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 architecture of plants of Helianthus annuus L. cv. Margot: evidence for plant segmentation in herbs

The hydraulic architecture of sunflower (Helianthus annuus L. cv. Margot) was studied in terms of the partitioning of the hydraulic conductance (Kleaf) of leaves inserted at progressively more apical nodes both in growing plants (GP) and in plants at full anthesis (mature plants, MP). Leaf conductance to water vapour (gL), leaf water potential (PsiL), leaf water potential at zero turgor (Psi tlp), and leaf osmotic potential at full turgor (pi0) were also measured. Sunflower plants showed gL and Kleaf values significantly increasing in the acropetal direction, while PsiL of basal leaves was significantly more negative than that of distal leaves; Psi tlp markedly decreased in the acropetal direction in MP so that leaves of MP retained increasingly more turgor the more apical they were. This hydraulic pattern, already present in very young plants (GP), strongly favours apical leaves. These data suggest that the progressive leaf dieback starting from the stem base, as observed when the inflorescence of sunflower reached maturity, might be due to time-dependent loss of hydraulic conductance. In fact, Kleaf loss was correlated with PsiL drop and stomatal closure. Leaf dehydration was aggravated by solute exportation from the basal towards the apical leaves, as revealed by the acropetal decrease of pi0. Kleaf was shown to be linearly and positively related to the prevailing ambient irradiance during plant growth, thus suggesting that leaf hydraulics is very sensitive to environmental conditions. It was concluded that the pronounced apical dominance of some sunflower cultivars is determined, among other factors, by plant hydraulic architecture.     

An abscisic acid-related reduced transpiration promotes gradual embolism repair when grapevines are rehydrated after drought

* Proposed mechanisms of embolism recovery are controversial for plants that are transpiring while undergoing cycles of dehydration and rehydration. * Here, water stress was imposed on grapevines (Vitis vinifera), and the course of embolism recovery, leaf water potential (Psi(leaf)), transpiration (E) and abscisic acid (ABA) concentration followed during the rehydration process. * As expected, Psi(leaf) and E decreased upon water stress, whereas xylem embolism and leaf ABA concentration increased. Upon rehydration, Psi(leaf) recovered in 5 h, whereas E fully recovered only after an additional 48 h. The ABA content of recovering leaves was higher than in droughted controls, both on the day of rewatering and the day after, suggesting that ABA accumulated in roots during drought was delivered to the rehydrated leaves. In recovering plants, xylem embolism in petioles, shoots, and roots decreased during the 24 h following rehydration. * A model is proposed to describe plant recovery after rehydration based on three main points: embolism repair occurs progressively in shoots and further in roots and in petioles, following an almost full recovery of Psi(leaf); hydraulic conductance recovers during diurnal transpiring hours, when formation and repair of embolisms occurs in all plant organs; an ABA residual signal in rehydrated leaves hinders stomatal opening even when water relations have recovered, suggesting that an ABA-induced transpiration control promotes gradual embolism repair in rehydrated grapevines.     

Changes in leaf hydraulics and stomatal conductance following drought stress and irrigation in Ceratonia siliqua (Carob tree)

Changes in leaf hydraulic conductance (K) were measured using the vacuum chamber technique during dehydration and rehydration of potted plants of Ceratonia siliqua . K of whole, compound leaves as well as that of rachides and leaflets decreased by 20–30% at leaf water potentials (Ψ_L) of −1.5 and −2.0 MPa, i.e. at Ψ_L values commonly recorded in field-growing plants of the species. Higher K losses (up to 50%) were measured for leaves at Ψ_L of −2.5 and −3.0 MPa, i.e. near or beyond the leaf turgor loss point. Leaves of plants rehydrated while in the dark for 30 min, 90 min and 12 h recovered from K loss with characteristic times and to extents inversely proportional to the initial water stress applied. Leaf conductance to water vapour of plants dehydrated to decreasing Ψ_L and rehydrated at low transpiration was inversely related to loss of K, thus suggesting that leaf vein embolism and refilling (and related changes in leaf hydraulics) may play a significant role in the stomatal response.     

