Xylem wall collapse in water-stressed pine needles

Wall reinforcement in xylem conduits is thought to prevent wall implosion by negative pressures, but direct observations of xylem geometry during water stress are still largely lacking. In this study, we have analyzed the changes in xylem geometry during water stress in needles of four pine species (Pinus spp.). Dehydrated needles were frozen with liquid nitrogen, and xylem cross sections were observed, still frozen, with a cryo-scanning electron microscope and an epifluorescent microscope. Decrease in xylem pressure during drought provoked a progressive collapse of tracheids below a specific threshold pressure (P(collapse)) that correlates with the onset of cavitation in the stems. P(collapse) was more negative for species with smaller tracheid diameter and thicker walls, suggesting a tradeoff between xylem efficiency, xylem vulnerability to collapse, and the cost of wall stiffening. Upon severe dehydration, tracheid walls were completely collapsed, but lumens still appeared filled with sap. When dehydration proceeded further, tracheids embolized and walls relaxed. Wall collapse in dehydrated needles was rapidly reversed upon rehydration. We discuss the implications of this novel hydraulic trait on the xylem function and on the understanding of pine water relations.     

Height-related trends in leaf xylem anatomy and shoot hydraulic characteristics in a tall conifer: safety versus efficiency in water transport

Hydraulic vulnerability of Douglas-fir (Pseudotsuga menziesii) branchlets decreases with height, allowing shoots at greater height to maintain hydraulic conductance (K shoot) at more negative leaf water potentials (Psi l). To determine the basis for this trend shoot hydraulic and tracheid anatomical properties of foliage from the tops of Douglas-fir trees were analysed along a height gradient from 5 to 55 m. Values of Psi l at which K shoot was substantially reduced, declined with height by 0.012 Mpa m(-1). Maximum K shoot was reduced by 0.082 mmol m(-2) MPa(-1) s(-1) for every 1 m increase in height. Total tracheid lumen area per needle cross-section, hydraulic mean diameter of leaf tracheid lumens, total number of tracheids per needle cross-section and leaf tracheid length decreased with height by 18.4 microm(2) m(-1), 0.029 microm m(-1), 0.42 m(-1) and 5.3 microm m(-1), respectively. Tracheid thickness-to-span ratio (tw/b)2 increased with height by 1.04 x 10(-3) m(-1) and pit number per tracheid decreased with height by 0.07 m(-1). Leaf anatomical adjustments that enhanced the ability to cope with vertical gradients of increasing xylem tension were attained at the expense of reduced water transport capacity and efficiency, possibly contributing to height-related decline in growth of Douglas fir.     

XYLEM ANATOMY AND HYDRAULIC CONDUCTANCE OF COSTA RICAN BLECHNUM FERNS

Hydraulic conductivities of stems, stipes, and elongate leaf stipes were determined for greenhouse-grown Blechnum (B. fraxineum, B. fragile, B. buchtienii, B. sprucei) and Salpichlaena (S. volubilis) plants collected in tropical rain forests of Costa Rica. Organ conductivity was examined in relation to morphology and tracheid characteristics in order to gain an understanding of factors influencing water flow. Hydraulic conductivity of plant organs was determined by measurement of transpiration rates, leaf areas, and water potential gradients. Erect stemmed species develop larger whole plant water potential gradients than elongate stemmed species for a similar transpiration rate. Elongate leaves develop even smaller water potential gradients for a given transpiration rate. Stems have larger hydraulic conductivities but smaller leaf-specific conductivities (LSCs) than stipes. Small conductivities and small LSCs are associated with short, erect stems. Elongate structures have large conductivities and large LSCs. Of the tracheid characteristics examined, the most important characteristics determining the magnitude of organ hydraulic conductivity are diameter, pit aperture area between tracheids, taper length, and cell length. Large conductivities of S. volubilis climbing leaf stipes are associated with very large-diameter tracheids (some > 200 μm), large tracheid number, exceptionally long tracheids (some > 4 cm), large pit aperture area between tracheids, short tracheid taper, and smooth tracheid lumen walls. Hagen-Poiseuille estimates of hydraulic conductivity range from 1.1 to 3.3 times the measured values. Conductivity of stipes is highly correlated with leaf area supplied by stipes. Conductivities of stems and elongate leaf stipes also correlate with leaf area supplied by these structures. Estimated hydraulic conductivities of field-grown Blechnum and Salpichlaena demonstrate that larger conductivities are associated with larger plants. This study contributes toward our knowledge of fern water relations and extends previous growth form/hydraulic architecture characterizations by providing a more comprehensive comparison of closely related species. In addition, this study provides evidence for the relative importance of tracheid characteristics in determining the magnitude of organ hydraulic conductivity.     

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.     

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.     

