Axial and radial profiles in conductivities, water storage and native embolism in trunks of young and old-growth ponderosa pine trees

Ponderosa pine has very wide sapwood, and yet the spatial and temporal use of that sapwood for water transport is poorly understood. Moreover, there have been few comparisons of function in tips of old-growth trees in comparison with young trees. In the present study, axial and radial specific conductivity (k_s), leaf specific conductivity (LSC), leaf specific conductance (k_l), native embolism and the compartmentalization of sapwood water storage were characterized in trunks of young and old-growth trees. Trunks of young trees had lower k_s, lower LSC and lower native embolism [corresponding to 5% loss of conductivity (PLC)] than trunks of old-growth trees. However, k_l in young trees was 3.5 times higher than in old-growth trees, supporting the hypothesis that tall trees have a reduced ability to transport water to their leaves. Water storage (capacitance) of young trees was not significantly different than at the base of old-growth trees. Although the top of the old-growth trees had similar k_s, LSC and k_l to the young trees for a given cambial age, they had higher native embolism and lower capacitance. There was no trade-off between k_s and native embolism at any height. In the tree crown, outer sapwood had 35–50% higher k_s than the inner sapwood and 17–25 PLC lower native embolism. At the base of the old trees, there was no significant difference in native embolism between the outer, middle and inner sapwood, showing that refilling of embolisms was complete despite the 130-year difference in wood age among these radial positions. Although during the dry season the inner sapwood tended to be more saturated than the outer sapwood, the outer part of the sapwood contributed up to 60% of the overall stored water. Safer xylem, higher capacitance and higher k_l would appear adaptive in the young trees for regulating their water resource, which is likely to be less reliable than the water availability of older trees with their more developed root system.     

Hydraulic redistribution of water from Pinus ponderosa trees to seedlings: evidence for an ectomycorrhizal pathway

While there is strong evidence for hydraulic redistribution (HR) of soil water by trees, it is not known if common mycorrhizal networks (CMN) can facilitate HR from mature trees to seedlings under field conditions. Ponderosa pine (Pinus ponderosa) seedlings were planted into root-excluding 61-microm mesh barrier chambers buried in an old-growth pine forest. After 2 yr, several mature trees were cut and water enriched in D(2)O and acid fuchsin dye was applied to the stumps. Fine roots and mycorrhizal root tips of source trees became heavily dyed, indicating reverse sap flow in root xylem transported water from stems throughout root systems to the root hyphal mantle that interfaces with CMN. Within 3 d, D(2)O was found in mesh-chamber seedling foliage > 1 m from source trees; after 3 wk, eight of 10 mesh-chamber seedling stem samples were significantly enriched above background levels. Average mesh-chamber enrichment was 1.8 x greater than that for two seedlings for which the connections to CMN were broken by trenching before D(2)O application. Even small amounts of water provided to mycorrhizas by HR may maintain hyphal viability and facilitate nutrient uptake under drying conditions, which may provide an advantage to seedlings hydraulically linked by CMN to large trees.     

Hydraulic redistribution of soil water in two old-growth coniferous forests: quantifying patterns and controls

Although hydraulic redistribution of soil water (HR) by roots is a widespread phenomenon, the processes governing spatial and temporal patterns of HR are not well understood. We incorporated soil/plant biophysical properties into a simple model based on Darcy's law to predict seasonal trajectories of HR. We investigated the spatial and temporal variability of HR across multiple years in two old-growth coniferous forest ecosystems with contrasting species and moisture regimes by measurement of soil water content (theta) and water potential (Psi) throughout the upper soil profile, root distribution and conductivity, and relevant climate variables. Large HR variability within sites (0-0.5 mm d(-1)) was attributed to spatial patterns of roots, soil moisture and depletion. HR accounted for 3-9% of estimated total site water depletion seasonally, peaking at 0.16 mm d(-1) (ponderosa pine; Pinus ponderosa) or 0.30 mm d(-1) (Douglas-fir; Pseudotsuga menziesii), then declining as modeled pathway conductance dropped with increasing root cavitation. While HR can vary tremendously within a site, among years and among ecosystems, this variability can be explained by natural variability in Psi gradients and seasonal courses of root conductivity.     