Declining hydraulic efficiency as transpiring leaves desiccate: two types of response

The conductance of transpiring leaves to liquid water (Kleaf) was measured across a range of steady-state leaf water potentials (Psileaf). Manipulating the transpiration rate in excised leaves enabled us to vary Psileaf in the range -0.1 MPa to less than -1.5 MPa while using a flowmeter to monitor the transpiration stream. Employing this technique to measure how desiccation affects Kleaf in 19 species, including lycophytes, ferns, gymnosperms and angiosperms, we found two characteristic responses. Three of the six angiosperm species sampled maintained a steady maximum Kleaf while Psileaf remained above -1.2 MPa, although desiccation of leaves beyond this point resulted in a rapid decline in Kleaf. In all other species measured, declining Psileaf led to a proportional decrease in Kleaf, such that midday Psileaf of unstressed plants in the field was sufficient to depress Kleaf by an average of 37%. It was found that maximum Kleaf was strongly correlated with maximum CO2 assimilation rate, while Kleaf = 0 occurred at a Psileaf slightly less negative than at leaf turgor loss. A strong linear correlation across species between Psileaf at turgor loss and Psileaf at Kleaf = 0 raises the possibility that declining Kleaf was related to declining cell turgor in the leaf prior to the onset of vein cavitation. The vulnerability of leaves rehydrating after desiccation was compared with vulnerability of leaves during steady-state evaporation, and differences between methods suggest that in many cases vein cavitation occurs only as Kleaf approaches zero.     

Impacts of tree height on leaf hydraulic architecture and stomatal control in Douglas-fir

This study investigated the mechanisms involved in the regulation of stomatal closure in Douglas-fir and evaluated the potential impact of compensatory adjustments in response to increasing tree height upon these mechanisms. In the laboratory, we measured leaf hydraulic conductance (K(leaf)) as leaf water potential (Psi(l)) declined for comparison with in situ diurnal patterns of stomatal conductance (g(s)) and Psi(l) in Douglas-fir across a height gradient, allowing us to infer linkages between diurnal changes in K(leaf) and g(s). A recently developed timed rehydration technique was used in conjunction with data from pressure-volume curves to develop hydraulic vulnerability curves for needles attached to small twigs. Laboratory-measured K(leaf) declined with increasing leaf water stress and was substantially reduced at Psi(l) values of -1.34, -1.45, -1.56 and -1.92 MPa for foliage sampled at mean heights of approximately 20, 35, 44 and 55 m, respectively. In situ g(s) measurements showed that stomatal closure was initiated at Psi(l) values of -1.21, -1.36, -1.74 and -1.86 MPa along the height gradient, which was highly correlated with Psi(l) values at loss of K(leaf). Cryogenic scanning electron microscopy (SEM) images showed that relative abundances of embolized tracheids in the central vein increased with increasing leaf water stress. Leaf embolism appeared to be coupled to changes in g(s) and might perform a vital function in stomatal regulation of plant water status and water transport in conifers. The observed trends in g(s) and K(leaf) in response to changes in Psi(l) along a height gradient suggest that the foliage at the tops of tall trees is capable of maintaining stomatal conductance at more negative Psi(l). This adaptation may allow taller trees to continue to photosynthesize during periods of greater water stress.     

Evidence for xylem embolism as a primary factor in dehydration-induced declines in leaf hydraulic conductance

Hydraulic conductance of leaves (K(leaf)) typically decreases with increasing water stress and recent studies have proposed different mechanisms responsible for decreasing K(leaf) . We measured K(leaf) concurrently with ultrasonic acoustic emissions (UAEs) in dehydrating leaves of several species to determine whether declining K(leaf) was associated with xylem embolism. In addition, we performed experiments in which the surface tension of water in the leaf xylem was reduced by using a surfactant solution. Finally, we compared the hydraulic vulnerability of entire leaves with the leaf lamina in three species. Leaf hydraulic vulnerability based on rehydration kinetics and UAE was very similar, except in Quercus garryana. However, water potentials corresponding to the initial decline in K(leaf) and the onset of UAE in Q. garryana were similar. In all species tested, reducing the surface tension of water caused K(leaf) to decline at less negative water potentials compared with leaves supplied with water. Microscopy revealed that as the fraction of embolized xylem increased, K(leaf) declined sharply in Q. garryana. Measurements on leaf discs revealed that reductions in lamina hydraulic conductance with dehydration were not as great as those observed in intact leaves, suggesting that embolism was the primary mechanism for reductions in K(leaf) during dehydration.     