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.     

XYLEM ANATOMY,WATER FLOW,AND HYDRAULIC CONDUCTANCE IN THE FERN CYRTOMIUM FALCATUM

Xylem anatomy and water relations were studied in holly fern (Cyrtomium falcatum, Aspidiaceae) to determine the details of the pathway for water flow through an entire plant and the influence of tracheid number and lumen diameter on water flow. Each leaf has two adaxial traces and an abaxial trace, which are supplied by diarch adventitious roots attached to the dictyostele of the rhizome near the leaf base. Anatomical observations and dye experiments showed that each adaxial bundle vascularizes the approximately seven pinnae on its side of a leaf. An abaxial bundle is intermittently connected to an adaxial bundle as well as other abaxial bundles, forming a minor vascular pathway between the bundles of the leaf axis. Changes in both number and diameter of tracheids result in an acropetal decrease in hydraulic conductance per unit length along the rachis, although tracheid number locally increases when the trace for a pinna is produced in an adaxial bundle. Water flow was determined from the transpiration distal to the point in question or by forcing a solution through an axis with applied pressure. The water potential gradient along the plant axis was quite constant, indicating that hydraulic conductance per unit length varied with leaf area to be supplied. About 40% of the overall water potential drop occurred from the rachis into the pinnae, which reflected factors controlling water potential gradients in the lamina and not a very low conductance in the petiolule xylem. Hydraulic conductances calculated using the Hagen-Poiseuille equation and tracheid diameters were generally double those of measured conductances. Since the values tended to vary by a constant factor, tracheid number and diameter may largely control water flow in the xylem.     

A new method for vulnerability analysis of small xylem areas reveals that compression wood of Norway spruce has lower hydraulic safety than opposite wood

Compression wood is formed at the underside of conifer twigs to keep branches at their equilibrium position. It differs from opposite wood anatomically and subsequently in its mechanical and hydraulic properties. The specific hydraulic conductivity (k_s) and vulnerability to drought‐induced embolism (loss of conductivity versus water potential ψ) in twigs of Norway spruce [Picea abies (L.) Karst.] were studied via cryo‐scanning electron microscope observations, dye experiments and a newly developed ‘Micro‐Sperry’ apparatus. This new technique enabled conductivity measurements in small xylem areas by insertion of syringe cannulas into twig samples. The hydraulic properties were related to anatomical parameters (tracheid diameter, wall thickness). Compression wood exhibited 79% lower k_s than opposite wood corresponding to smaller tracheid diameters. Vulnerability was higher in compression wood despite its narrower tracheids and thicker cell walls. The P₅₀ (ψ at 50% loss of conductivity) was ?3.6 MPa in opposite but only ?3.2 MPa in compression wood. Low hydraulic efficiency and low hydraulic safety indicate that compression wood has primarily a mechanical function.     

Analysis of circular bordered pit function II. Gymnosperm tracheids with torus-margo pit membranes

A model of xylem conduit function was applied to gymnosperm tracheids with torus-margo pit membranes for comparison with angiosperm vessels. Tracheids from 17 gymnosperm tree species with circular bordered pits and air-seed pressures from 0.8 to 11.8 MPa were analyzed. Tracheids were more reinforced against implosion than vessels, consistent with their double function in transport and support. Tracheid pits were 3.3 to 44 times higher in hydraulic conductivity than vessel pits because of greater membrane conductivity of the torus-margo configuration. Tight scaling between torus and pit size maximized pit conductivity. Higher pit conductivity allowed tracheids to be 1.7-3.4 times shorter than vessels and still achieve 95% of their lumen-limited maximum conductivity. Predicted tracheid lengths were consistent with measured lengths. The torus-margo structure is important for maximizing the conductivity of the inherently length-limited tracheid: replacing the torus-margo membrane with a vessel membrane caused stem tracheid conductivity to drop by 41%. Tracheids were no less hydraulically efficient than vessels if they were long enough to reach their lumen-limiting conductivity. However, this may only be possible for lumen diameters below approximately 60-70 μm.     

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.     

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%.     

Minimum hydraulic safety leads to maximum water-use efficiency in a forage grass

Understanding how water-use regulation relates to biomass accumulation is imperative for improving crop production in water-limited environments. Here, we examine how the vulnerability of xylem to water stress-induced cavitation and the coordination between water transport capacity and assimilation (A) influences diurnal water-use efficiency (WUE) and dry-matter production in Lolium perenne L. - a commercial forage grass. Plants were exposed to a range of water stresses, causing up to 90% leaf death, by withholding water and then rewatering to observe the recovery process. Leaf hydraulic conductance (K(leaf) ) declined to 50% of maximum at a leaf water potential (ψ(leaf) ) of -1 MPa, whereas complete stomatal closure occurred well after this point, at -2.35 MPa, providing no protection against hydraulic dysfunction. Instantaneous A remained maximal until >70% of hydraulic conductivity had been lost. Post-stress rewatering showed that 95% loss of K(leaf) could be incurred before the recovery of gas exchange exceeded 1 d, with a rapid transition to leaf death after this point. Plants exposed to sustained soil water deficits through restricted nightly watering regimes did not suffer cumulative losses in K(leaf) ; instead, ψ(leaf) and gas exchange recovered diurnally. The effect was improved WUE during the day and optimal ψ(leaf) during the night for the maintenance of growth.     