Tree proximity,soil pathways and common mycorrhizal networks: their influence on the utilization of redistributed water by understory seedlings

Hydraulic redistribution (HR) is a process by which water moves through plant roots from moist to dry soils. An experiment was conducted to quantify the influence of common mycorrhizal networks (CMNs) and proximity to mature HR-source trees on the water relations of surrounding seedlings. Douglas-fir (Pseudotsuga menziesii var glauca (Mirb.) Franco) seedlings were planted at four distances (0.5, 1, 2.5, and 5 m) from six mature Douglas-fir trees, either directly into soil (soil plus CMN pathway) or inside 0.5 μm mesh bags (soil-only pathway). Deuterated water was used to irrigate soil beside mature trees in order to identify different HR water pathways to surrounding seedlings. This was followed by measurements of seedling deuterium enrichment, seedling water potential, soil water potential_, gravimetric soil water content, and tree root density surrounding the seedlings. There was no significantly detectable difference in the quantity of HR water transferred to seedlings having access to soil and CMN pathways or soil-only pathways of water movement. Water from the irrigation plot contributed up to 1.4% of the water of Douglas-fir seedlings. Based on the assumption that the only pathway through which seedlings could access irrigation water was through the mature trees, we estimate that as much as 21.6% of the seedling water was supplied by the nearby tree. Seedling water potential was not significantly affected either by proximity to mature trees or pathway, suggesting HR may have compensated for increasing tree competitive effects with proximity. It is also possible that the lack of difference was due to a relatively moist summer. Our results suggest that residual mature trees are potentially important for hydraulic redistribution to regenerating seedlings in harvested dry interior Douglas-fir forests.     

Effects of Restoration Thinning on Presettlement Pinus ponderosa in Northern Arizona

In the 100 years following the arrival of Euro-American settlers in northern Arizona, Pinus ponderosa (ponderosa pine) forests changed from open, low-density stands to closed, high-density stands. The increase in tree density has been detrimental to the vigor of old-growth trees that established before settlement (presettlement trees). In this study, we examined whether the vigor of presettlement trees could be improved by restoring the original stand structure by thinning the ponderosa pines that established after settlement (postsettlement trees). The restoration treatment caused the following changes in the presettlement trees and their environment in the first year following thinning: an increase in volumetric soil water content between May and August, an increase in predawn xylem water potential in July and August, a decrease in midday xylem water potential in June and August, an increase in net photosynthetic rate in August, an increase in foliar nitrogen concentration in July and August, and an increase in bud and needle size. The results show that the thinning restoration treatment improved the condition of presettlement ponderosa pines by increasing canopy growth and the uptake of water, nitrogen, and carbon.     

The canopy water relations of old-growth Douglas-fir trees

 We investigated whole tree water relations in 56–65 m tall, old-growth Pseudotsuga menziesii trees within the Wind River Canopy Crane site, Carson, Washington, USA. We measured at predawn and solar noon the vertical gradients in xylem pressure potential using a pressure chamber. On an Abies amabilis sapling located in the understory at the base of one of the study trees, predawn and solar noon xylem pressure potentials were also measured. Xylem pressure potential data were measured from late June through early September 1996 on foliage sampled from 1 to 64.5 m. Over this height gradient, predawn water potentials ranged from –0.23 to –1.10 MPa. Solar noon values showed an even greater range (from –0.44 to –2.51 MPa). At predawn, the water potential gradient approached the theoretical hydrostatic gradient (–0.0105 vs –0.010 MPa m^–1). The gradient at solar noon was steeper (–0.0331 MPa m^–1). Instantaneous stomatal conductances were not greatly different between young, sapling-sized and old-growth trees [0.094±0.033 (SD) vs 0.086±0.045 cm s^–1, respectively]. Stomata of both size classes of trees appeared very sensitive to increasing vapor pressure deficits. A comparison of stable carbon isotope values from the old-growth and sapling-sized trees indicated lower stomatal conductances in the old-growth. This study provides sound documentation regarding the utility of the cohesion theory in the interpretation of water potential gradients. This study also emphasizes inherent differences between sapling-sized and tall, old-growth trees. Received: 10 January 1998 / Accepted: 12 October 1998     