Vein cavitation and stomatal behaviour of sunflower (Helianthus annuus) leaves under water limitation

The impact of leaf vein cavitation and embolism on stomatal response and leaf hydraulic conductance was studied in potted plants of sunflower subjected to water limitation. Plant dehydration was achieved either by cutting well‐watered plants near their base and leaving them dehydrating in air or by depriving intact plants of irrigation. The vein cavitation threshold (Ψ_CAV) was estimated in terms of ultrasound acoustic emissions (UAE) from the leaf blade versus leaf water potential (Ψ_L). This was found to be the same (Ψ_CAV ≈ ?0.6 MPa) for leaves of both cut and intact plants where stomata began to close in coincidence with starting vein cavitation. Vein embolism was detected by infiltrating leaves at different Ψ_L with 0.7 mM fluorescein and measuring the percentage fluorescent area as percentage of total leaf surface area. A distinct loss of vein functionality (up to 50%) was found to occur in leaves at progressively decreasing Ψ_L, starting when leaves reached Ψ_CAV. A linear positive relationship with high statistical significance was found to exist between g_L and percentage leaf fluorescent area, thus indicating that stomata were sensitive to vein embolism. The hydraulic conductance (K_L) of the leaf was affected by leaf dehydration less than expected (K_L decreased by about 20% between near full turgor and Ψ_L = ?1.3 MPa). When the extravascular leaf compartment was excluded either by killing cells by immersing leaves in 70% ethanol or by cutting the main leaf venous system through to allow flow to bypass it, K_L turned out to increase 5.5 times, thus suggesting that the high dominance of the hydraulic resistance of the extravascular leaf compartment over the total leaf resistance might buffer or mask possibly large local changes in K_L inducing stomatal closure.     

散孔材与环孔材树种枝干、叶水力学特性的比较研究

为揭示散孔材与环孔材树种树木水分生理特性的差异,选取了常见的3种散孔材落叶树种(毛白杨、法国梧桐和樱花)和3种环孔材落叶树种(刺槐、合欢和白蜡),研究了其枝干与叶水力学性质的差异及其协调性。结果表明:3种环孔材树种枝干横截面积基础上的最大比导水率(Ks-max)大于3种散孔材树种,但其木质部对空穴化的脆弱性(P50branch)高于散孔材树种,6种树木枝干的水分传输能力和抵抗空穴化能力之间存在一种相互制约的权衡关系。3种散孔材与3种环孔材树种的叶最大水力导度(Kl-max)和水力脆弱性(P50leaf)并无显著差异;对于3种散孔材树种,叶的水力脆弱性要高于枝干,但对3种环孔材树种而言,枝干的水力脆弱性要高于叶。6种树木枝干和叶的水力学性质(Kmax、P50)之间并无相关关系。这些结果表明:散孔材与环孔材树种的枝干水力学特性有明显差异,但叶水力学特性无差异;枝干与叶水力学性质之间是相互独立的。     

Decline of leaf hydraulic conductance with dehydration: relationship to leaf size and venation architecture

Across plant species, leaves vary enormously in their size and their venation architecture, of which one major function is to replace water lost to transpiration. The leaf hydraulic conductance (K(leaf)) represents the capacity of the transport system to deliver water, allowing stomata to remain open for photosynthesis. Previous studies showed that K(leaf) relates to vein density (vein length per area). Additionally, venation architecture determines the sensitivity of K(leaf) to damage; severing the midrib caused K(leaf) and gas exchange to decline, with lesser impacts in leaves with higher major vein density that provided more numerous water flow pathways around the damaged vein. Because xylem embolism during dehydration also reduces K(leaf), we hypothesized that higher major vein density would also reduce hydraulic vulnerability. Smaller leaves, which generally have higher major vein density, would thus have lower hydraulic vulnerability. Tests using simulations with a spatially explicit model confirmed that smaller leaves with higher major vein density were more tolerant of major vein embolism. Additionally, for 10 species ranging strongly in drought tolerance, hydraulic vulnerability, determined as the leaf water potential at 50% and 80% loss of K(leaf), was lower with greater major vein density and smaller leaf size (|r| = 0.85-0.90; P < 0.01). These relationships were independent of other aspects of physiological and morphological drought tolerance. These findings point to a new functional role of venation architecture and small leaf size in drought tolerance, potentially contributing to well-known biogeographic trends in leaf size.     