Inter-tracheid pitting and the hydraulic efficiency of conifer wood: the role of tracheid allometry and cavitation protection

Plant xylem must balance efficient delivery of water to the canopy against protection from air entry into the conduits via air-seeding. We investigated the relationship between tracheid allometry, end wall pitting, safety from air-seeding, and the hydraulic efficiency of conifer wood in order to better understand the trade-offs between effective transport and protection against air entry. Root and stem wood were sampled from conifers belonging to the Pinaceae, Cupressaceae, Podocarpaceae, and Araucariaceae. Hydraulic resistivity of tracheids decreased with increasing tracheid diameter and width, with 64 ± 4% residing in the end wall pitting regardless of tracheid size or phylogenetic affinity. This end-wall percentage was consistent with a near-optimal scaling between tracheid diameter and length that minimized flow resistance for a given tracheid length. There was no evidence that tracheid size and hydraulic efficiency were constrained by the role of the pits in protecting against cavitation by air-seeding. An increase in pit area resistance with safety from cavitation was observed only for species of the northern hemisphere (Pinaceae and Cupressaceae), but this variable was independent of tracheid size, and the increase in pit resistance did not significantly influence tracheid resistance. In contrast to recent work on angiosperm vessels, protection against air-seeding in conifer tracheids appears to be uncoupled from conduit size and conducting efficiency.     

Kinetics of recovery of leaf hydraulic conductance and vein functionality from cavitation-induced embolism in sunflower

The kinetics of leaf vein recovery from cavitation-induced embolism was studied in plants of sunflower cv. Margot, together with the impact of vein embolism on the overall leaf hydraulic conductance (Kleaf). During the air-dehydration of leaves to leaf water potentials (Psi L) of -1.25 MPa, Kleaf was found to decrease by about 46% with respect to values recorded in well-hydrated leaves. When leaves, previously dehydrated to Psi L= -1.1 MPa (corresponding to the turgor loss point), were put in contact with water, Kleaf recovered completely in 10 min and so did leaf water potential. Functional vein density was estimated in both dehydrating and rehydrating leaves in terms of total length of red-stained veins infiltrated with a Phloxine B solution per unit leaf surface area. Veins were found to embolize (unstained) with kinetics showing a linear relationship with Kleaf so that about a 70% loss of functional veins corresponded with a Kleaf loss of 46%. Cavitated veins recovered from embolism within 10 min from the beginning of leaf rehydration. These data indicate that: (a) leaves of sunflower underwent substantial vein embolism during dehydration; (b) vein embolism and leaf hydraulic efficiency apparently recovered from dehydration completely and rapidly upon rehydration; (c) vein refilling occurred while conduits were still at more negative xylem pressures than those required for spontaneous bubble dissolution on the basis of Henry's law. The possible consistent contribution of vital mechanisms for vein refilling is discussed.     

Morphological adaptations of tracheid structure to water stress gradients in the crown of Sequoiadendron giganteum

Summary Mean radial diameter of tracheids in young branches of a 90 m Sequoiadendron giganteum decreases linearly with height along a gradient correlated linearly with decreasing xylem pressure potential. These smaller tracheid diameters provide strength to resist strong mechanical tensions in the xylem column and hypothetically allow greater efficiencies of water conduction. Tracheid length is not significantly correlated with either water stress or tracheid diameter.     

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.     

干旱胁迫对油松和侧柏水分运输安全性和有效性的影响

通过采用改良的冲洗法测定5年生苗木油松(Pinus tabulaeformis)和侧柏(Platycladus orientalis)在不同干旱胁迫时期的水力结构参数,结果表明随着干旱胁迫增加,不同分枝级和茎段所在区域的导水率损失(PLC)增加,比导率(Ks)减少。油松和侧柏在0级,1级和2级分枝级的发生栓塞的水势阈值分别为-0.55、-0.49、-0.43和-0.90、-0.78、-0.74MPa。随着相对分枝级的增加,油松和侧柏的水势阈值增大,栓塞脆弱性变大。非限速区PLC大而Ks小,限速区PLC小而Ks大。油松和侧柏相对分枝级和茎段所在区域的栓塞脆弱性大小为2级1级0级,限速区非限速区,且油松大于侧柏。油松和侧柏在不同干旱胁迫,不同相对分枝级,不同茎段所在区域采取不同的方式来适应由水势降低而引起的栓塞变化。其采取的生态策略包括:保持较高的水分安全性;减轻安全性而对有效性的折衷;同时降低有效性和安全性但不终止任何生产力或树高的组织生长所需水的限制。     