Patterns of tree dieback in Queensland,Australia: the importance of drought stress and the role of resistance to cavitation

During the extreme 1992–1997 El Niño drought event, widespread stem mortality, or tree dieback, of both mature and juvenile eucalypts occurred within the tropical savannas of northeast Australia. Most of the dieback occurred in individuals of the ironbark species complex (Eucalyptus crebra – E. xanthoclada) while individuals of the bloodwood species Corymbia erythrophloia, exhibited significantly less stem mortality. Indicative of greater water stress, predawn and midday xylem water potentials of ironbark adults and saplings were significantly more negative than predawn values of bloodwoods. The very negative xylem water potentials in ironbarks suggest that stem mortality in both adult and juvenile ironbarks results from drought-induced embolism and that ironbarks perhaps have a shallower and less extensive root system than bloodwoods. Although predawn and midday water potentials for ironbark adults and saplings were similar, a census of mature and juvenile ironbark trees indicated that mortality was higher in adult trees. Cavitation vulnerability curves indicated that ironbark saplings may be better buffered against cavitation than adult trees. If they possess smaller root systems, saplings are more likely than adults to experience low xylem water potentials, even in non-drought years. Xylem conduits produced in adult trees during periods of normal rainfall, although perhaps more efficient in water conduction, may be more vulnerable to cavitation during infrequent severe droughts.     

摘叶造成的碳限制对刺槐碳素分配和水力学特性的影响

以3年生刺槐（Robinia pseudoacacia Linn.）为研究对象,通过对其进行连续3次摘叶造成严重碳限制,检测摘叶后刺槐的生物量分配、叶片形态和不同部位的非结构性碳（NSC）浓度,同时检测其根压和根系导水率、枝条水势和导水率损失值（PLC）及茎的抗栓塞能力,研究摘叶造成的碳限制对刺槐碳素分配和水力学特性的影响。结果显示,摘叶显著降低了刺槐不同部位的生物量,其中细根生物量降低程度最大;摘叶还造成了刺槐不同部位NSC浓度显著降低,茎韧皮部、茎木质部、根韧皮部和根木质部的NSC浓度分别为对照的29.6%、20.2%、10.2%和8.7%,且根部NSC的降低程度显著高于茎;碳限制显著降低了刺槐苗木的根压和根系导水率,增加了枝条凌晨和正午的PLC,降低了其抗栓塞能力。研究结果表明摘叶造成的碳限制改变了刺槐的碳素分配模式,限制了碳素向根的分配,抑制细根的发生,进而限制根的水分吸收能力,加重枝条栓塞程度,同时还会导致枝条抗栓塞能力下降,从而降低植物水分输导的安全性。     

Hydraulic properties of Pinus halepensis,Pinus pinea and Tetraclinis articulata in a dune ecosystem of Eastern Spain

The hydraulic properties of Pinus pinea, Pinus halepensis and Tetraclinis articulata were studied in a coastal dune area from Eastern Spain. The measured variables include vulnerability to xylem embolism (vulnerability curves), hydraulic conductivity and carbon isotopic discrimination in leaves. Leaf water potentials were also monitored in the three studied populations during an extremely dry period. Our results showed that roots had always wider vessels and higher hydraulic conductivity than branches. Roots were also more vulnerable to xylem embolism and operated closer to their hydraulic limit (i.e., with narrower safety margins). Although it was not quantified, extensive root mortality was observed in the two pines during the study period, in agreement with the high values of xylem embolism (> 75%) predicted from vulnerability curves and the water potentials measured in the field. T. articulata was much more resistant to embolism than P. pinea and P. halepensis. Since T. articulata experienced also lower water potentials, safety margins from hydraulic failure were only slightly wider in this species than in the pines. Combining species and tissues, high resistance to xylem embolism was associated with low hydraulic conductivity and with high wood density. Both relationships imply a cost of having a resistant xylem. The study outlined very different water-use strategies for T. articulata and the pines. Whereas T. articulata had a conservative strategy that relied on the low vulnerability of its conducting system to drought-induced xylem embolism, the two pines showed regulatory mechanisms at different levels (i.e., embolism, root demography) that constrained the absorption of water when it became scarce. This revised version was published online in August 2006 with corrections to the Cover Date.     