Leaf hydraulic capacity in ferns, conifers and angiosperms: impacts on photosynthetic maxima

* The hydraulic plumbing of vascular plant leaves varies considerably between major plant groups both in the spatial organization of veins, as well as their anatomical structure. * Five conifers, three ferns and 12 angiosperm trees were selected from tropical and temperate forests to investigate whether the profound differences in foliar morphology of these groups lead to correspondingly profound differences in leaf hydraulic efficiency. * We found that angiosperm leaves spanned a range of leaf hydraulic conductance from 3.9 to 36 mmol m2 s-1 MPa-1, whereas ferns (5.9-11.4 mmol m-2 s-1 MPa-1) and conifers (1.6-9.0 mmol m-2 s-1 MPa-1) were uniformly less conductive to liquid water. Leaf hydraulic conductance (Kleaf) correlated strongly with stomatal conductance indicating an internal leaf-level regulation of liquid and vapour conductances. Photosynthetic capacity also increased with Kleaf, however, it became saturated at values of Kleaf over 20 mmol m-2 s-1 MPa-1. * The data suggest that vessels in the leaves of the angiosperms studied provide them with the flexibility to produce highly conductive leaves with correspondingly high photosynthetic capacities relative to tracheid-bearing species.     

Dynamics of leaf hydraulic conductance with water status: quantification and analysis of species differences under steady state

Leaf hydraulic conductance (K(leaf)) is a major determinant of photosynthetic rate in well-watered and drought-stressed plants. Previous work assessed the decline of K(leaf) with decreasing leaf water potential (Ψ(leaf)), most typically using rehydration kinetics methods, and found that species varied in the shape of their vulnerability curve, and that hydraulic vulnerability correlated with other leaf functional traits and with drought sensitivity. These findings were tested and extended, using a new steady-state evaporative flux method under high irradiance, and the function for the vulnerability curve of each species was determined individually using maximum likelihood for 10 species varying strongly in drought tolerance. Additionally, the ability of excised leaves to recover in K(leaf) with rehydration was assessed, and a new theoretical framework was developed to estimate how rehydration of measured leaves may affect estimation of hydraulic parameters. As hypothesized, species differed in their vulnerability function. Drought-tolerant species showed shallow linear declines and more negative Ψ(leaf) at 80% loss of K(leaf) (P(80)), whereas drought-sensitive species showed steeper, non-linear declines, and less negative P(80). Across species, the maximum K(leaf) was independent of hydraulic vulnerability. Recovery of K(leaf) after 1 h rehydration of leaves dehydrated below their turgor loss point occurred only for four of 10 species. Across species without recovery, a more negative P(80) correlated with the ability to maintain K(leaf) through both dehydration and rehydration. These findings indicate that resistance to K(leaf) decline is important not only in maintaining open stomata during the onset of drought, but also in enabling sustained function during drought recovery.     

Effects of the experimental blockage of the major veins on hydraulics and gas exchange of Prunus laurocerasus L. leaves

The impact of leaf vein blockage on leaf hydraulic conductance (K(L)), gas exchange (g(L)) and water potential (Psi(L)) was studied in Prunus laurocerasus L., a broad-leaved evergreen. For this purpose, leaves were measured for the three variables above, either with an intact leaf blade (controls) or with the midrib cut a third of the way up (cut a), or with the midrib cut at three different points and the first-order veins cut through near their insertion to the midrib (cut b), or with the midrib cut at 2 mm from the leaf base (cut c). All the cut surfaces were sealed with cyanoacrylate. A serial decrease of K(L) was recorded from cut a to cut c with respect to that measured for the controls, i.e. a K(L) loss of about 37% (cut a), 57% (cut b) and 87% (cut c). A positive linear relationship appeared to exist between g(L) and K(L) with a high correlation coefficient (r(2)=0.99) and a high statistical significance (P <0.01). Even under a severe drop in K(L) (as that induced by cut c), leaf water potential remained approximately constant and not statistically different from Psi(L) measured for the controls. In fact, Psi(L) ranged between -0.83 and -0.98 MPa, i.e. within the cavitation threshold of leaves in terms of the critical Psi(L) inducing a significant production of ultrasound acoustic emissions which was -0.94+/-0.09 MPa. The conclusion was that stomata were very sensitive to changes in K(L) and that stomatal closure led to the homeostatic maintenance of Psi(L) and cavitation avoidance.     

