution profile andit increased the root weight. The increase in root weight wasmainly due to a substantial shift in assimilates allocated infavour of roots with total biomass being unaffected. Leaf growthwas maintained for six weeks without any significant declinein expansion rate. However, the development of severe waterstress reduced both leaf production and expansion.     

Hydraulic redistribution in a stand of <Emphasis Type="Italic">Artemisia tridentata</Emphasis>: evaluation of benefits to transpiration assessed with a simulation model

The significance of soil water redistribution facilitated by roots (an extension of "hydraulic lift", here termed hydraulic redistribution) was assessed for a stand of Artemisia tridentata using measurements and a simulation model. The model incorporated water movement within the soil via unsaturated flow and hydraulic redistribution and soil water loss from transpiration. The model used Buckingham-Darcy's law for unsaturated flow while hydraulic redistribution was developed as a function of the distribution of active roots, root conductance for water, and relative soil-root (rhizosphere) conductance for water. Simulations were conducted to compare model predictions with time courses of soil water potential at several depths, and to evaluate the importance of root distribution, soil hydraulic conductance and root xylem conductance on transpiration rates and the dynamics of soil water. The model was able to effectively predict soil water potential during a summer drying cycle, and the rapid redistribution of water down to 1.5 m into the soil column after rainfall events. Results of simulations indicated that hydraulic redistribution could increase whole canopy transpiration over a 100-day drying cycle. While the increase was only 3.5% over the entire 100-day period, hydraulic redistribution increased transpiration up to 20.5% for some days. The presence of high soil water content within the lower rooting zone appears to be necessary for sizeable increases in transpiration due to hydraulic redistribution. Simulation results also indicated that root distributions with roots concentrated in shallow soil layers experienced the greatest increase in transpiration due to hydraulic redistribution. This redistribution had much less effect on transpiration with more uniform root distributions, higher soil hydraulic conductivity and lower root conductivity. Simulation results indicated that redistribution of water by roots can be an important component in soil water dynamics, and the model presented here provides a useful approach to incorporating hydraulic redistribution into larger models of soil processes.     

The effects of acclimation to sunlight on the xylem vulnerability to embolism in Fagus sylvatica L.

We assessed the effects of irradiance received during growth on the vulnerability of Fagus sylvatica L. xylem vessels to water-stress-induced embolism. The measurements were conducted on (1) potted saplings acclimated for 2 years under 100% and 12% incident global radiation and (2) branches collected from sun-exposed and shaded sides of adult trees. Both experiments yielded similar results. Light-acclimated shoots were less vulnerable to embolism. Xylem water potential levels producing 50% loss of hydraulic conductivity were lower in sun-exposed branches and seedlings than in shade-grown ones (–3·0 versus –2·3 MPa on average). The differences in vulnerability were not correlated with differences in xylem hydraulic conductivity nor vessel diameter. Resistance to cavitation was correlated with transpiration rates, midday xylem and leaf water potentials in adult trees. We concluded that vulnerability to cavitation in Fagus sylvatica may acclimate to contrasting ambient light conditions.