Bayesian analysis of Douglas-fir hydraulic architecture at multiple scales

We used a Bayesian hierarchical model to analyze the variation in xylem anatomy, hydraulic properties, and the relationship between anatomy and properties within Douglas-fir trees. The hierarchical scales in our study included fertilization treatments (fertilized and unfertilized), trees within the treatments, and positions within the trees. We measured tracheid diameter, tracheid length, percent latewood, number of pits per cell, density, and specific conductivity (K _s) on seven positions in each of 16 fertilized and 16 unfertilized trees: the trunk at cambial age 52 (breast height), 25, and 5; a branch at cambial age 20 and 7; and a root at cambial age 42 and 22. Vulnerability to embolism was also measured on the oldest trunk, branch, and root positions. For any measurement, there was little variation between treatments, however, there was great variation among positions. Tracheid diameter, tracheid length, number of pits per cell, K _s, and vulnerability to embolism decreased vertically from the roots to the branches. Correlations were evident between some positions for tracheid diameter, percent earlywood, pits per cell, and vulnerability to embolism, mostly in the fertilized treatment. We found evidence for large-scale relationships (among all observations from all trees) between density and tracheid diameter, K _s and diameter, vulnerability and diameter, K _s and pits per cell, and vulnerability and pits per cell. At a smaller scale of within position, however, usually only the branches and roots maintained the relationship.     

Ectomycorrhizal fungi associated with ponderosa pine and Douglas-fir: a comparison of species richness in native western North American forests and Patagonian plantations from Argentina

The putative ectomycorrhizal fungal species registered from sporocarps associated with ponderosa pine and Douglas-fir forests in their natural range distribution (i.e., western Canada, USA, and Mexico) and from plantations in south Argentina and other parts of the world are listed. One hundred and fifty seven taxa are reported for native ponderosa pine forests and 514 taxa for native Douglas-fir forests based on available literature and databases. A small group of genera comprises a high proportion of the species richness for native Douglas-fir (i.e., Cortinarius, Inocybe, and Russula), whereas in native ponderosa pine, the species richness is more evenly distributed among several genera. The comparison between ectomycorrhizal species richness associated with both trees in native forests and in Patagonia (Argentina) shows far fewer species in the latter, with 18 taxa for the ponderosa pine and 15 for the Douglas-fir. Epigeous species richness is clearly dominant in native Douglas-fir, whereas a more balanced relation epigeous/hypogeous richness is observed for native ponderosa pine; a similar trend was observed for Patagonian plantations. Most fungi in Patagonian Douglas-fir plantations have not been recorded in plantations elsewhere, except Suillus lakei and Thelephora terrestris, and only 56% of the fungal taxa recorded in Douglas-fir plantations around the world are known from native forests, the other taxa being new associations for this host, suggesting that new tree + ectomycorrhizal fungal taxa associations are favored in artificial situations as plantations.     

Vulnerability to embolism differs in roots and shoots and among three Mediterranean conifers: consequences for stomatal regulation of water loss?