Corticular photosynthesis drives bark water uptake to refill embolized vessels in dehydrated branches of Salix matsudana

It is well known that xylem embolism can be repaired by bark water uptake and that the sugar required for embolism refilling can be provided by corticular photosynthesis. However, the relationship between corticular photosynthesis and embolism repair by bark water uptake is still poorly understood. In this study, the role of corticular photosynthesis in embolism repair was assessed using Salix matsudana branch segments dehydrated to ?1.9 MPa (P₅₀, water potential at 50% loss of conductivity). The results indicated that corticular photosynthesis significantly promoted water uptake and nonstructural carbohydrate (NSC) accumulation in the bark and xylem during soaking, thereby effectively enhancing the refilling of the embolized vessels and the recovery of hydraulic conductivity. Furthermore, the influence of the extent of dehydration on the embolism refilling enhanced by corticular photosynthesis was investigated. The enhanced refilling effects were much higher in the mildly dehydrated (?1.5 MPa) and moderately dehydrated (?1.9 MPa) branch segments than in the severely dehydrated (?2.2 MPa) branch segments. This study provides evidence that corticular photosynthesis plays a crucial role in xylem embolism repair by bark water uptake for mildly and moderately dehydrated branches.     

Are needles of Pinus pinaster more vulnerable to xylem embolism than branches? New insights from X‐ray computed tomography

Plants can be highly segmented organisms with an independently redundant design of organs. In the context of plant hydraulics, leaves may be less embolism resistant than stems, allowing hydraulic failure to be restricted to distal organs that can be readily replaced. We quantified drought‐induced embolism in needles and stems of Pinus pinaster using high‐resolution computed tomography (HRCT). HRCT observations of needles were compared with the rehydration kinetics method to estimate the contribution of extra‐xylary pathways to declining hydraulic conductance. High‐resolution computed tomography images indicated that the pressure inducing 50% of embolized tracheids was similar between needle and stem xylem (P_{50 needle xylem} = ?3.62 MPa, P_{50 stem xylem} = ?3.88 MPa). Tracheids in both organs showed no difference in torus overlap of bordered pits. However, estimations of the pressure inducing 50% loss of hydraulic conductance at the whole needle level by the rehydration kinetics method were significantly higher (P_{50 needle} = ?1.71 MPa) than P_{50 needle xylem} derived from HRCT. The vulnerability segmentation hypothesis appears to be valid only when considering hydraulic failure at the entire needle level, including extra‐xylary pathways. Our findings suggest that native embolism in needles is limited and highlight the importance of imaging techniques for vulnerability curves.     

Stomatal closure during leaf dehydration,correlation with other leaf physiological traits

The question as to what triggers stomatal closure during leaf desiccation remains controversial. This paper examines characteristics of the vascular and photosynthetic functions of the leaf to determine which responds most similarly to stomata during desiccation. Leaf hydraulic conductance (K(leaf)) was measured from the relaxation kinetics of leaf water potential (Psi(l)), and a novel application of this technique allowed the response of K(leaf) to Psi(l) to be determined. These "vulnerability curves" show that K(leaf) is highly sensitive to Psi(l) and that the response of stomatal conductance to Psi(l) is closely correlated with the response of K(leaf) to Psi(l). The turgor loss point of leaves was also correlated with K(leaf) and stomatal closure, whereas the decline in PSII quantum yield during leaf drying occurred at a lower Psi(l) than stomatal closure. These results indicate that stomatal closure is primarily coordinated with K(leaf). However, the close proximity of Psi(l) at initial stomatal closure and initial loss of K(leaf) suggest that partial loss of K(leaf) might occur regularly, presumably necessitating repair of embolisms.     

Changes in leaf hydraulic conductance correlate with leaf vein embolism in<Emphasis Type="Italic"> Cercis siliquastrum</Emphasis> L.