Plasticity of stomatal distribution pattern and stem tracheid dimensions in Podocarpus lambertii: an ecological study

Background and Aims

Leaf and wood plasticity are key elements in the survival of widely distributed plant species. Little is known, however, about variation in stomatal distribution in the leaf epidermis and its correlation with the dimensions of conducting cells in wood. This study aimed at testing the hypothesis that Podocarpus lambertii, a conifer tree, possesses a well-defined pattern of stomatal distribution, and that this pattern can vary together with the dimensions of stem tracheids as a possible strategy to survive in climatically different sites.

Methods

Leaves and wood were sampled from trees growing in a cold, wet site in south-eastern Brazil and in a warm, dry site in north-eastern Brazil. Stomata were thoroughly mapped in leaves from each study site to determine a spatial sampling strategy. Stomatal density, stomatal index and guard cell length were then sampled in three regions of the leaf: near the midrib, near the leaf margin and in between the two. This sampling strategy was used to test for a pattern and its possible variation between study sites. Wood and stomata data were analysed together via principal component analysis.

Key Results

The following distribution pattern was found in the south-eastern leaves: the stomatal index was up to 25 % higher in the central leaf region, between the midrib and the leaf margin, than in the adjacent regions. The inverse pattern was found in the north-eastern leaves, in which the stomatal index was 10 % higher near the midrib and the leaf margin. This change in pattern was accompanied by smaller tracheid lumen diameter and length.

Conclusions

Podocarpus lambertii individuals in sites with higher temperature and lower water availability jointly regulate stomatal distribution in leaves and tracheid dimensions in wood. The observed stomatal distribution pattern and variation appear to be closely related to the placement of conducting tissue in the mesophyll.     

沙打旺根系提水作用及其机理研究

采用上下桶分根法研究了3年生沙打旺的根系提水作用及土壤水势与植物组织水势、植物渗透调节物质之间的关系。结果表明,当上下桶土壤体积含水量和水势分别在14.9%和-1.28MPa、19%和-0.6MPa左右时,下桶土壤水势>下桶根水势>上桶根水势>上桶土壤水势>叶水势,出现沙打旺根系提水现象。上桶土壤含水量和土壤水势的日变化在晴天表现为:7:00～16:00急剧下降,16:00～22:00上升较快并于22:00达到最大值,之后缓慢下降;而其在阴天随时间的推移呈现缓慢下降趋势。沙打旺叶片中K 、可溶性糖和脯氨酸含量等渗透调节物质在16:00和22:00均高于根中,16:00上桶根和叶片中脯氨酸、可溶性糖、K 和Na 的含量显著高于22:00,由此产生了上桶根水势、叶水势的日变化,促进了沙打旺的提水作用。     

Unraveling the Effects of Plant Hydraulics on Stomatal Closure during Water Stress in Walnut

The objectives of the study were to identify the relevant hydraulic parameters associated with stomatal regulation during water stress and to test the hypothesis of a stomatal control of xylem embolism in walnut (Juglans regia x nigra) trees. The hydraulic characteristics of the sap pathway were experimentally altered with different methods to alter plant transpiration (Eplant) and stomatal conductance (gs). Potted trees were exposed to a soil water depletion to alter soil water potential (Psisoil), soil resistance (Rsoil), and root hydraulic resistances (Rroot). Soil temperature was changed to alter Rroot alone. Embolism was created in the trunk to increase shoot resistance (Rshoot). Stomata closed in response to these stresses with the effect of maintaining the water pressure in the leaf rachis xylem (P(rachis)) above -1.4 MPa and the leaf water potential (Psileaf) above -1.6 MPa. The same dependence of Eplant and gs on P(rachis) or Psileaf was always observed. This suggested that stomata were not responding to changes in Psisoil, Rsoil, Rroot, or Rshoot per se but rather to their impact on P(rachis) and/or Psileaf. Leaf rachis was the most vulnerable organ, with a threshold P(rachis) for embolism induction of -1.4 MPa. The minimum Psileaf values corresponded to leaf turgor loss point. This suggested that stomata are responding to leaf water status as determined by transpiration rate and plant hydraulics and that P(rachis) might be the physiological parameter regulated by stomatal closure during water stress, which would have the effect of preventing extensive developments of cavitation during water stress.