We investigated the potential links between stomatal control of transpiration and the risk of embolism in root and shoot xylem of seedlings of three Mediterranean conifers (Cupressus sempervirens, Pinus halepensis and P. nigra) grown in a greenhouse under semi-controlled conditions. We measured the intrinsic vulnerability to embolism in roots and current year shoots by the air injection method. Root and shoot segments were subjected to increasing pressures, and the induced loss of hydraulic conductivity recorded. The three species displayed very different vulnerabilities in shoots, with P. nigra being much more vulnerable than P. halepensis and C. sempervirens. Roots were distinctly more vulnerable than shoots in C. sempervirens and P. halepensis (50% loss of conductivity induced at 3.0 MPa and 1.7 MPa higher xylem water potential in roots vs shoots). In P. nigra, no significant difference of vulnerability between shoots and roots was found. Seedlings were subjected to soil drought, and stomatal conductance, twig hydraulic conductivity and needle water potential were measured. The water potential resulting in almost complete stomatal closure (90%) was very close to the threshold water potential inducing loss of conductivity (10%) in twigs in P nigra, resulting in a very narrow safety margin between stomatal closure and embolism induction. The safety margin was larger in P. halepensis and greatest in C. sempervirens. Unexpectedly, this water potential threshold produced a 30–50% loss of conductivity in 3–5 mm diameter roots, depending on the species. The implications of this finding are discussed.     

Hydraulic architecture of deciduous and evergreen dry rainforest tree species from north-eastern Australia

Hydraulic conductivity and xylem anatomy were examined in stems of two evergreen species, Alphitonia excelsa (Fenzal) Benth. and Austromyrtus bidwillii (Benth.) Burret., and two drought-deciduous species, Brachychiton australis (Schott and Endl.) A. Terracc. and Cochlospermum gillivraei Benth., from a seasonally dry rainforest in north Queensland, Australia. The deciduous species possessed hydraulic architecture typical of drought-sensitive plants, i.e. low wood density, wider xylem vessels, higher maximal rates of sapwood specific hydraulic conductivity (K_s) and high vulnerability to drought-induced embolism. In contrast, the evergreen species had lower rates of K_h and leaf specific conductivity (K_L) but were less susceptible to embolism. The evergreen species experienced leaf water potentials <–4.0 MPa during the dry season, while the deciduous species shed their leaves before leaf water potentials declined below –2.0 MPa. Thus, the hydraulic architecture of the evergreens allows them to withstand the greater xylem pressure gradients required to maintain water transport to the canopy during the dry season. Our results are consistent with observations made in neotropical dry forests and demonstrate that drought-deciduous species with low wood density and high water storage capacity are likely to be more hydraulically efficient, but more vulnerable to embolism, than coexisting evergreens.     

Different responses to drought and freeze stress of Quercus ilex L. growing along a latitudinal gradient

The vulnerability to drought and freeze stress was measured in young plants of Quercus ilex L. growing in the field in two natural sites within the Italian distribution area of this species, i.e. Sicily (Southern Mediterranean Basin) and Venezia Giulia (Northeastern Italy), respectively. In particular, the resistance strategies adopted by Q. ilex to withstand the two stresses were estimated in terms of seasonal and/or diurnal changes in leaf conductance to water vapour (g_L), water potential (_L) and relative water content (RWC) as well as of xylem embolism in the stem and root hydraulic conductance (K_RL). Sicilian (SI) plants showed to reduce water loss by stomatal closure (g_L decreased) in summer, thus maintaining average RWCs at 88–90%. Moreover, SI plants showed considerable resistance to xylem cavitation in the stem (the loss of hydraulic conductance, PLC, was less than 12% throughout the year) and to maintain the hydraulic conductance of their roots (K_RL), constantly high even in summer. Plants growing in Venezia Giulia (VG plants), on the contrary, underwent leaf dehydration in the winter due to freeze stress so that RWC measured in April was still 78% on a diurnal basis. This was apparently due to consistent xylem embolism in the stem. In fact, PLC was as high as 40% between November and March. Only in the summer was PLC similar to that recorded in SI plants. Moreover, K_RL of VG plants decreased in November from about 1.5 to 0.8×10^–4 kg s^–1 m^–2 MPa^–1, i.e. about 50%, and in February K_RL dropped further to 0.4×10^–4 kg s^–1 m^–2 MPa^–1. On the basis of the above, we conclude that: (a) Q. ilex was more sensitive to freeze than to drought stress so that freeze stress can be considered as a factor limiting the distribution area of this species; (b) drought and freeze stress were faced by Q. ilex adopting two different resistance strategies, i.e. drought avoidance based on water saving in Sicily and freeze tolerance in Venezia Giulia.     