The impact of xylem cavitation and embolism on leaf (K _leaf) and stem (K _stem) hydraulic conductance was measured in current-year shoots of Cercis siliquastrum L. (Judas tree) using the vacuum chamber technique. K _stem decreased at leaf water potentials (Ψ_L) lower than ?1.0 MPa, while K _leaf started to decrease only at Ψ_L L K _leaf changes. Field measurements of leaf conductance to water vapour (g _L) and Ψ_L showed that stomata closed when Ψ_L decreased below the Ψ_L threshold inducing loss of hydraulic conductance in the leaf. The partitioning of hydraulic resistances within shoots and leaves was measured using the high-pressure flow meter method. The ratio of leaf to shoot hydraulic resistance was about 0.8, suggesting that stem cavitation had a limited impact on whole shoot hydraulic conductance. We suggest that stomatal aperture may be regulated by the cavitation-induced reduction of hydraulic conductance of the soil-to-leaf water pathway which, in turn, strongly depends on the hydraulic architecture of the plant and, in particular, on leaf hydraulics.     

Contrasting hydraulic strategies in <Emphasis Type="Italic">Salix psammophila</Emphasis> and <Emphasis Type="Italic">Caragana korshinskii</Emphasis> in the southern Mu Us Desert,China

Salix psammophila and Caragana korshinskii are two common shrubs in the southern Mu Us Desert, China. Their hydraulic strategies for adapting to this harsh, dry desert environment are not yet clear. This study examined the hydraulic transport efficiency, vulnerability to cavitation, and daily embolism refilling in the leaves and stems of these two shrubs during the dry season. In order to gain insight into water use strategies of whole plants, other related traits were also considered, including daily changes in stomatal conductance, leaf mass per area, leaf pressure–volume parameters, wood density and the Huber value. The leaves and stems of S. psammophila had greater hydraulic efficiency, but were more vulnerable to drought-induced hydraulic dysfunction than C. korshinskii. The difference between leaf and stem water potential at 50 % loss of conductivity was 0.12 MPa for S. psammophila and 0.81 MPa for C. korshinskii. Midday stomatal conductance decreased by 74 % compared to that at 8:30 in S. psammophila, whereas no change occurred in C. korshinskii. Daily embolism and refilling occurred in the stems of S. psammophila and leaves of C. korshinskii. These results suggest that a stricter stomatal regulation, daily embolism repair in stems, and a higher stem water capacitance could be partially compensating for the greater susceptibility to xylem embolism in S. psammophila, whereas higher leaf elastic modulus, greater embolism resistance in stems, larger difference between leaf and stem hydraulic safety, and drought-induced leaf shedding in C. korshinskii were largely responsible for its more extensive distribution in arid and desert steppes.     

Root pressurization affects growth-induced water potentials and growth in dehydrated maize leaves

Profiles of water potential (Psi w) were measured from the soil to the tips of growing leaves of maize (Zea mays L.) when pressure (P) was applied to the soil/root system. At moderately low soil Psi w, leaf elongation was somewhat inhibited, large tensions existed in the xylem, and Psi w were slightly lower in the elongating leaf tissues than in the xylem, i.e. a growth-induced Psi w was present but small. With P, the tension was relieved, enlarging the difference in Psi w between the xylem and the elongating tissues, i.e. enlarging the growth-induced Psi w, which is critical for growth. Guttation occurred, confirming the high Psi w of the xylem, and the mature leaf tissue rehydrated. Water uptake increased and met the requirements of transpiration. Leaf elongation recovered to control rates. Under more severe conditions at lower soil Psi w, P induced only a brief elongation and the growth-induced Psi w responded only slightly. Guttation did not occur, water flow did not meet the requirements of transpiration, and the mature leaf tissues did not rehydrate. A rewatering experiment indicated that a low conductance existed in the severely dehydrated soil, which limited water delivery to the root and shoot. Therefore, the initial growth inhibition appeared to be hydraulic because the enlargement of the growth-induced Psi w by P together with rehydration of the mature leaf tissue were essential for growth recovery. In more severe conditions, P was ineffective because the soil could not supply water at the required rate, and metabolic factors began to contribute to the inhibition.     

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.