Relationship between growth rates and xylem hydraulic characteristics in young, mature and old-growth ponderosa pine trees

The first objective of the present study was to quantify the effects of tree age and stem position on specific conductivity (k_s), vulnerability to embolism and water storage capacity (capacitance) in trunks of young, mature and old‐growth ponderosa pine. The second objective was to determine relationships between hydraulic characteristics and radial and height growth rates to increase the understanding of possible tradeoffs. Within sapwood at all heights and in all ages of trees, outer sapwood had 25–60% higher k_s than inner sapwood. The water potential at which embolism started (air entry point) was 1.3 MPa lower in inner sapwood than outer sapwood within the mature trees, but there was no difference in the other trees. There was no significant difference in capacitances between the tops of the old growth trees, the mature trees and the young trees. Taking all data together, the capacitances increased sharply with an increase in k_s and an increase in vulnerability to embolism. The hydraulic characteristics of the three age classes were correlated with the height growth rate but not with the diameter growth rate. Within these age classes, high k_s was associated with the slowest yearly increase in sapwood area and with a low percentage of latewood, whereas high vulnerability to embolism and high capacitance were more closely associated with high height growth rates.     

极端干旱环境下的胡杨木质部水力特征

胡杨作为我国西北干旱区重要的乔木树种,研究其木质部水力特征对了解此树种适应极端干旱环境的生物学背景具有较重要的意义。本研究以塔里木河下游的胡杨成株和2年生胡杨幼苗为研究材料,对其木质部最大导水能力（ks(max)）和自然栓塞程度（PLC）等木质部水力特征及其水力特征有关的木质部导管(或管饱)数量特征进行研究。结果表明,成株胡杨多年生枝条和侧根（2≤d<5 mm）木质部自然栓塞程度均较高,PLC均值高于50%,其中多年生枝条栓塞程度具有一定的日变化规律,清晨的PLC均值（58%）小于正午的（67%）;河道边上成株胡杨侧根的均ks(max)和PLC均值都小于距河道200 m处的。随着土壤干旱程度的加剧,幼苗胡杨侧根的自然栓塞程度随之增加,而叶片气孔导度随之降低,土壤含水率与侧根自然栓塞程度,叶片气孔导度之间分别存在显著负相关关系（R =-0.9、R =-0.811）。在统一直径范围内(2≤d<5 mm),成株胡杨侧根均导管直径（dmean）和水力直径均大于（d95%、dh）胡杨幼苗,而导管密度胡杨幼苗高于成株胡杨;胡杨侧根木质部最大导水能力与均导管直径、水力直径之间具有显著正相关关系（R>0.9）.     

Hydraulic propagation of pressure along immature and mature xylem vessels of roots of Zea mays measured by pressure-probe techniques

Turgor pressure was measured in cortical cells and in xylem elements of excised roots and roots of intact plants of Zea mays L. by means of a cell pressure probe. Turgor of living and hence not fully differentiated late metaxylem (range 0.6–0.8 MPa) was consistently higher than turgor of cortical cells (range 0.4–0.6 MPa) at positions between 40 and 180 mm behind the root tip. Closer to the tip, no turgor difference between the cortex and the stele was measured. The turgor difference indicated that late-metaxylem elements may function as nutrient-storage compartments within the stele. Excised roots were attached to the root pressure probe to precisely manipulate the xylem water potential. Root excision did not affect turgor of cortical cells for at least 8 h. Using the cell pressure probe, the propagation of a hydrostatic pressure change effected by the root pressure probe was recorded in mature and immature xylem elements at various positions along the root. Within seconds, the pressure change propagated along both early and late metaxylems. The half-times of the kinetics, however, were about five times smaller for the early metaxylem, indicating they are likely the major pathway of longitudinal water flow. The hydraulic signal dissipated from the source of the pressure application (cut end of the root) to the tip of the root, presumably because of radial water movement along the root axis. The results demonstrate that the water status of the growth zone and other positions apical to 20 mm is mainly uncoupled from changes of the xylem water potential in the rest of the plant.Abbreviations and Symbols CPP cell pressure probe - EMX early metaxylem - LMX Late metaxylem - P_c cell turgor - P_r root pressure - RPP root pressure probe - t_1/2,c half-time of water exchange across a single cell - t_1/2 half-time of water exchange across multiple cells We thank Antony Matista for his expert assistance in the construction and modification of instruments. The work was supported by grant DCB8802033 from the National Science Foundation and grant 91-37100-6671 from USDA, and by the award of a Feodor Lynen-Fellowship from the Alexander von Humboldt-Foundation (Germany) to J.F.     

Resistance to water flow in xylem of Picea abies (L.) Karst. trees grown under contrasting light conditions

Summary The relative hydraulic conductivity (k) of xylem and resistance (R) to water flow through trunk, primary roots and branches in Picea abies trees growing under contrasting light conditions were investigated. The xylem permeability to water was measured by forcing 10 mM water solution of KC1 through excised wood specimens. From the values of k, the sapwood transverse area and the length of conducting segments, R of the whole trunk, branches and roots was calculated. The relative conductivity of xylem in open-grown trees exceeded that of shade-grown trees by 1.4–3.1 times, while k was closely correlated with the hydraulically effective radius (R _e) of the largest tracheids (R ² was 0.85–0.94 for open- and 0.51–0.79 for shade-grown trees). Because of both a low k and a smaller sapwood area in shade-grown trees the resistance to water movement through their trunk, roots and branches was many times higher. The distribution of R between single segments of the water-conducting pathway differed considerably in trees from different sites. At high water status the largest share of the total resistance in open- as well as shade-grown trees resides in the apical part of the trunk. The contribution of the branches to total xylem resistance is supposed to increase with developing water deficit.     

Root and stem xylem embolism,stomatal conductance,and leaf turgor in Acer grandidentatum populations along a soil moisture gradient

The objective of this study was to determine how adjustment in stomatal conductance (g ^s) and turgor loss point (_tlp) between riparian (wet) and neighboring slope (dry) populations of Acer grandidentum Nutt. was associated with the susceptibility of root versus stem xylem to embolism. Over two summers of study (1993–1994), the slope site had substantially lower xylem pressures (_px) and g ^s than the riparian site, particularly during the drought year of 1994. The _tlp was also lower at the slope (-2.9±0.1 MPa; all errors 95% confidence limits) than at riparian sites (-1.9±0.2 MPa); but it did not drop in response to the 1994 drought. Stem xylem did not differ in vulnerability to embolism between sites. Although slope-site stems lost a greater percentage of hydraulic conductance to embolism than riparian stems during the 1994 drought (46±11% versus 27±3%), they still maintained a safety margin of at least 1.7 MPa between midday _px and the critical pressure triggering catastrophic xylem embolism (_pxCT). Root xylem was more susceptible to embolism than stem xylem, and there were significant differences between sites: riparian roots were completely cavitated at -1.75 MPa, compared with -2.75 MPa for slope roots. Vulnerability to embolism was related to pore sizes in intervessel pit membranes and bore no simple relationship to vessel diameter. Safety margins from _pxCT averaged less than 0.6 MPa in roots at both the riparian and slope sites. Minimal safety margins at the slope site during the drought of 1994 may have led to the almost complete closure of stomata (g _s=9±2 versus 79±15 mmol m^-2 s^-1 at riparian site) and made any further osmotic adjustment of _tlp non-adaptive. Embolism in roots was at least partially reversed after fall rains. Although catastrophic embolism in roots may limit the minimum for gas exchange, partial (and reversible) root embolism may be adaptive in limiting water use as soil water is exhausted.     

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.